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Neuroscience and Biobehavioral Reviews 35 (2011) 19461958

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Neuroscience and Biobehavioral Reviews


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

Review

The brains emotional foundations of human personality and the Affective


Neuroscience Personality Scales
Kenneth L. Davis a, , Jaak Panksepp b
a
b

Department of Psychology, The University of North Carolina at Charlotte, Charlotte, NC 28223-0001, USA
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

Keywords:
Personality
Affective neuroscience
Five Factor Model
Subcortical brain emotion systems
Affective neuroscience
Affective Neuroscience Personality Scales

a b s t r a c t
Six of the primary-process subcortical brain emotion systems SEEKING, RAGE, FEAR, CARE, GRIEF and
PLAY are presented as foundational for human personality development, and hence as a potentially
novel template for personality assessment as in the Affective Neurosciences Personality Scales (ANPS),
described here. The ANPS was conceptualized as a potential clinical research tool, which would help
experimentalists and clinicians situate subjects and clients in primary-process affective space. These
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) processing. Supporting neuroscience research is presented along with comparisons to Cloningers Temperament
and Character Inventory and the Five Factor Model (FFM). Suggestions are made for grounding the internal 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 ve 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.
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Introduction: Darwin, McDougal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .


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

Corresponding author. Tel.: +1 336 379 9828; fax: +1 336 379 9835.
E-mail address: ken@pegasusintl.com (K.L. Davis).
0149-7634/$ see front matter 2011 Elsevier Ltd. All rights reserved.
doi:10.1016/j.neubiorev.2011.04.004

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Affective neuroscience and therapeutic effectiveness . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .


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

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1. Introduction: Darwin, McDougal


Charles Darwin provided the rst modern scientic treatise on
emotion in 1872 with The Expression of the Emotions in Man
and Animals (Darwin, 1872/1967). In this classic work, Darwin
documented cross species commonalities that occurred for many
primary emotions such as fear, anger, grief, and joy, which included
a discussion of playful tickling in human children and many other
descriptions of human emotional subtleties. Darwins keen observations, which remain relevant to this day, allowed him to extend
his robust case for the continuity of species from body structures to the domain of emotions. In line with his views, a recent
spectrographic analysis of the tickle-induced laughter response
on four species of young great apes plus humans yielded data
that effectively mapped the scientically established evolutionary
branching of these closely related anthropoids (Ross et al., 2009).
Likewise, in synergy with Darwins vision, cross-species affective
neuroscientists have started to map out the continuity of primal
emotional systems in subcortical-limbic circuits of selected mammalian brains (Panksepp, 1998) from mice to men, so to speak.
The anatomical, neurochemical, and functional homologies of
subcortical emotional networks strongly indicate underlying evolutionary continuities in affective principles in all mammalian
brains (Panksepp and Biven, 2011). Since it is widely believed that
individuals emotional inclinations (temperaments) are foundational for human personality development, the goal of the following
work was to try to ground personality assessment on the affective circuits that all mammals share. But before we summarize
work with the Affective Neuroscience Personality Scales (ANPS),
we briey note historical antecedents of this way of thinking about
human personality.
In 1908 William McDougal published his classic work Social
Psychology in which he reviewed the inuence of instincts on
human personality. He was the rst personality theorist to be
guided by Darwins principle of the continuity of human and animal mental evolution, and he used two principles for determining
whether an emotion should be considered a primary emotion,
i.e. simple instinctive impulse. The rst was whether the emotion
was clearly displayed in the instinctive activities of the higher animals (McDougal, 1908/1963, p. 42). Hence, he rejected that there
was any religious instinct, and various other debatable higherorder affective-cognitive processes. His second principle was that
a primary emotion should be evident in human psychopathologies,
with emotional behaviors being manifested in extreme, exaggerated, and hence abnormal displays, which is consistent with
certain current views of mental disorders (see below). He included
only seven behaviorally well-dened emotional instincts, which he
labeled ight, repulsion, curiosity, pugnacity, self-abasement, selfassertion, and the parental instinct, a list based on comparative
observations that is arguably still relevant to personality theory a
century later, although those strands of early ethological thought
were never fully developed.
2. Subcortical emotion systems
Walter Hesss seminal work on evoking emotions with localized
electrical stimulation of the brain (research summarized in Hess,
1957) demonstrated the importance of discrete subcortical areas

