Personality and Neuroscience - Panksepp

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Neuroscience and Biobehavioral Reviews 35 (2011) 1946–1958

Contents lists available at ScienceDirect

Neuroscience and Biobehavioral Reviews


journal homepage: www.elsevier.com/locate/neubiorev

Review

The brain’s emotional foundations of human personality and the Affective


Neuroscience Personality Scales
Kenneth L. Davis a,∗ , Jaak Panksepp b
a
Department of Psychology, The University of North Carolina at Charlotte, Charlotte, NC 28223-0001, USA
b
Department of Veterinary and Comparative Anatomy, College of Veterinary Medicine, Washington State University, Pullman, WA 99164-6520, USA

a r t i c l e i n f o a b s t r a c t

Keywords: Six of the primary-process subcortical brain emotion systems – SEEKING, RAGE, FEAR, CARE, GRIEF and
Personality PLAY – are presented as foundational for human personality development, and hence as a potentially
Affective neuroscience novel template for personality assessment as in the Affective Neurosciences Personality Scales (ANPS),
Five Factor Model
described here. The ANPS was conceptualized as a potential clinical research tool, which would help
Subcortical brain emotion systems
experimentalists and clinicians situate subjects and clients in primary-process affective space. These
Affective neuroscience
Affective Neuroscience Personality Scales emotion systems are reviewed in the context of a multi-tiered framing of consciousness spanning from
primary affect, which encodes biological valences, to higher level tertiary (thought mediated) process-
ing. Supporting neuroscience research is presented along with comparisons to Cloninger’s Temperament
and Character Inventory and the Five Factor Model (FFM). Suggestions are made for grounding the inter-
nal structure of the FFM on the primal emotional systems recognized in affective neuroscience, which
may promote substantive dialog between human and animal research traditions. Personality is viewed
in the context of Darwinian “continuity” with the inherited subcortical brain emotion systems being
foundational, providing major forces for personality development in both humans and animals, and
providing an affective infrastructure for an expanded five factor descriptive model applying to normal
and clinical human populations as well as mammals generally. Links with ontogenetic and epigenetic
models of personality development are also presented. Potential novel clinical applications of the CARE
maternal-nurturance system and the PLAY system are also discussed.
© 2011 Elsevier Ltd. All rights reserved.

Contents

1. Introduction: Darwin, McDougal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1947


2. Subcortical emotion systems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1947
3. The Five Factor Model and the ANPS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1948
3.1. Cloninger’s biologically based personality theory . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1948
3.2. ANPS data . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1949
3.2.1. Conscientiousness . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1951
3.2.2. Non-human personalities . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1952
3.3. Expansion of the FFM . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1952
3.4. Examples of studies using ANPS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1952
4. Affective neuroscience trait versus theoretical expansion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1953
4.1. Primary affective consciousness . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1953
4.2. Could the fundamental nature of personality be affective? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1953
4.3. State vs. channel functions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1953
4.3.1. Evidence . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1954
4.3.2. Social rejection and the SADNESS system . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1954
4.3.3. Evidence summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1954
5. Clinical assessments from affective neuroscience trajectories . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1954

∗ Corresponding author. Tel.: +1 336 379 9828; fax: +1 336 379 9835.
E-mail address: [email protected] (K.L. Davis).

0149-7634/$ – see front matter © 2011 Elsevier Ltd. All rights reserved.
doi:10.1016/j.neubiorev.2011.04.004
K.L. Davis, J. Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 1946–1958 1947

6. Affective neuroscience and therapeutic effectiveness . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1955


7. Summary of affective neuroscience perspectives on the foundations of personality . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1955
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1956
Appendix A. Scoring the ANPS 2.4, copyright 2004 version . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1956
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1956

1. Introduction: Darwin, McDougal of the brain for generating specific emotional-behavioral episodes,
along with characteristic accompanying autonomic arousals. In
Charles Darwin provided the first modern scientific treatise on 1949, he received the Nobel Prize for his work on the central rep-
emotion in 1872 with “The Expression of the Emotions in Man resentations of the autonomic nervous system. As ruefully related
and Animals” (Darwin, 1872/1967). In this classic work, Darwin in his autobiographical materials, Hess resisted, with regrets, any
documented cross species commonalities that occurred for many explicit discussion of emotional experiences in the animals he stud-
primary emotions such as fear, anger, grief, and joy, which included ied, because he feared marginalization by behaviorists. He indicated
a discussion of playful tickling in human children and many other that he always believed these irritated animals were experienc-
descriptions of human emotional subtleties. Darwin’s keen obser- ing a state akin to human anger. He never inquired whether the
vations, which remain relevant to this day, allowed him to extend animals found such evoked states of anger to constitute aversive
his robust case for the “continuity of species” from body struc- experiences. Panksepp (1971) later demonstrated an homologous
tures to the domain of emotions. In line with his views, a recent anatomy in rats, and proceeded to show that, given the opportunity,
spectrographic analysis of the tickle-induced laughter response animals would escape the RAGE evoking stimulation.
on four species of young great apes plus humans yielded data Panksepp and some others continued these lines of research,
that effectively mapped the scientifically established evolutionary with explicit discussions of the accompanying affective experi-
branching of these closely related anthropoids (Ross et al., 2009). ences, based on whether the ESB (electrical stimulation of the brain)
Likewise, in synergy with Darwin’s vision, cross-species affective evoked ‘reward’ or ‘punishment’ effects in simple learning tasks.
neuroscientists have started to map out the continuity of primal Panksepp’s (1982, 1998, 2005) synthesis of such work provides
emotional systems in subcortical-limbic circuits of selected mam- documentation for the existence of seven “blue ribbon” subcorti-
malian brains (Panksepp, 1998) – from mice to men, so to speak. cal emotion systems in the brain, which he labeled SEEKING, RAGE,
The anatomical, neurochemical, and functional homologies of FEAR, LUST, maternal CARE, separation-distress PANIC/GRIEF, and
subcortical emotional networks strongly indicate underlying evo- physical PLAY. The capitalizations were premised on the need for
lutionary continuities in affective principles in all mammalian a specialized terminology for the primary-process anoetic experi-
brains (Panksepp and Biven, 2011). Since it is widely believed that ential processes of the brain, which together constitute forms of
individual’s emotional inclinations (temperaments) are founda- affective consciousness (Panksepp, 2007a)
tional for human personality development, the goal of the following This nomenclature allows the use of common emotional terms
work was to try to ground personality assessment on the affec- that convey the gist of the emotional meaning, while minimizing
tive circuits that all mammals share. But before we summarize mereological fallacies (part-whole confusions) that are endemic to
work with the Affective Neuroscience Personality Scales (ANPS), the use of emotional terms in describing animal experiences, thus
we briefly note historical antecedents of this way of thinking about providing some distance from dictionary definitions and every-
human personality. day understandings. We will never know how any other animal
In 1908 William McDougal published his classic work “Social or any other human actually feels. This is the classic dilemma
Psychology” in which he reviewed the influence of instincts on that subjectivity poses for modern neuroscience. All mammalian
human personality. He was the first personality theorist to be brains may be fundamentally subjective organs – they not only
guided by Darwin’s principle of the continuity of human and ani- generate behaviors, but “mental” processes such as basic feelings
mal mental evolution, and he used two principles for determining and perceptions (anoetic qualia), which control learning and mem-
whether an emotion should be considered a “primary emotion,” ory (noetic secondary-processes) as well as higher mental abilities
i.e. “simple instinctive impulse.” The first was whether the emotion (autonoetic tertiary processes) such as thoughts, plans and insights,
was “clearly displayed in the instinctive activities of the higher ani- aspects of mind that can only really be well-studied in humans
mals” (McDougal, 1908/1963, p. 42). Hence, he rejected that there (for a coverage of such nomenclature issues, see Vandekerckhove
was any religious instinct, and various other debatable higher- and Panksepp, 2009 and this issue). In contrast, feelings, because of
order affective-cognitive processes. His second principle was that their capacity to engender rewarding and punishing states in the
a primary emotion should be evident in human psychopathologies, brain, can be objectively studied in a substantive phenomenologi-
with emotional behaviors being manifested in extreme, exag- cal manner. Neuroscience advances will be substantially premised
gerated, and hence abnormal displays, which is consistent with on our ability to conceptualize brain functions both from the
certain current views of mental disorders (see below). He included external behavioral view as well as the internal psychological
only seven behaviorally well-defined emotional instincts, which he perspective.
labeled flight, repulsion, curiosity, pugnacity, self-abasement, self- We sustained the capitalization convention in our attempt (i.e.,
assertion, and the parental instinct, a list based on comparative the ANPS) to translate anoetic (without knowledge) emotional
observations that is arguably still relevant to personality theory a experiences to noetic (knowledge linked) and autonoetic (thought
century later, although those strands of early ethological thought linked), language-based evaluations of the potential influences of
were never fully developed. primal affects in the personality structures of human minds. How-
ever, rather than RAGE and PANIC/GRIEF, we decided to use the
2. Subcortical emotion systems terms ANGER and SADNESS in this paper and in our construction of
the Affective Neuroscience Personality Scales (ANPS), since those
Walter Hess’s seminal work on evoking emotions with localized are labels that are generally more understandable for most indi-
electrical stimulation of the brain (research summarized in Hess, viduals. Still, the theoretical implication is that these systems are
1957) demonstrated the importance of discrete subcortical areas critically important for the various feelings highlighted by these sci-
1948 K.L. Davis, J. Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 1946–1958

entific terms – e.g., the ANGER system energizes states commonly be informed by these subcortical affects. Specifically, studies of
labeled as ‘irritability’ and ‘anger’. primary-process emotional aspects of personality may provide a
Neuroanatomical, neurochemical, pharmacological, and physi- heuristic intersection for the Darwinian “continuity of species”,
ological brain research have helped to more precisely define each of with the cross-species emotion systems of mammalian brains, and
these seven emotion systems (see Panksepp, 1998, 2005; Panksepp the need for neurobiologically meaningful psychological assess-
and Biven, 2011 for detailed reviews). We assumed these brain ment of human temperaments/personalities.
systems could serve as an empirical roadmap for advances in the But this is not the first attempt to bridge basic neuroscience
study of personality and serve as a template for emotional person- and human personality evaluation. The two individuals who have
ality assessment, as in the ANPS (Davis et al., 2003), which was attempted to construct neurobiologically based personality assess-
conceptualized largely as a research tool aspiring to, and perhaps ment instruments in the past are Cloninger (1986) who based
capable of, situating adult human temperaments within the com- his thinking on functional animal work on some of the first well
plexities of primary-process affective space shared by all mammals. characterized neurochemical systems of all mammalian brains –
If so, that could also open up a common terminology for study per- dopamine, norepinephrine, and serotonin (see below). In con-
sonality across species. It is a major supposition of this paper that trast, Carver and White (1994) developed the Behavioral Activation
these basic emotions are fundamental powers of the human Brain- and Behavioral Inhibition Scales on the basis of Jeffrey Gray’s
Mind, homologous with feelings experienced by other mammals, (1981) early brain research recognizing the existence of behavioral
and hence potentially of critical importance for understanding the approach/arousal and withdrawal/inhibition (aka, reward and pun-
foundations of personality (we intentionally conflate ‘brain’ and ishment) systems in the brain. The present work is an extension of
‘mind’, and double capitalized BrainMind to reflect our monistic this strategy, as our understanding of brain affective systems has
ontology). become more resolved. Because of space constraints, we will only
There is strong evidence for these seven well-defined emo- contrast our work on the ANPS, designed to estimate six affects rele-
tion systems in all mammalian brains. Several of these emotions vant for defining temperamental/personality dimensions, with the
have evolutionally deep reptilian roots (SEEKING, ANGER, FEAR and popular Cloninger assessment tool following a brief introduction of
LUST), but three (CARING, SADNESS, and PLAY) are more uniquely the five factor personality model.
mammalian adaptations, but surely with antecedents in “lower”
vertebrates, perhaps invertebrates also (e.g., see Huber et al., 2011). 3. The Five Factor Model and the ANPS
Each of these seven emotions can be evoked by stimulating distinct
but partly overlapping subcortical regions of mammalian brains. At the very outset of this work, the ANPS was hypothesized
However, they typically work interactively in concert to increase to provide affective underpinnings to the widely used Five Fac-
the adaptiveness of individual feelings, perceptions, thoughts, and tor Model (FFM). The FFM has provided a personality assessment
behaviors. (see Panksepp, 1998, 2005; Panksepp and Biven, 2011 standard, which while not universally accepted (Block, 1995) has
for fuller descriptions). offered a robust personality model in general population and clin-
There are several principles basic to understanding these seven ical research (Costa and Widiger, 2002). The FFM has origins in
brain systems: (1) These emotion systems are subcortical networks adjective descriptors of personality (Allport and Odbert, 1936)
and lower brain regions have evolutionary primacy in generating and the early history of factor analysis (Cattell, 1947). The five
these basic emotions and their affects, while learning and higher personality dimensions are usually labeled Extraversion or Sur-
brain functions can be deemed to be secondary and tertiary pro- gency, Agreeableness, Conscientiousness, Emotional Stability, and
cesses. (2) To the best of our knowledge, these emotion systems, Intellect or Openness to Experience (Hofstee et al., 1992). These
situated in ancient brain regions, are largely homologous in all five dimensions have been consistently derived from a variety of
mammals. (3) These basic emotions also have similar chemistries in descriptive data using factor analysis (Digman, 1990) and have
all mammals. (4) These brain systems generate instinctual behav- been replicated in several languages (Saucier and Goldberg, 2006).
ioral responses that are closely linked to the raw, primal affects However, the first four dimensions have been more robust with
that accompany those responses. (5) The integrity of these seven Openness to Experience typically accounting for the least variance
systems is demonstrated by the ability to elicit coherent specific of the five factors (Goldberg, 1990). A comprehensive presenta-
emotional responses and/or the associated affects with localized tion of the five factor “descriptive” model was provided by Hofstee
brain stimulation – as evaluated by the capacity of the subcortical et al. (1992) and consisted of an analysis of over 500 adjectives
arousals to mediate ‘reward’ and ‘punishment’ functions that con- placed on five orthogonal dimensions as well as numerous off-axis,
trol learning. (6) Lastly, these systems remain relatively unscathed two-factor “blends,” such as “angry” emerging as a “blend” of low
in animals whose neocortices were surgically removed in early Emotional Stability and low Agreeableness.
development (see Panksepp, 1998).
In summary, emotional responses from each of the primary 3.1. Cloninger’s biologically based personality theory
process emotions can be activated by localized subcortical ESB or
chemical brain stimulation. That decortication of young animals Cloninger was the first to attempt an integration of personality
generally leaves the expression of these emotions intact (Deyo and brain systems with his biogenic amine-based “biosocial” theory
et al., 1990; Panksepp et al., 1994), further reinforces the sub- of personality (Cloninger, 1987) and the Tridimensional Personal-
cortical nature of these emotion systems. Also, each instinctual ity Questionnaire (TPQ), which included three scales: (1) Novelty
emotional system engenders affective valence since animals termi- Seeking based on a postulated dopaminergically based behavioral
nate the stimulation of ANGER, FEAR, and SADNESS, while actively activation system that promotes interest in new experiences, (2)
working to obtain SEEKING, LUST, CARE, and PLAY system arousals Harm Avoidance, postulated to primarily reflect brain serotonin
(Panksepp, 2005). However, there are reasons to believe the SEEK- activity, the brain’s presumed punishment system (which is highly
ING system is foundational for aspects of all of the other primal debatable, see Panksepp, 1986, 1998), and the resulting sensitiv-
emotions (e.g., during FEAR intensification, the shift from freezing ity to threatening situations, and (3) Reward Dependence featuring
to flight may be due to recruitment of SEEKING related dopamin- behavioral maintenance and sensitivity to reinforcement contin-
ergic psychomotor drive). gencies with norepinephrine as the major neuromodulator. Each of
It is our premise that these emotions form an important these “temperament” traits was seen as highly heritable and stable
foundation of personality. As such, personality assessment can across the developmental stages of life.
K.L. Davis, J. Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 1946–1958 1949