of the brain for generating specic emotional-behavioral episodes,


along with characteristic accompanying autonomic arousals. In
1949, he received the Nobel Prize for his work on the central representations of the autonomic nervous system. As ruefully related
in his autobiographical materials, Hess resisted, with regrets, any
explicit discussion of emotional experiences in the animals he studied, because he feared marginalization by behaviorists. He indicated
that he always believed these irritated animals were experiencing a state akin to human anger. He never inquired whether the
animals found such evoked states of anger to constitute aversive
experiences. Panksepp (1971) later demonstrated an homologous
anatomy in rats, and proceeded to show that, given the opportunity,
animals would escape the RAGE evoking stimulation.
Panksepp and some others continued these lines of research,
with explicit discussions of the accompanying affective experiences, based on whether the ESB (electrical stimulation of the brain)
evoked reward or punishment effects in simple learning tasks.
Panksepps (1982, 1998, 2005) synthesis of such work provides
documentation for the existence of seven blue ribbon subcortical emotion systems in the brain, which he labeled SEEKING, RAGE,
FEAR, LUST, maternal CARE, separation-distress PANIC/GRIEF, and
physical PLAY. The capitalizations were premised on the need for
a specialized terminology for the primary-process anoetic experiential processes of the brain, which together constitute forms of
affective consciousness (Panksepp, 2007a)
This nomenclature allows the use of common emotional terms
that convey the gist of the emotional meaning, while minimizing
mereological fallacies (part-whole confusions) that are endemic to
the use of emotional terms in describing animal experiences, thus
providing some distance from dictionary denitions and everyday understandings. We will never know how any other animal
or any other human actually feels. This is the classic dilemma
that subjectivity poses for modern neuroscience. All mammalian
brains may be fundamentally subjective organs they not only
generate behaviors, but mental processes such as basic feelings
and perceptions (anoetic qualia), which control learning and memory (noetic secondary-processes) as well as higher mental abilities
(autonoetic tertiary processes) such as thoughts, plans and insights,
aspects of mind that can only really be well-studied in humans
(for a coverage of such nomenclature issues, see Vandekerckhove
and Panksepp, 2009 and this issue). In contrast, feelings, because of
their capacity to engender rewarding and punishing states in the
brain, can be objectively studied in a substantive phenomenological manner. Neuroscience advances will be substantially premised
on our ability to conceptualize brain functions both from the
external behavioral view as well as the internal psychological
perspective.
We sustained the capitalization convention in our attempt (i.e.,
the ANPS) to translate anoetic (without knowledge) emotional
experiences to noetic (knowledge linked) and autonoetic (thought
linked), language-based evaluations of the potential inuences of
primal affects in the personality structures of human minds. However, rather than RAGE and PANIC/GRIEF, we decided to use the
terms ANGER and SADNESS in this paper and in our construction of
the Affective Neuroscience Personality Scales (ANPS), since those
are labels that are generally more understandable for most individuals. Still, the theoretical implication is that these systems are
critically important for the various feelings highlighted by these sci-

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K.L. Davis, J. Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 19461958

entic terms e.g., the ANGER system energizes states commonly


labeled as irritability and anger.
Neuroanatomical, neurochemical, pharmacological, and physiological brain research have helped to more precisely dene each of
these seven emotion systems (see Panksepp, 1998, 2005; Panksepp
and Biven, 2011 for detailed reviews). We assumed these brain
systems could serve as an empirical roadmap for advances in the
study of personality and serve as a template for emotional personality assessment, as in the ANPS (Davis et al., 2003), which was
conceptualized largely as a research tool aspiring to, and perhaps
capable of, situating adult human temperaments within the complexities of primary-process affective space shared by all mammals.
If so, that could also open up a common terminology for study personality across species. It is a major supposition of this paper that
these basic emotions are fundamental powers of the human BrainMind, homologous with feelings experienced by other mammals,
and hence potentially of critical importance for understanding the
foundations of personality (we intentionally conate brain and
mind, and double capitalized BrainMind to reect our monistic
ontology).
There is strong evidence for these seven well-dened emotion systems in all mammalian brains. Several of these emotions
have evolutionally deep reptilian roots (SEEKING, ANGER, FEAR and
LUST), but three (CARING, SADNESS, and PLAY) are more uniquely
mammalian adaptations, but surely with antecedents in lower
vertebrates, perhaps invertebrates also (e.g., see Huber et al., 2011).
Each of these seven emotions can be evoked by stimulating distinct
but partly overlapping subcortical regions of mammalian brains.
However, they typically work interactively in concert to increase
the adaptiveness of individual feelings, perceptions, thoughts, and
behaviors. (see Panksepp, 1998, 2005; Panksepp and Biven, 2011
for fuller descriptions).
There are several principles basic to understanding these seven
brain systems: (1) These emotion systems are subcortical networks
and lower brain regions have evolutionary primacy in generating
these basic emotions and their affects, while learning and higher
brain functions can be deemed to be secondary and tertiary processes. (2) To the best of our knowledge, these emotion systems,
situated in ancient brain regions, are largely homologous in all
mammals. (3) These basic emotions also have similar chemistries in
all mammals. (4) These brain systems generate instinctual behavioral responses that are closely linked to the raw, primal affects
that accompany those responses. (5) The integrity of these seven
systems is demonstrated by the ability to elicit coherent specic
emotional responses and/or the associated affects with localized
brain stimulation as evaluated by the capacity of the subcortical
arousals to mediate reward and punishment functions that control learning. (6) Lastly, these systems remain relatively unscathed
in animals whose neocortices were surgically removed in early
development (see Panksepp, 1998).
In summary, emotional responses from each of the primary
process emotions can be activated by localized subcortical ESB or
chemical brain stimulation. That decortication of young animals
generally leaves the expression of these emotions intact (Deyo
et al., 1990; Panksepp et al., 1994), further reinforces the subcortical nature of these emotion systems. Also, each instinctual
emotional system engenders affective valence since animals terminate the stimulation of ANGER, FEAR, and SADNESS, while actively
working to obtain SEEKING, LUST, CARE, and PLAY system arousals
(Panksepp, 2005). However, there are reasons to believe the SEEKING system is foundational for aspects of all of the other primal
emotions (e.g., during FEAR intensication, the shift from freezing
to ight may be due to recruitment of SEEKING related dopaminergic psychomotor drive).
It is our premise that these emotions form an important
foundation of personality. As such, personality assessment can

be informed by these subcortical affects. Specically, studies of


primary-process emotional aspects of personality may provide a
heuristic intersection for the Darwinian continuity of species,
with the cross-species emotion systems of mammalian brains, and
the need for neurobiologically meaningful psychological assessment of human temperaments/personalities.
But this is not the rst attempt to bridge basic neuroscience
and human personality evaluation. The two individuals who have
attempted to construct neurobiologically based personality assessment instruments in the past are Cloninger (1986) who based
his thinking on functional animal work on some of the rst well
characterized neurochemical systems of all mammalian brains
dopamine, norepinephrine, and serotonin (see below). In contrast, Carver and White (1994) developed the Behavioral Activation
and Behavioral Inhibition Scales on the basis of Jeffrey Grays
(1981) early brain research recognizing the existence of behavioral
approach/arousal and withdrawal/inhibition (aka, reward and punishment) systems in the brain. The present work is an extension of
this strategy, as our understanding of brain affective systems has
become more resolved. Because of space constraints, we will only
contrast our work on the ANPS, designed to estimate six affects relevant for dening temperamental/personality dimensions, with the
popular Cloninger assessment tool following a brief introduction of
the ve factor personality model.
3. The Five Factor Model and the ANPS
At the very outset of this work, the ANPS was hypothesized
to provide affective underpinnings to the widely used Five Factor Model (FFM). The FFM has provided a personality assessment
standard, which while not universally accepted (Block, 1995) has
offered a robust personality model in general population and clinical research (Costa and Widiger, 2002). The FFM has origins in
adjective descriptors of personality (Allport and Odbert, 1936)
and the early history of factor analysis (Cattell, 1947). The ve
personality dimensions are usually labeled Extraversion or Surgency, Agreeableness, Conscientiousness, Emotional Stability, and
Intellect or Openness to Experience (Hofstee et al., 1992). These
ve dimensions have been consistently derived from a variety of
descriptive data using factor analysis (Digman, 1990) and have
been replicated in several languages (Saucier and Goldberg, 2006).
However, the rst four dimensions have been more robust with
Openness to Experience typically accounting for the least variance
of the ve factors (Goldberg, 1990). A comprehensive presentation of the ve factor descriptive model was provided by Hofstee
et al. (1992) and consisted of an analysis of over 500 adjectives
placed on ve orthogonal dimensions as well as numerous off-axis,
two-factor blends, such as angry emerging as a blend of low
Emotional Stability and low Agreeableness.
3.1. Cloningers biologically based personality theory
Cloninger was the rst to attempt an integration of personality
and brain systems with his biogenic amine-based biosocial theory
of personality (Cloninger, 1987) and the Tridimensional Personality Questionnaire (TPQ), which included three scales: (1) Novelty
Seeking based on a postulated dopaminergically based behavioral
activation system that promotes interest in new experiences, (2)
Harm Avoidance, postulated to primarily reect brain serotonin
activity, the brains presumed punishment system (which is highly
debatable, see Panksepp, 1986, 1998), and the resulting sensitivity to threatening situations, and (3) Reward Dependence featuring
behavioral maintenance and sensitivity to reinforcement contingencies with norepinephrine as the major neuromodulator. Each of
these temperament traits was seen as highly heritable and stable
across the developmental stages of life.