Our affective neuroscience perspective would align the Novelty ally liking to explore. CARING centered on nurturing tendencies
Seeking dopaminergic brain incentive dimension with the SEEK- including liking to care for others, being drawn to young chil-
ING system. The SEEKING system not only promotes exploration, dren and pets, and feeling softhearted towards animals and people
investigation and foraging but probably ‘energizes’ all basic emo- in need. The FEAR scale incorporated experiencing anxiety, wor-
tional systems with forms of appetitive and anticipatory arousal rying, difficulty making decisions, ruminating, feeling tense, and
(e.g., as already noted, seeking of safety when FEAR is aroused). losing sleep. ANGER included feeling hotheaded, being easily irri-
We suspect that serotonin and norepinephrine may function too tated and frustrated, and expressing anger verbally or physically.
broadly to be isolated as specific contributors to very specific SADNESS was conceptualized as feeling social separation distress,
types of affects the way Cloninger proposed, even though they feeling lonely, and thinking about loved ones and past relation-
figure heavily in the arousal dimensions of all personality control ships including crying. The Spirituality scale focused on feelings of
systems. Norepinephrine promotes attention by amplifying signal- connectedness with all of life and oneness with creation.
to-noise processing in most sensory-perceptual channels and also The revised ANPS 2.4 items are listed in Fig. 1 (for details, see
promotes emotionality in general. In contrast, serotonin dampens Appendix A). These scales are provided for free use in all scien-
and restricts neural processing in the brain, resulting generally in tific endeavors and non-commercial clinical use (e.g., individual
reduced emotionality (Panksepp, 1986). psychotherapy practitioners) without need to obtain permission
Cloninger subsequently expanded his assessment instrument of the copyright owners (Davis and Panksepp). Those wishing to
adding an additional temperament dimension – Persistence – and use it for commercial use are requested to contact the first author.
three controversial “character” traits – Self-Directedness, Coop- Again, a detailed description of the scales and scoring instructions
erativeness, and Self-Transcendence – hypothesized to have low are provided in the appendix at the end of this paper.
heritabilities and be more sensitive to individual development (we Based on our first study using these scales, reliabilities for the
would conceptualize these as higher-order, tertiary-process type ANPS scales, computed as Cronbach’s alpha, were reported rang-
BrainMind functions). The revised instrument was called the Tem- ing from .65 to .86 with the PLAY and SEEK scales below .70 and
perament and Character Inventory (TCI; Cloninger, 1999). Although the FEAR, ANGER, and Spirituality scales above .80 (Davis et al.,
Cloninger’s earlier TPQ factors have been related to many genetic 2003). Multiple data sets using the revised ANPS 2.4, have shown
variables, many of Cloninger’s later theoretical claims have not all reliabilities now over .70 (unpublished data).
been supported. In other words, heritabilities did not differenti- Contrasting self-rating data for ANPS and FFM scales, as seen
ate the temperament and character scales (Gillespie et al., 2003; in Table 1, revealed that each of the ANPS scales except Spirituality
Cloninger, 2004). Other research found that the Self-Directedness correlated strongly with at least one of the FFM scales as follows: (1)
character scale also responded to serotonin treatments (Peirson SEEK with Openness to Experience (r = .47), (2) PLAY with Extraver-
et al., 1999; Wai and Bond, 2001). Factor analysis placed the tem- sion (r = .46), (3) CARE with Agreeableness (r = .50), (4) FEAR with
perament Harm Avoidance and the character Self-Directedness on Emotional Stability (r = −.75), (5) SADNESS with Emotional Stabil-
the same factor, and hence did not support a distinction between ity (r = −.68), and (6) ANGER with Emotional Stability (r = −.65)
“temperament” and “character” dimensions, and accounted for as well as with Agreeableness (r = −.48) (Davis et al., 2003). The
the revised TCI scales with five factors (Farmer and Goldberg, data supported strong relationships between primary emotions
2008a,b). Others also reported considerable convergence between and the most widely accepted model of human personality, which
Cloninger’s revised TCI (Cloninger, 1999) and the FFM (Ball et al., was consistent with the hypothesis that these six brain emotion
1999; De Fruyt et al., 2000). systems form a foundation for the adult five-factor personality
model.
3.2. ANPS data When factor analyzing ANPS scales with FFM scales, we reported
a five factor solution with the FFM Conscientiousness scale on a
The Affective Neuroscience Personality Scales (ANPS) were factor by itself. Since there were only four eigenvalues > 1.0, we
introduced to explore the potential primary-process brain emo- removed the Conscientiousness scale and published a four factor
tional system foundations of personality. The scales were rationally solution with (1) FEAR SADNESS, ANGER, and Emotional Stability,
defined to potentially provide biological underpinnings for the (2) CARE and Agreeableness, (3) PLAY and Extraversion, and (4)
descriptive FFM of personality (Davis et al., 2003). Our view is that SEEK and Openness to Experience loading on four separate scales
these personality dimensions escape the problem of definitional with .68 being the lowest primary factor loading. The only scale
circularity since comparative research has demonstrated how each having a loading on a second factor greater than .30 was ANGER,
primary dimension can be manipulated in humans as indepen- which also loaded −.53 on the Agreeableness factor. In summary,
dent variables including experientially (Eisenberger et al., 2003; the ANPS offers personality primes that align with the Big Five
Panksepp, 1985) or pharmacologically as well as through brain model but target the six specific primary emotion brain systems
damage research (e.g., Bechara et al., 1999). that have been well documented with comparative brain research.
Each ANPS dimension corresponded to one of the “blue ribbon” A factor analysis of just the six primary ANPS scales yielded
primal emotions identified by cross-species ESB-based affective two eigenvalues greater than 1.0 with FEAR, SADNESS, and ANGER
neuroscience research. We excluded LUST, which seems less rel- on the first component and PLAY, CARE, and SEEK on the second
evant to current conceptualizations of human personality and we with all primary factor loadings greater than .55 and all secondary
also suspected that it may potentially be an affective factor that loadings less than .20. This clustering, corresponds nicely to other
people would not wish to be frank about, and thus may contami- scales which simply try to parse human personality into positive
nate frankness on the other scales. In addition, a Spirituality scale and negative affective dimensions (Watson et al., 1988) and gen-
was included to acknowledge the importance of this factor in drug erally replicates Gray’s (1970) Behavioral Inhibition and Approach
addiction (Panksepp et al., 2004) and many drug addiction recovery Systems. We feel this is a striking demonstration of the multifac-
programs (e.g., 12-step programs) and what might be considered torial nature of any general positive and negative valence type of
the highest human emotion. measurement tool. In this vein, it is also important to note Goldberg
The PLAY scale focused on playing social games with physical and colleague’s argument (Ashton et al., 2009, p. 80) that higher
contact plus laughter, humor, and generally having fun. SEEKING order factors such as Positive Affect, Negative Affect, and overall
was defined as anticipating new positive experiences including Emotional Stability “may represent artifacts rather than substan-
being curious, liking to strive for solutions to problems, and gener- tive dimensions of personality.”
1950 K.L. Davis, J. Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 1946–1958