K.L. Davis, J. Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 19461958

Our affective neuroscience perspective would align the Novelty


Seeking dopaminergic brain incentive dimension with the SEEKING system. The SEEKING system not only promotes exploration,
investigation and foraging but probably energizes all basic emotional systems with forms of appetitive and anticipatory arousal
(e.g., as already noted, seeking of safety when FEAR is aroused).
We suspect that serotonin and norepinephrine may function too
broadly to be isolated as specic contributors to very specic
types of affects the way Cloninger proposed, even though they
gure heavily in the arousal dimensions of all personality control
systems. Norepinephrine promotes attention by amplifying signalto-noise processing in most sensory-perceptual channels and also
promotes emotionality in general. In contrast, serotonin dampens
and restricts neural processing in the brain, resulting generally in
reduced emotionality (Panksepp, 1986).
Cloninger subsequently expanded his assessment instrument
adding an additional temperament dimension Persistence and
three controversial character traits Self-Directedness, Cooperativeness, and Self-Transcendence hypothesized to have low
heritabilities and be more sensitive to individual development (we
would conceptualize these as higher-order, tertiary-process type
BrainMind functions). The revised instrument was called the Temperament and Character Inventory (TCI; Cloninger, 1999). Although
Cloningers earlier TPQ factors have been related to many genetic
variables, many of Cloningers later theoretical claims have not
been supported. In other words, heritabilities did not differentiate the temperament and character scales (Gillespie et al., 2003;
Cloninger, 2004). Other research found that the Self-Directedness
character scale also responded to serotonin treatments (Peirson
et al., 1999; Wai and Bond, 2001). Factor analysis placed the temperament Harm Avoidance and the character Self-Directedness on
the same factor, and hence did not support a distinction between
temperament and character dimensions, and accounted for
the revised TCI scales with ve factors (Farmer and Goldberg,
2008a,b). Others also reported considerable convergence between
Cloningers revised TCI (Cloninger, 1999) and the FFM (Ball et al.,
1999; De Fruyt et al., 2000).
3.2. ANPS data
The Affective Neuroscience Personality Scales (ANPS) were
introduced to explore the potential primary-process brain emotional system foundations of personality. The scales were rationally
dened to potentially provide biological underpinnings for the
descriptive FFM of personality (Davis et al., 2003). Our view is that
these personality dimensions escape the problem of denitional
circularity since comparative research has demonstrated how each
primary dimension can be manipulated in humans as independent variables including experientially (Eisenberger et al., 2003;
Panksepp, 1985) or pharmacologically as well as through brain
damage research (e.g., Bechara et al., 1999).
Each ANPS dimension corresponded to one of the blue ribbon
primal emotions identied by cross-species ESB-based affective
neuroscience research. We excluded LUST, which seems less relevant to current conceptualizations of human personality and we
also suspected that it may potentially be an affective factor that
people would not wish to be frank about, and thus may contaminate frankness on the other scales. In addition, a Spirituality scale
was included to acknowledge the importance of this factor in drug
addiction (Panksepp et al., 2004) and many drug addiction recovery
programs (e.g., 12-step programs) and what might be considered
the highest human emotion.
The PLAY scale focused on playing social games with physical
contact plus laughter, humor, and generally having fun. SEEKING
was dened as anticipating new positive experiences including
being curious, liking to strive for solutions to problems, and gener-

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ally liking to explore. CARING centered on nurturing tendencies


including liking to care for others, being drawn to young children and pets, and feeling softhearted towards animals and people
in need. The FEAR scale incorporated experiencing anxiety, worrying, difculty making decisions, ruminating, feeling tense, and
losing sleep. ANGER included feeling hotheaded, being easily irritated and frustrated, and expressing anger verbally or physically.
SADNESS was conceptualized as feeling social separation distress,
feeling lonely, and thinking about loved ones and past relationships including crying. The Spirituality scale focused on feelings of
connectedness with all of life and oneness with creation.
The revised ANPS 2.4 items are listed in Fig. 1 (for details, see
Appendix A). These scales are provided for free use in all scientic endeavors and non-commercial clinical use (e.g., individual
psychotherapy practitioners) without need to obtain permission
of the copyright owners (Davis and Panksepp). Those wishing to
use it for commercial use are requested to contact the rst author.
Again, a detailed description of the scales and scoring instructions
are provided in the appendix at the end of this paper.
Based on our rst study using these scales, reliabilities for the
ANPS scales, computed as Cronbachs alpha, were reported ranging from .65 to .86 with the PLAY and SEEK scales below .70 and
the FEAR, ANGER, and Spirituality scales above .80 (Davis et al.,
2003). Multiple data sets using the revised ANPS 2.4, have shown
all reliabilities now over .70 (unpublished data).
Contrasting self-rating data for ANPS and FFM scales, as seen
in Table 1, revealed that each of the ANPS scales except Spirituality
correlated strongly with at least one of the FFM scales as follows: (1)
SEEK with Openness to Experience (r = .47), (2) PLAY with Extraversion (r = .46), (3) CARE with Agreeableness (r = .50), (4) FEAR with
Emotional Stability (r = .75), (5) SADNESS with Emotional Stability (r = .68), and (6) ANGER with Emotional Stability (r = .65)
as well as with Agreeableness (r = .48) (Davis et al., 2003). The
data supported strong relationships between primary emotions
and the most widely accepted model of human personality, which
was consistent with the hypothesis that these six brain emotion
systems form a foundation for the adult ve-factor personality
model.
When factor analyzing ANPS scales with FFM scales, we reported
a ve factor solution with the FFM Conscientiousness scale on a
factor by itself. Since there were only four eigenvalues > 1.0, we
removed the Conscientiousness scale and published a four factor
solution with (1) FEAR SADNESS, ANGER, and Emotional Stability,
(2) CARE and Agreeableness, (3) PLAY and Extraversion, and (4)
SEEK and Openness to Experience loading on four separate scales
with .68 being the lowest primary factor loading. The only scale
having a loading on a second factor greater than .30 was ANGER,
which also loaded .53 on the Agreeableness factor. In summary,
the ANPS offers personality primes that align with the Big Five
model but target the six specic primary emotion brain systems
that have been well documented with comparative brain research.
A factor analysis of just the six primary ANPS scales yielded
two eigenvalues greater than 1.0 with FEAR, SADNESS, and ANGER
on the rst component and PLAY, CARE, and SEEK on the second
with all primary factor loadings greater than .55 and all secondary
loadings less than .20. This clustering, corresponds nicely to other
scales which simply try to parse human personality into positive
and negative affective dimensions (Watson et al., 1988) and generally replicates Grays (1970) Behavioral Inhibition and Approach
Systems. We feel this is a striking demonstration of the multifactorial nature of any general positive and negative valence type of
measurement tool. In this vein, it is also important to note Goldberg
and colleagues argument (Ashton et al., 2009, p. 80) that higher
order factors such as Positive Affect, Negative Affect, and overall
Emotional Stability may represent artifacts rather than substantive dimensions of personality.

1950

K.L. Davis, J. Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 19461958

Pahlavan et al. (2008) conrmed the psychometric properties


of the ANPS in a French population including scale reliabilities, factor structure, and correlations with the FFM. Berthoz
and colleagues (personal communication) replicated the Pahlavan
et al. (2008) ndings on a large sample (n = 830) and also
performed an item-level conrmatory factor analysis obtaining an acceptable t modeling 6 factors (standardized root
mean square residual = .06, root mean square error of approximation = .039, and the comparative t index = .78). Savitz et al.

(2008c) obtained signicant evidence of heritability for four of


the ANPS scales: CARE, SEEK, FEAR, and Spirituality. Through personal correspondence, we know the scale has been translated into
a dozen languages, and has already proved its utility in evaluating certain psychiatric problems (e.g., Savitz et al., 2008a,b;
see below). For a while Martin Reuter (University of Bonn) had
all the original scales readily available on the web, and also for
data collection purposes (www.anps.de), but this site has been
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.
mdanps24

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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 gure legend and Appendix A.

K.L. Davis, J. Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 19461958

Please mark bubbles like this

and not like this

or

Disagree
Str Disagree

1951

Agree
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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mdanps24
Copyright 2004, Kenneth L. Davis, Ph.D., Jaak Panksepp, Ph.D., Pegasus International, Inc. All rights reserved.
Fig. 1. (Continued ).

3.2.1. Conscientiousness
Conscientiousness was the only FFM dimension that did
not correlate strongly with the ANPS. Lower correlations with
ANGER (r = .30), FEAR (r = .24), and SADNESS (r = .30) suggested that while Conscientiousness may be associated with the
regulation of negative affects, it seems unlikely that a single
primary brain affect serves as the foundation for Conscientiousness. From animal FFM studies, Conscientiousness only appeared

in chimpanzees (Gosling and John, 1999), which suggests that


Conscientiousness is a more cerebral dimension emerging late
in mammalian evolution. Congdon and Canlis (2008) view of
impulsiveness as a top down lack of behavioral inhibition
involving the inferior frontal cortex and subthalamic nucleus ts
well with Kings (2007) description of chimpanzee Conscientiousness characterized by predictable, not impulsive or erratic,
behaviors.

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K.L. Davis, J. Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 19461958

Table 1
ANPS Scale correlations with Five Factor Model scales.