Pahlavan et al. (2008) confirmed the psychometric properties (2008c) obtained significant evidence of heritability for four of
of the ANPS in a French population including scale reliabili- the ANPS scales: CARE, SEEK, FEAR, and Spirituality. Through per-
ties, factor structure, and correlations with the FFM. Berthoz sonal correspondence, we know the scale has been translated into
and colleagues (personal communication) replicated the Pahlavan a dozen languages, and has already proved its utility in eval-
et al. (2008) findings on a large sample (n = 830) and also uating certain psychiatric problems (e.g., Savitz et al., 2008a,b;
performed an item-level confirmatory factor analysis obtain- see below). For a while Martin Reuter (University of Bonn) had
ing an acceptable fit modeling 6 factors (standardized root all the original scales readily available on the web, and also for
mean square residual = .06, root mean square error of approxi- data collection purposes (www.anps.de), but this site has been
mation = .039, and the comparative fit index = .78). Savitz et al. closed.

Affective Neuroscience Personality Scale 2.4 Name:


Age: Sex: Please mark bubbles like this Disagree Agree
Str Disagree Str Agree
1. Almost any little problem or puzzle stimulates my interest.
2. People who know me well would say I am an anxious person.
3. I often feel a strong need to take care of others.
4. When I am frustrated, I usually get angry.
5. I am a person who is easily amused and laughs a lot.
6. I often feel sad.
7. Feeling a oneness with all of creation helps give more meaning to my life.
8. I like to be the one in a group making the decisions.
9. I do not get much pleasure out of looking forward to special events.
10. I am not frequently jittery and nervous.
11. I think it is ridiculous the way some people carry on around baby animals.
12. I never stay irritated at anyone for very long.
13. My friends would probably describe me as being too serious.
14. I seem to be affected very little by personal rejection.
15. Feeling like a part of creation is not an important source of meaning for my life.
16. I will gossip a little at times.
17. I really enjoy looking forward to new experiences.
18. I often think of what I should have done after the opportunity has passed.
19. I like taking care of children.
20. My friends would probably describe me as hotheaded.
21. I am known as one who keeps work fun.
22. I often have the feeling that I am going to cry.
23. I am often spiritually touched by the beauty of creation.
24. I usually avoid activities in which I would be the center of attention.
25. I am usually not highly curious.
26. I would not describe myself as a worrier.
27. Caring for a sick person would be a burden for me.
28. I cannot remember a time when I became so angry that I wanted to break something.
29. I generally do not like vigorous games which require physical contact.
30. I rarely become sad.
31. I rarely rely on spiritual inspiration to help me meet important challenges.
32. I always tell the truth.
33. Seeking an answer is as enjoyable as finding the solution.
34. I often cannot fall right to sleep because something is troubling me.
35. I love being around baby animals.
36. When I get angry, I often feel like swearing.
37. I like to joke around with other people.
38. I often feel lonely.
39. For me, experiencing a connection to all of life is an important source of inspiration.
40. When I play games, it is important for me to win.
41. I usually feel little eagerness or anticipation.
42. I have very few fears in my life.
43. I do not especially like being around children.
44. When I am frustrated, I rarely become angry.
45. I dislike humor that gets really silly.
46. I never become homesick.
47. For me, spirituality is not a primary source of inner peace and harmony.
48. Sometimes I feel like swearing.
49. I enjoy anticipating and working towards a goal almost as much as achieving it.
50. I sometimes cannot stop worrying about my problems.
51. I feel softhearted towards stray animals.
52. When someone makes me angry, I tend to remain fired up for a long time.
53. People who know me would say I am a very fun-loving person.
54. I often think about people I have loved who are no longer with me.
55. Contemplating spiritual issues often fills me with a sense of intense awe and possibility.
56. If my peers have outperformed me, I would still be happy, if I have nearly met my goals.
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Copyright © 2004, Kenneth L. Davis, Ph.D., Jaak Panksepp, Ph.D., Pegasus International, Inc. All rights reserved.

Fig. 1. (a) Affective Neuroscience Personality Scale 2.4 items and response scale. The ANPS items are arranged in fourteen blocks using the following item sequence: SEEK,
FEAR, CARE, ANGER, PLAY, SADNESS as described in Appendix A. (b) ANPS continued: see description in above figure legend and Appendix A.
K.L. Davis, J. Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 1946–1958 1951

Please mark bubbles like this and not like this or


Disagree Agree
Str Disagree Str Agree
57. I am usually not interested in solving problems and puzzles just for the sake of solving them.
58. My friends would say that it takes a lot to frighten me.
59. I would generally consider pets in my home to be more trouble than they are worth.
60. People who know me well would say I almost never become angry.
61. I do not particularly enjoy kidding around and exchanging "wisecracks."
62. It does not particularly sadden me when friends or family members are disapproving of me.
63. My sense of significance and purpose in life does not come from my spiritual beliefs.
64. I have never "played sick" to get out of something.
65. My curiosity often drives me to do things.
66. I often worry about the future.
67. I feel sorry for the homeless.
68. I tend to get irritated if someone tries to stop me from doing what I want to do.
69. I am very playful.
70. I tend to think about losing loved ones often.
71. Feeling a connection with the rest of humanity motivates me to make more ethical choices.
72. When I play games, I do not mind losing.
73. I rarely feel the need just to get out and explore things.
74. There are very few things that make me anxious.
75. I do not like to feel "needed" by other people.
76. I rarely get angry enough to want to hit someone.
77. I do not tend to see the humor in things many people consider funny.
78. I rarely have the feeling that I am close to tears.
79. The goals I set for myself are not influenced by my spirituality.
80. There have been times in my life when I was afraid of the dark.
81. Whenever I am in a new place, I like to explore the area and get a better feel for my surroundings.
82. I often worry about whether I am making the correct decision.
83. I am the kind of person that likes to touch and hug people.
84. When things do not work out the way I want, I sometimes feel like kicking or hitting something.
85. I like all kinds of games including those with physical contact.
86. I frequently feel downhearted when I cannot be with my friends or loved ones.
87. Spiritual inspiration helps me transcend my limitations.
88. I am not satisfied unless I can stay ahead of my peers.
89. I am not the kind of person that likes probing and investigating problems.
90. I rarely worry about my future.
91. I do not especially want people to be emotionally close to me.
92. I hardly ever become so angry at someone that I feel like yelling at them.
93. I do not frequently ask other people to join me for fun activities.
94. I rarely think about people or relationships I have lost.
95. My choices are not guided by a sense of connectedness with all of life.
96. I have never intentionally told a lie.
97. I often feel like I could accomplish almost anything.
98. I often feel nervous and have difficulty relaxing.
99. I am a person who strongly feels the pain of other people.
100. Sometimes little quirky things people do really annoy me.
101. I see life as being full of opportunities to have fun.
102. I am a person who strongly feels the pain from my personal losses.
103. When working on a project, I like having authority over others.
104. Being embarrassed or looking stupid are among my worst fears.
105. I am not an extremely inquisitive person.
106. I almost never lose sleep worrying about things.
107. I am not particularly affectionate.
108. When people irritate me, I rarely feel the urge to say nasty things to them.
109. Playing games with other people is not especially enjoyable for me.
110. It would not bother me to spend the holidays away from family and friends.
111. Striving to be better than my peers is not important for me.
112. Fear of embarrassment often causes me to avoid doing things or speaking to others.
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Copyright © 2004, Kenneth L. Davis, Ph.D., Jaak Panksepp, Ph.D., Pegasus International, Inc. All rights reserved.

Fig. 1. (Continued ).

3.2.1. Conscientiousness in chimpanzees (Gosling and John, 1999), which suggests that
Conscientiousness was the only FFM dimension that did Conscientiousness is a more cerebral dimension emerging late
not correlate strongly with the ANPS. Lower correlations with in mammalian evolution. Congdon and Canli’s (2008) view of
ANGER (r = −.30), FEAR (r = −.24), and SADNESS (r = −.30) sug- impulsiveness as a “top down” lack of behavioral inhibition
gested that while Conscientiousness may be associated with the involving the inferior frontal cortex and subthalamic nucleus fits
regulation of negative affects, it seems unlikely that a single well with King’s (2007) description of chimpanzee Conscientious-
primary brain affect serves as the foundation for Conscientious- ness – characterized by predictable, not impulsive or erratic,
ness. From animal FFM studies, Conscientiousness only appeared behaviors.
1952 K.L. Davis, J. Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 1946–1958

Table 1
ANPS Scale correlations with Five Factor Model scales.

Extraversion Agreeableness Conscientiousness Emotional stability Openness to experience

PLAY .46* .29* .00 .12 .13


SEEK .13 −.01 −.01 .01 .47*
CARE .25 .50* .12 −.07 .06
FEAR −.19 −.17 −.24 −.75* −.05
ANGER −.04 −.48* −.30* −.65* −.08
SADNESS −.21 −.13 −.30* −.68* −.00
Spirituality .15 .26* .14 .09 .17

Student sample: n = 170 (50 males, 121 females).The Five Factor Model was measured using Goldberg (1992) adjectives, selecting 14 markers for each Big Five dimension
(data adapted from Davis et al., 2003).
*
p < .001, two-tailed.

These results provide encouraging evidence that the FFM reflects a tertiary cognitive-type reconfiguration of the underlying
reflects primary subcortical affects. Although ANPS items attempt primary brain emotion systems.
to address primary affects directly, since all self-report assessments
must include cognitive reflection, we interpret the ANPS scales as 3.4. Examples of studies using ANPS
tertiary (thought-mediated) approximations of the influence of the
various primary emotional systems in people’s lives. However, it Several of the ANPS scales have been further validated by
is our working hypothesis that the subcortical primary-processes independent research. As an example of ANPS research, study-
neural systems, where the foundations of emotions reside, can gen- ing families with a member having bipolar disorder (BPD), Savitz
erate individual differences in normal personality as well as the et al. (2008a) found that the ANPS SADNESS scale was significantly
affective imbalances characterizing mental disorders. higher in BPD I diagnosed individuals than two control groups
made up of unaffected family members or family members with a
DSM-IV diagnosis other than depression such as alcoholism or gen-
3.2.2. Non-human personalities
eralized anxiety disorder. Consistent with the quantitative genetic
If subcortical primary-processes are foundational for the FFM,
model of bipolar spectrum illness (Evans et al., 2005), the ANPS
one should be able to identify these same FFM dimensions in
SADNESS and FEAR scores trended highest for individuals diag-
non-human mammals. In a review of 19 studies of personality
nosed as BPD I or II, lower for individuals with recurrent major
in non-human animals, Gosling and John (1999) found consistent
depression or a single depressive episode, and lowest for the two
cross-species evidence for the FFM in mammals ranging from rats to
control groups. However, after controlling for self-rated depres-
chimpanzees. The most consistent FFM temperament dimensions
sion (Beck Depression Inventory (Beck and Steer, 1993)) or mania
observed were Extraversion and Emotional Stability. Agreeable-
(Altman Self-Rating Mania Scale (Altman et al., 1997)), only the
ness was the next most common. Consistent with human data
SADNESS scale continued to show significant differences across
(Goldberg, 1990), the evidence for Openness to Experience was
groups.
weaker than for the previously named three dimensions. As noted
Savitz et al. (2008b) also studied the distribution of hypo-
previously, Conscientiousness was only observed in chimpanzees,
manic, cyclothymic, and hostile traits in BPD probands and their
human’s closest evolutionary relative. These findings support the
families. After controlling for age, gender, depression, and mania,
Darwinian concept of continuity of species that “the difference in
BPD II patients had significantly higher ANPS ANGER scores than
the mind between man and the higher animals, great as it is, cer-
individuals with only a single previous depressive episode, and
tainly is one of degree and not of kind.” (Darwin, 1872/1967, p.
higher ANGER scores than unaffected family members. Savitz
104)
et al. (2008b) also reported that Beck Depression Inventory
scores were significantly correlated with higher ANGER scores and
3.3. Expansion of the FFM lower SEEK and PLAY scores while the Altman Self-Rating Mania
Scale significantly correlated with higher ANGER, SEEK, and PLAY
These ANPS results also revealed FFM limitations such as FFM scores.
Emotional Stability associating with all three of the ANPS nega- The construct validity of the ANPS ANGER scale was also sup-
tive affects, FEAR, SADNESS, and ANGER, which makes Emotional ported by Reuter et al. (2009) investigating links between ANGER
Stability equivalent to Negative Affect. A recent meta-analysis of and the rs907094 (aC → T single nucleotide polymorphism of the
past brain imaging studies also provided strong evidence that dis- DARPP-32 gene). Dopamine- and cAMP-regulated phosphoprotein,
tinct anatomical systems in human brains can be related to basic 32 kDa (DARPP-32) is a key regulatory molecule in the dopaminer-
affective processes including support for sadness, fear, and anger gic signaling pathway. The C-allele is more common in Sub-Saharan
(Vytal and Hamann, 2010). At the tertiary process level, humans Africa but rather infrequent in European populations. The genetic
may not always differentiate well between the distressful feelings analysis of German subjects without psychopathology showed that
associated with FEAR, SADNESS, and ANGER, which could lead to carriers of the T-allele had significantly higher ANPS ANGER scores,
statistical lumping, but that does not mean it is desirable to com- with CC genotypes having significantly lower ANGER scores. MRI
bine three of our most powerful emotional drivers into a single data on a subsample of subjects revealed that ANGER scores cor-
dimension. related negatively with gray matter volume in the left amygdala.
ANGER also correlated with Agreeableness. Despite the fact that Reuter’s group has also recently observed that the homozygous
the ANPS ANGER and CARE did not correlate (r = −.036 in Davis long-variant of the serotonin transporter polymorphism and the
et al., 2003), the two scales correlated in opposite directions with TT variant of the single nucleotide polymorphism rs2268498 on
the FFM Agreeableness scale. Human minds seem to generate a the oxytocin receptor gene showed significantly lower scores on
descriptive “love/hate” Agreeableness construct with caring, nur- the personality dimensions FEAR and SADNESS of the ANPS, as well
turing feelings on the positive pole and hostile, demanding feelings as on the overall super-factor of Negative Emotionality, than carri-
on the negative pole. Our view is that while the FFM is an impor- ers of the other serotonin transporter and oxytocin gene variants
tant theory-free, empirical step forward in parsing personality, it (Montag et al., 2011).
K.L. Davis, J. Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 1946–1958 1953