PLAY
SEEK
CARE
FEAR
ANGER
SADNESS
Spirituality

Extraversion

Agreeableness

Conscientiousness

Emotional stability

Openness to experience

.46*
.13
.25
.19
.04
.21
.15

.29*
.01
.50*
.17
.48*
.13
.26*

.00
.01
.12
.24
.30*
.30*
.14

.12
.01
.07
.75*
.65*
.68*
.09

.13
.47*
.06
.05
.08
.00
.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


reects primary subcortical affects. Although ANPS items attempt
to address primary affects directly, since all self-report assessments
must include cognitive reection, we interpret the ANPS scales as
tertiary (thought-mediated) approximations of the inuence of the
various primary emotional systems in peoples lives. However, it
is our working hypothesis that the subcortical primary-processes
neural systems, where the foundations of emotions reside, can generate individual differences in normal personality as well as the
affective imbalances characterizing mental disorders.
3.2.2. Non-human personalities
If subcortical primary-processes are foundational for the FFM,
one should be able to identify these same FFM dimensions in
non-human mammals. In a review of 19 studies of personality
in non-human animals, Gosling and John (1999) found consistent
cross-species evidence for the FFM in mammals ranging from rats to
chimpanzees. The most consistent FFM temperament dimensions
observed were Extraversion and Emotional Stability. Agreeableness was the next most common. Consistent with human data
(Goldberg, 1990), the evidence for Openness to Experience was
weaker than for the previously named three dimensions. As noted
previously, Conscientiousness was only observed in chimpanzees,
humans closest evolutionary relative. These ndings support the
Darwinian concept of continuity of species that the difference in
the mind between man and the higher animals, great as it is, certainly is one of degree and not of kind. (Darwin, 1872/1967, p.
104)
3.3. Expansion of the FFM
These ANPS results also revealed FFM limitations such as FFM
Emotional Stability associating with all three of the ANPS negative affects, FEAR, SADNESS, and ANGER, which makes Emotional
Stability equivalent to Negative Affect. A recent meta-analysis of
past brain imaging studies also provided strong evidence that distinct anatomical systems in human brains can be related to basic
affective processes including support for sadness, fear, and anger
(Vytal and Hamann, 2010). At the tertiary process level, humans
may not always differentiate well between the distressful feelings
associated with FEAR, SADNESS, and ANGER, which could lead to
statistical lumping, but that does not mean it is desirable to combine three of our most powerful emotional drivers into a single
dimension.
ANGER also correlated with Agreeableness. Despite the fact that
the ANPS ANGER and CARE did not correlate (r = .036 in Davis
et al., 2003), the two scales correlated in opposite directions with
the FFM Agreeableness scale. Human minds seem to generate a
descriptive love/hate Agreeableness construct with caring, nurturing feelings on the positive pole and hostile, demanding feelings
on the negative pole. Our view is that while the FFM is an important theory-free, empirical step forward in parsing personality, it

reects a tertiary cognitive-type reconguration of the underlying


primary brain emotion systems.
3.4. Examples of studies using ANPS
Several of the ANPS scales have been further validated by
independent research. As an example of ANPS research, studying families with a member having bipolar disorder (BPD), Savitz
et al. (2008a) found that the ANPS SADNESS scale was signicantly
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 generalized anxiety disorder. Consistent with the quantitative genetic
model of bipolar spectrum illness (Evans et al., 2005), the ANPS
SADNESS and FEAR scores trended highest for individuals diagnosed as BPD I or II, lower for individuals with recurrent major
depression or a single depressive episode, and lowest for the two
control groups. However, after controlling for self-rated depression (Beck Depression Inventory (Beck and Steer, 1993)) or mania
(Altman Self-Rating Mania Scale (Altman et al., 1997)), only the
SADNESS scale continued to show signicant differences across
groups.
Savitz et al. (2008b) also studied the distribution of hypomanic, cyclothymic, and hostile traits in BPD probands and their
families. After controlling for age, gender, depression, and mania,
BPD II patients had signicantly higher ANPS ANGER scores than
individuals with only a single previous depressive episode, and
higher ANGER scores than unaffected family members. Savitz
et al. (2008b) also reported that Beck Depression Inventory
scores were signicantly correlated with higher ANGER scores and
lower SEEK and PLAY scores while the Altman Self-Rating Mania
Scale signicantly correlated with higher ANGER, SEEK, and PLAY
scores.
The construct validity of the ANPS ANGER scale was also supported by Reuter et al. (2009) investigating links between ANGER
and the rs907094 (aC T single nucleotide polymorphism of the
DARPP-32 gene). Dopamine- and cAMP-regulated phosphoprotein,
32 kDa (DARPP-32) is a key regulatory molecule in the dopaminergic signaling pathway. The C-allele is more common in Sub-Saharan
Africa but rather infrequent in European populations. The genetic
analysis of German subjects without psychopathology showed that
carriers of the T-allele had signicantly higher ANPS ANGER scores,
with CC genotypes having signicantly lower ANGER scores. MRI
data on a subsample of subjects revealed that ANGER scores correlated negatively with gray matter volume in the left amygdala.
Reuters group has also recently observed that the homozygous
long-variant of the serotonin transporter polymorphism and the
TT variant of the single nucleotide polymorphism rs2268498 on
the oxytocin receptor gene showed signicantly lower scores on
the personality dimensions FEAR and SADNESS of the ANPS, as well
as on the overall super-factor of Negative Emotionality, than carriers of the other serotonin transporter and oxytocin gene variants
(Montag et al., 2011).

K.L. Davis, J. Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 19461958

Reuter et al. (2005) also demonstrated a link between the ANPS


SEEK scale and creativity. Using gural, verbal, and numeric creativity tests, subjects with high SEEK scores had higher numeric
creativity scores and were signicantly superior on gural and verbal creativity. SEEK scores also explained more than 15% of the
variance of total creativity. Using intelligence tests as a covariate indicated that the relationship of SEEK to creativity was not
moderated by intelligence.
4. Affective neuroscience trait versus theoretical expansion
In addition to suggesting a reprioritized selection of personality traits in the FFM, affective neuroscience (Panksepp, 1998)
suggests an enhanced neuroscientically premised personality
research model. By providing a putative physiological basis for
FFM traits, affective neuroscience helps ve-factor theory escape
circular reasoning by suggesting brain mechanisms that can be
manipulated to inuence the expression of personality. However,
even more important may be helping to advance personality theory
from trait and situationist approaches to interactionist and ontogenetic personality models (see Ballantyne, 1995 for a more detailed
discussion of personality theory levels).
One ontogenetic model is Scarr and McCartneys (1983) theory
of genotype-environment effects hypothesizing that individuals
make their own environments. The theory predicts that earlier
in life, individuals experience more passive environmental effects
through the environment provided by their parents. However,
there is also an evoked effect, since infant individual differences
can inuence the kind of care infants receive from their parents.
While the passive effect is hypothesized to decrease with ontogenetic development, the evoked effect continues throughout an
individuals life. However, there is also an active effect in which
the individual selects from different environments. This active
selection of environments becomes relatively more important as
individuals mature and are more able to choose the circumstances
and experiences they nd attractive, which accounts for adopted
siblings not resembling each others personalities by late adolescence (Scarr et al., 1981; Scarr and McCartney, 1983).
This genotype-environment interaction model can be tested
with an affective neuroscience approach to personality. At the
secondary process level characterized by simple conditioning and
learning, conditioned place preferences or aversions provide a
mechanism for dening the attractiveness of environments. Pleasurable experiences result in nding similar situations attractive
in the future with traumatic or painful experiences having the
opposite effect. One example prediction would be that individuals who were more sensitive to social pain (for a full discussion of
this construct, see Panksepp, 2011 as well as other contributions
to understanding psychic pain in that book edited by MacDonald
and Jensen-Campbell) would nd such distressing situations more
aversive perhaps at lower thresholds and exhibit more avoidant
behavior after those experiences than individuals who were less
sensitive to social pain.
Early experience epigenetic models have already been tested
in rats. Focusing on the PLAY system, extra provisioning of roughand-tumble play decreased ADHD-type impulsiveness (Panksepp
et al., 2003). Centering on the CARE system, superior maternal
nurturing positively contributed to the stress tolerance of offspring (Francis et al., 1999). Since personality traits can also be
assessed in animal subjects (Gosling and John, 1999; Gosling, 2008),
such longitudinal effects of primary-process experiences can be
tested with animal models that incorporate variables and controls
that would not be possible with human subjects. In shorterlived species, one could also test Scarr and McCartneys (1983)
theory that the personalities of fraternal and adoptive siblings
would become less similar as they mature whereas genetically