Reuter et al. (2005) also demonstrated a link between the ANPS identical siblings (whether adopted or not) would converge over
SEEK scale and creativity. Using figural, verbal, and numeric cre- time.
ativity tests, subjects with high SEEK scores had higher numeric
creativity scores and were significantly superior on figural and ver- 4.1. Primary affective consciousness
bal creativity. SEEK scores also explained more than 15% of the
variance of total creativity. Using intelligence tests as a covari- Another affective neuroscience concept related to personality is
ate indicated that the relationship of SEEK to creativity was not the multi-tiered framing of consciousness. At the primary emotion
moderated by intelligence. level of consciousness, it is clear that young animals and humans
without a neocortex are still conscious creatures with primi-
4. Affective neuroscience trait versus theoretical expansion tive affective and perceptual capacities (Merker, 2007; Shewmon
et al., 1999) including the capacity to play (Panksepp et al., 1994).
In addition to suggesting a reprioritized selection of person- This primary-process affective consciousness may include various
ality traits in the FFM, affective neuroscience (Panksepp, 1998) sensory/perceptual feelings but especially includes ancient emo-
suggests an enhanced neuroscientifically premised personality tional/motivational experiences all mammals share that provide
research model. By providing a putative physiological basis for the basis of a “primordial evaluative system” (Kahneman, 2003,
FFM traits, affective neuroscience helps five-factor theory escape p. 701). This primary level makes itself felt throughout our lives
circular reasoning by suggesting brain mechanisms that can be but is soon augmented by secondary consciousness reflecting basic
manipulated to influence the expression of personality. However, conditioning-learning. In humans, and perhaps some other corti-
even more important may be helping to advance personality theory cally well-endowed animals, thoughts about how external events
from trait and situationist approaches to interactionist and ontoge- relate to internal events yield even higher tertiary forms of con-
netic personality models (see Ballantyne, 1995 for a more detailed sciousness consisting of thoughts about thoughts, awareness of
discussion of personality theory levels). awareness, and the linguistic/symbolic transformation of experi-
One ontogenetic model is Scarr and McCartney’s (1983) theory ence at which humans excel (for overview, see Vandekerckhove
of genotype-environment effects hypothesizing that individuals and Panksepp, 2009 and in this issue). However, it is the primary-
make their own environments. The theory predicts that earlier process level of affective consciousness that encodes biological
in life, individuals experience more passive environmental effects values through a diversity of raw positively and negatively valenced
through the environment provided by their parents. However, emotional experiences, from pain to joy, that may be foundational
there is also an evoked effect, since infant individual differences for personality as well as disorders of personality, which reflect
can influence the kind of care infants receive from their parents. imbalances in these intrinsic value systems of the BrainMind.
While the passive effect is hypothesized to decrease with onto-
genetic development, the evoked effect continues throughout an 4.2. Could the fundamental nature of personality be affective?
individual’s life. However, there is also an active effect in which
the individual selects from different environments. This active A key question from a purely cognitive perspective is how
selection of environments becomes relatively more important as non-humans with their more limited cerebral cortical capacities
individuals mature and are more able to choose the circumstances compared to humans can have distinct personalities at all, let alone
and experiences they find attractive, which accounts for adopted personalities defined by the well accepted FFM dimensions (see
siblings not resembling each other’s personalities by late adoles- Gosling and John, 1999). Our theory is related to the idea that
cence (Scarr et al., 1981; Scarr and McCartney, 1983). the affective foundations of personality lie in MacLean’s (1990)
This genotype-environment interaction model can be tested sub-neocortical “limbic” and “reptilian” areas of the central ner-
with an affective neuroscience approach to personality. At the vous system. It is our position that in these evolutionally older
secondary process level characterized by simple conditioning and parts of the brain, much more ancient than “neomammalian”
learning, conditioned place preferences or aversions provide a cortex, one finds the most important evolutionary “roots” of per-
mechanism for defining the attractiveness of environments. Plea- sonality. Furthermore, this homologous foundation consists of
surable experiences result in finding similar situations attractive instinctual emotion action systems shared by all mammals. These
in the future with traumatic or painful experiences having the “ancient tools for living” engender a within-brain type of emo-
opposite effect. One example prediction would be that individu- tional affective valence, as well as behavioral reactive guidance,
als who were more sensitive to social pain (for a full discussion of and general physical adaptation to survival challenges faced by
this construct, see Panksepp, 2011 as well as other contributions our ancestors for millions of years. Thus, they were built into
to understanding psychic pain in that book edited by MacDonald the BrainMind as ancestral “memories”. These “primary” systems
and Jensen-Campbell) would find such distressing situations more are undoubtedly elaborated during human development by “sec-
aversive – perhaps at lower thresholds – and exhibit more avoidant ondary” conditioning and “tertiary” thoughts and self-reflections,
behavior after those experiences than individuals who were less but the evolutionary origin and foundational power of these dis-
sensitive to social pain. crete, persistent systems is pre-human.
Early experience epigenetic models have already been tested
in rats. Focusing on the PLAY system, extra provisioning of rough- 4.3. State vs. channel functions
and-tumble play decreased ADHD-type impulsiveness (Panksepp
et al., 2003). Centering on the CARE system, superior maternal The influence of these basic emotion systems is pervasive, affect-
nurturing positively contributed to the stress tolerance of off- ing not only our actions and reactions but also our perceptions,
spring (Francis et al., 1999). Since personality traits can also be thoughts, and memories. The mechanism mediating these old brain
assessed in animal subjects (Gosling and John, 1999; Gosling, 2008), influences may lie in the distinction between “channel” and “state”
such longitudinal effects of primary-process experiences can be brain functions (Mesulam, 2000). For “state” modulation, each cor-
tested with animal models that incorporate variables and controls tical area receives inputs arising from limbic neurons, which can
that would not be possible with human subjects. In shorter- modulate activity in the entire cerebral cortex. These influences
lived species, one could also test Scarr and McCartney’s (1983) determine the “state” of information processing rather than the
theory that the personalities of fraternal and adoptive siblings content being transmitted along the point-to-point “channels.”
would become less similar as they mature whereas genetically These modulatory “corticopetal” projections to the cortex are not
1954 K.L. Davis, J. Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 1946–1958

balanced by corresponding “corticofugal” connections from the be elicited by social rejection by peers and general social isolation.
cerebral cortex to the subcortex. This asymmetry allows the limbic Eisenberger et al. (2003) did an fMRI analysis of brain changes as she
system to “rapidly shift information processing states throughout manipulated “painful” social exclusion experienced by participants
the cerebral cortex . . . [and] alter the tone, coloring, and interpre- supposedly playing a virtual ball-tossing game with other partic-
tation of experience rather than its content” (Mesulam, 2000, pp. ipants. During an initial control scan, subjects just watched other
78–79). participants play the virtual ball-tossing game, ostensibly because
“technical difficulties” prevented them from participating. In the
4.3.1. Evidence second scan, subjects were linked in with “virtual” participants sup-
Evidence supporting such “state” modulation comes from many posedly in other scanners and “included” in the virtual ball-tossing
sources, and we just touch on a few: Wu et al. (1991) report that game. To create a sense of social exclusion, during the final scan
patients with generalized anxiety disorder exhibited stronger tem- subjects were socially excluded when the other players (actually a
poral and frontal cortex PET responses to a passive viewing task computer program) stopped tossing the ball to the subject. Subjects
compared to normal controls. Also, individuals screened for lack of showed significantly more dACC activation during the social exclu-
a mental health treatment history but with higher Beck Depression sion condition compared to inclusion. Measures of self-reported
Inventory (BDI) scores exhibited heightened resting fMRI amygdala distress also correlated positively with dACC activity supporting
activity to neutral visual stimuli (Way et al., 2010). Furthermore, the relationship between dACC activity and felt emotional distress
when compared to subjects with lower BDI scores, these higher during the social exclusion condition. In addition, right ventral pre-
scoring BDI subjects also showed greater amygdala reactivity when frontal cortex (RVPFC) was also more active during social exclusion
viewing fearful or angry faces. In other words, differences in sub- compared to inclusion conditions but negatively associated with
cortical “states” biased perceptions and generated greater reactive self-reported emotional distress suggesting that the RVPFC may
tendencies. Such tendencies would likely result in observed indi- play a role in regulating the dACC.
vidual differences in the many behavioral dimensions of individual Eisenberger and Lieberman (2005) have also linked social and
lives. physical pain arguing that increasing or decreasing social harm will
Canli and Lesch (2007) have analyzed the influence of the sero- correspond to increased or decreased sensitivity to physical pain.
tonin transporter gene (5-HTT) short variant on fMRI measured For example, college students who were told they had performed
activity of amygdala and related brain areas and found that well- poorly on important exams also reported higher pain ratings to a
adjusted and phobic-prone short variant carriers showed greater cold pressor task (Levine et al., 1993; van den Hout et al., 2000). Con-
amygdala activity during tasks such as passive viewing of negative versely, Brown et al. (2003) found that undergraduates reported
vs. neutral pictures or matching fearful and angry faces vs. geo- less pain performing the cold pressor task in the presence of a
metric shapes. Analysis of emotionally negative or neutral stimuli non-evaluative supportive friend or stranger than when alone or
vs. a fixation rest condition showed decreased amygdala activation just with an “interactive” control participant. In medical settings,
to neutral relative to fixation stimuli, which was “interpreted as social support and social activities were associated with less can-
indicating elevated amygdala activation during the processing of cer pain (Zaza and Baine, 2002). Women with continuous social
emotionally undefined stimuli (our italics) in short variant carriers.” support reported less labor pain (Kennell et al., 1991). Coronary-
(p. 1105). bypass patients with more social support reported less pain and
Furthermore, Canli and Lesch proposed a “tonic activation” took less pain medication (Kulik and Mahler, 1989). Given these
model that suggested short variant 5HTT carriers exhibit elevated findings, personality theorists and practitioners must distinguish
amygdala activity at rest compared to noncarriers, which they sup- between distress reflecting social harm avoidance and physical
ported with a perfusion imaging study that measured absolute harm avoidance likely linked with the SADNESS or FEAR systems
amygdala blood flow at rest. Higher baseline amygdala activity lev- respectively. For extensive discussions of the social-pain construct,
els of short variant 5HTT carriers suggested these short variant see MacDonald and Jensen-Campbell (2011).
carriers may experience a different more threatening world than
their noncarrier counterparts. For example, Canli et al. (2006) found
4.3.3. Evidence summary
that short variant 5HTT carriers showed increased rumination in
The above studies link primary emotions to limbic brain activity.
response to life stress compared to noncarriers.
Furthermore, individuals whose brain systems have higher medial
While studies like Canli and Lesch demonstrate the capacity of
cortical “resting-state” levels of activity (see Northoff et al., 2011)
limbic structures to modulate cognitive interpretations of neutral
are more reactive to neutral stimuli and respond differently to
or negative stimuli, the general brain neurotransmitter serotonin
events. It is our position that individual differences in such higher
is not thought to play a specific role in modulating any of the
affective as well as lower primary-process aversive affective brain
primary emotional responses but functions as a general modula-
systems (RAGE, FEAR, and SADNESS) along with the positive affect
tor of all of the primary affective systems, and many other brain
systems of PLAY, CARING, and SEEKING are foundational for per-
functions such as cortical processing (see Panksepp, 1998). Also,
sonality expression as well as the emergence of mental anguish and
mixing fearful and angry face stimuli makes it difficult to inte-
pathology. Individuals with different levels of responsiveness in
grate results into primary brain emotion systems, since affective
these primary brain systems not only react differently to the same
neuroscience would predict that, while the FEAR and ANGER sys-
stimuli, they will experience these stimuli differently and develop
tems are closely related anatomically in lower brain regions, they
different conditioned response tendencies and ongoing personal
remain distinctly separate systems, and my have distinct higher
preferences.
brain regions of influence where raw affects are blended with cog-
nitive information-processing types of strategies.
5. Clinical assessments from affective neuroscience
4.3.2. Social rejection and the SADNESS system trajectories
Similar fMRI research has focused on the SADNESS system and
the relationship of dorsal anterior cingulate cortex (dACC) activ- The FFM is a factor analytic approach to personality, which has
ity in response to social pain. The SADNESS system during infancy been derived from studying “normal” populations. However, psy-
is characterized by distress calls in response to separation from chometric approaches to psychopathology (Livesley, 1986, 1987;
parents and group members. In older individuals, it is thought to Clark, 1990) have demonstrated that personality disorder (PD)
K.L. Davis, J. Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 1946–1958 1955

diagnosis can be conceptualized in terms of the FFM (Clark and In addition to the importance of the CARE system being dis-
Livesley, 2002). Widiger and Costa (2002) reviewed 56 studies that played by therapists, the PLAY system could have powerful and
related the FFM to PD symptomology. Of these studies, 30 used clin- underutilized therapeutic effects as well (Panksepp, 2009). Acti-
ical populations. The authors concluded that “much of this research vating the PLAY system requires the bodily vigor, spontaneity, and
indicates strong support for understanding PD symptomatology as creativity of real “rough-and-tumble” PLAY, which may be most
maladaptive variants of the personality traits included with the easily applicable to children. There is good reason to believe that
FFM” (Widiger and Costa, 2002, p. 80), which suggests that the FFM one of the main functions of the PLAY system is the epigenetic con-
does represent a meaningful continuum of behavior extending into struction of the social brain (Panksepp, 2007b, 2008). Play deprived
the realm of clinical pathology. human children, just like other mammalian young, develop height-
The NEO PI-Revised (Costa and McCrae, 1992) includes six facets ened motivations to play. In humans, this can result in receiving
for each of the FFM dimensions. Some feel that facet level analyses psychostimulant drugs, which are effective inhibitors of physical
are required for optimal discrimination between PDs (Axelrod et al., play urges. Animal models confirm that ADHD impulsivity can be
1997; De Fruyt et al., 2006; Dyce and O’Connor, 1998). However, a reduced with rough-and-tumble play during early development
challenge for the FFM may be identifying facets that are congru- (Panksepp et al., 2003). Ample physical play time for children may
ent with neuroscience research rather than relying on an arbitrary be one of the best ways to protect them against ADHD as well as
number of rationally defined facets, some of which may be best depression.
defined as factor “blends,” e.g. Hostility. We may be at a point “in While adult brains are not as plastic as those of children, play-
which software capabilities now exceed both the quality of data fulness can be a valuable resource for redirecting adults onto a
and the scope of conceptualization” (Wiggins and Pincus, 2002, p. more adaptive life track. Robust physical activity by itself may be as
109) and in which it may be more productive to use functional neu- effective an antidepressant as the medications that dampen emo-
roscience rather than factor analysis to inform and identify new tionality (see Watt and Panksepp, 2009). Play urges in adults may
trait dimensions. For example “empathy” (Chen et al., 2009) and be reenergized by physical activities like sports or dance accompa-
“jealousy” (Panksepp, 2010), and the other socially constructed nied by music that stimulates the rhythmic motor impulses of the
emotions, could emerge as newly defined brain emotions in ani- body (Panksepp and Trevarthen, 2009).
mals that can be used to guide higher-order affective personality
scale construction. In our estimation, it is most important to be very 7. Summary of affective neuroscience perspectives on the
clear about the primary-process issues that undergird personality foundations of personality
development.
Our scientific understanding of personality began with Darwin
and McDougal with their focus on emotion and instinct and the
6. Affective neuroscience and therapeutic effectiveness recognition that the differences between humans and their non-
human ancestors were of “degree” and not of “kind.”
Affective Neuroscience also has important implications regard- In one sense, affective neuroscience helps reconnect personal-
ing therapy for affective imbalances (see Coenen et al., 2011). ity theory back to our evolutionary roots by showing how closely
Panksepp (2009) has emphasized the critical nature of the CARE our personalities are linked to the ancestral affective forces we
system for effective therapy and has even considered how oxytocin share with other mammals and by relating psychopathology to
administration might be used to enhance a supportive nurtur- disturbances in the primary emotional subcortical brain systems.
ing therapist demeanor. Shedler (2010) has summarized data on Effective treatment of behavioral disorders, and long-term thera-
the long-term effectiveness of psychodynamic therapy and like- peutic change, lies in the sensitive exploration of troubling feelings
wise emphasized the importance of a nurturing “working alliance” and reactive patterns and perhaps not as much in changing dis-
between the patient and the therapist. He also discussed research torted cognitions. Consequently, recognizing that our emotional
(see Castonguay et al., 1996) that linked poor therapeutic out- instincts are fundamentally action systems suggests the impor-
comes to rigid insensitive implementation of therapies. In a second tance of incorporating bodily activity into therapy. If a person
point, Shedler also emphasized the importance of “labeled expe- is frozen in an emotional state, getting them to move around
rience” structured so the client “gains awareness of previously and physically interact with the therapists may help promote a
implicit feelings and meaning” (Shedler, 2010, p. 104; quoted shift in affective balance. After all we are dynamically ‘embod-
from Castonguay et al., 1996, p. 499). This suggests that a better ied’ creatures, whose emotionality is closely linked to motor
understanding of primary-process affective feelings at a thoughtful, action systems. In adult psychotherapy, the PLAY system may
tertiary-process awareness level may be therapeutically beneficial. have considerable untapped potential for helping patients reinte-
In support of this approach, Lieberman showed that affect labeling grate troublesome emotional experiences towards more adaptive
decreased amygdala response that was inversely correlated with and emotionally comfortable affective trajectories. For many
the right ventrolateral prefrontal cortex activity, which has been people, bringing the body and emotional actions on to the ther-
previously associated with emotional regulation (Lieberman et al., apeutic field, may open up many opportunities for change that
2007), and that those more skilled in mindfulness enhanced the would not exist as long as therapy consists largely of “talking
effect (Creswell et al., 2007). These two themes have also been heads.”
shown to be important for the success of cognitive behavior ther- An exploration of cross-species, primary-process emotional
apy (CBT), a finding that was replicated by Hayes et al. (1996). A systems of the brain suggests that the FFM itself needs further clar-
third theme, which dealt with CBT’s emphasis on cognitive distor- ification. Combining the key affective dimensions of FEAR, ANGER,
tions, was not supported in these studies. In fact the therapeutic and SADNESS into a general Neuroticism factor detracts from their
aim of cognitive change predicted poorer therapeutic outcomes. unique contributions to personality expression and mental dis-
In a review of evidence-based explanations for psychotherapeu- order. Furthermore, ANGER and CARING need to remain distinct
tic interventions, Kazdin (2007) concluded “whatever may be the dimensions rather than fusing them onto opposite poles of a scien-
basis of changes with CBT, it does not seem to be the cognitions tifically confusing Agreeableness factor. One advantage of distinct
as originally proposed.” (p. 8). Thus, effective therapeutic interven- emotional dimensions is their underlying unipolar nature, inter-
tions may typically impact the affective dimensions and levels of active though they are with other emotions, and having separate
the BrainMind more than cognitive, information-processing ones. dimensions for the ANGER and CARE systems might contribute
1956 K.L. Davis, J. Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 1946–1958

to clearer primary-process personality thinking. Above all, it may Using this 4-point scale with 1 for “Strongly Agree” and 4 for
be helpful to have more precise tools to systematically assess the “Strongly Disagree” (but computing scores equivalent to hand scor-
location of experimental subjects and therapeutic clients in affec- ing above with a base score of 0) the following formulas were used
tive space. And that was our original motivation for developing for “computer scoring” the 6 primary ANPS scales plus Spirituality:
the ANPS. We hope it will be useful for many other research and
practical endeavors. SEEK score = (+21 −ans1 −ans17 −ans33 −ans49 −ans65 −ans81
Note: Parkinson’s disease has been linked with distinctive per- −ans97 +ans9 +ans25 +ans41 +ans57 +ans73 +ans89 +ans105).
sonality differences that include an increased risk for depression. FEAR score = (+21 −ans2 −ans18 −ans34 −ans50 −ans66 −ans82
Remy et al. (2005) used PET analysis to investigate the brain cor- −ans98 +ans10 +ans26 +ans42 +ans58 +ans74 +ans90 +ans106).
relates of depression in Parkinson’s patients and matched controls CARE score = (+21 −ans3 −ans19 −ans35 −ans51 −ans67 −ans83
and found that depressed Parkinson’s subjects had lower activity in −ans99 +ans11 +ans27 +ans43 +ans59 +ans75 +ans91 +ans107).
the locus coeruleus, anterior cingulate cortex, thalamus, amygdala, ANGER score = (+21 −ans4 −ans20 −ans36 −ans52 −ans68
and ventral striatum–all very ancient brain areas. They hypoth- −ans84 −ans100 +ans12 +ans28 +ans44 +ans60 +ans76 +ans92
esized that depression and anxiety in Parkinson’s patients were +ans108).
associated with loss of dopamine and noradrenaline activity in PLAY score = (+21 −ans5 −ans21 −ans37 −ans53 −ans69 −ans85
the limbic system. Damholdt et al. (2011) collected FFM data in −ans101+ans13 +ans29 +ans45 +ans61 +ans77 +ans93 +ans109).
a study of Parkinson’s disease and found that Parkinson’s patients SADNESS score = (+21 −ans6 −ans22 −ans38 −ans54 −ans70
with depression displayed lower Extraversion and higher Neuroti- −ans86 −ans102 +ans14 +ans30 +ans46 +ans62 +ans78 +ans94
cism scores. It would have been interesting to have supplemented +ans110).
the Neuroticism measure with measures of SADNESS, FEAR, and Spirituality score = (+18 −ans7 −ans23 −ans39 −ans55 −ans71
ANGER to assess likely links to discrete brain systems. Perhaps link- −ans87 +ans15 +ans31 +ans47 +ans63 +ans79 +ans95).
ing human personality to our ancestral emotional urges will bring
further clarity to the understanding of human personality. Algebraically, the “hand scoring” and “computer scoring” pro-
cedures are identical.
Acknowledgements Copyright © 2004, Kenneth L. Davis, Ph.D., Jaak Panksepp, Ph.D.,
Pegasus International, Inc. All rights reserved.
We appreciate all the investigators who have taken an inter-
est in the Affective Neuroscience Personality Scales. This work was References
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