1953

identical siblings (whether adopted or not) would converge over


time.
4.1. Primary affective consciousness
Another affective neuroscience concept related to personality is
the multi-tiered framing of consciousness. At the primary emotion
level of consciousness, it is clear that young animals and humans
without a neocortex are still conscious creatures with primitive affective and perceptual capacities (Merker, 2007; Shewmon
et al., 1999) including the capacity to play (Panksepp et al., 1994).
This primary-process affective consciousness may include various
sensory/perceptual feelings but especially includes ancient emotional/motivational experiences all mammals share that provide
the basis of a primordial evaluative system (Kahneman, 2003,
p. 701). This primary level makes itself felt throughout our lives
but is soon augmented by secondary consciousness reecting basic
conditioning-learning. In humans, and perhaps some other cortically well-endowed animals, thoughts about how external events
relate to internal events yield even higher tertiary forms of consciousness consisting of thoughts about thoughts, awareness of
awareness, and the linguistic/symbolic transformation of experience at which humans excel (for overview, see Vandekerckhove
and Panksepp, 2009 and in this issue). However, it is the primaryprocess level of affective consciousness that encodes biological
values through a diversity of raw positively and negatively valenced
emotional experiences, from pain to joy, that may be foundational
for personality as well as disorders of personality, which reect
imbalances in these intrinsic value systems of the BrainMind.
4.2. Could the fundamental nature of personality be affective?
A key question from a purely cognitive perspective is how
non-humans with their more limited cerebral cortical capacities
compared to humans can have distinct personalities at all, let alone
personalities dened by the well accepted FFM dimensions (see
Gosling and John, 1999). Our theory is related to the idea that
the affective foundations of personality lie in MacLeans (1990)
sub-neocortical limbic and reptilian areas of the central nervous system. It is our position that in these evolutionally older
parts of the brain, much more ancient than neomammalian
cortex, one nds the most important evolutionary roots of personality. Furthermore, this homologous foundation consists of
instinctual emotion action systems shared by all mammals. These
ancient tools for living engender a within-brain type of emotional affective valence, as well as behavioral reactive guidance,
and general physical adaptation to survival challenges faced by
our ancestors for millions of years. Thus, they were built into
the BrainMind as ancestral memories. These primary systems
are undoubtedly elaborated during human development by secondary conditioning and tertiary thoughts and self-reections,
but the evolutionary origin and foundational power of these discrete, persistent systems is pre-human.
4.3. State vs. channel functions
The inuence of these basic emotion systems is pervasive, affecting not only our actions and reactions but also our perceptions,
thoughts, and memories. The mechanism mediating these old brain
inuences may lie in the distinction between channel and state
brain functions (Mesulam, 2000). For state modulation, each cortical area receives inputs arising from limbic neurons, which can
modulate activity in the entire cerebral cortex. These inuences
determine the state of information processing rather than the
content being transmitted along the point-to-point channels.
These modulatory corticopetal projections to the cortex are not

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K.L. Davis, J. Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 19461958

balanced by corresponding corticofugal connections from the


cerebral cortex to the subcortex. This asymmetry allows the limbic
system to rapidly shift information processing states throughout
the cerebral cortex . . . [and] alter the tone, coloring, and interpretation of experience rather than its content (Mesulam, 2000, pp.
7879).
4.3.1. Evidence
Evidence supporting such state modulation comes from many
sources, and we just touch on a few: Wu et al. (1991) report that
patients with generalized anxiety disorder exhibited stronger temporal and frontal cortex PET responses to a passive viewing task
compared to normal controls. Also, individuals screened for lack of
a mental health treatment history but with higher Beck Depression
Inventory (BDI) scores exhibited heightened resting fMRI amygdala
activity to neutral visual stimuli (Way et al., 2010). Furthermore,
when compared to subjects with lower BDI scores, these higher
scoring BDI subjects also showed greater amygdala reactivity when
viewing fearful or angry faces. In other words, differences in subcortical states biased perceptions and generated greater reactive
tendencies. Such tendencies would likely result in observed individual differences in the many behavioral dimensions of individual
lives.
Canli and Lesch (2007) have analyzed the inuence of the serotonin transporter gene (5-HTT) short variant on fMRI measured
activity of amygdala and related brain areas and found that welladjusted and phobic-prone short variant carriers showed greater
amygdala activity during tasks such as passive viewing of negative
vs. neutral pictures or matching fearful and angry faces vs. geometric shapes. Analysis of emotionally negative or neutral stimuli
vs. a xation rest condition showed decreased amygdala activation
to neutral relative to xation stimuli, which was interpreted as
indicating elevated amygdala activation during the processing of
emotionally undened stimuli (our italics) in short variant carriers.
(p. 1105).
Furthermore, Canli and Lesch proposed a tonic activation
model that suggested short variant 5HTT carriers exhibit elevated
amygdala activity at rest compared to noncarriers, which they supported with a perfusion imaging study that measured absolute
amygdala blood ow at rest. Higher baseline amygdala activity levels of short variant 5HTT carriers suggested these short variant
carriers may experience a different more threatening world than
their noncarrier counterparts. For example, Canli et al. (2006) found
that short variant 5HTT carriers showed increased rumination in
response to life stress compared to noncarriers.
While studies like Canli and Lesch demonstrate the capacity of
limbic structures to modulate cognitive interpretations of neutral
or negative stimuli, the general brain neurotransmitter serotonin
is not thought to play a specic role in modulating any of the
primary emotional responses but functions as a general modulator of all of the primary affective systems, and many other brain
functions such as cortical processing (see Panksepp, 1998). Also,
mixing fearful and angry face stimuli makes it difcult to integrate results into primary brain emotion systems, since affective
neuroscience would predict that, while the FEAR and ANGER systems are closely related anatomically in lower brain regions, they
remain distinctly separate systems, and my have distinct higher
brain regions of inuence where raw affects are blended with cognitive information-processing types of strategies.
4.3.2. Social rejection and the SADNESS system
Similar fMRI research has focused on the SADNESS system and
the relationship of dorsal anterior cingulate cortex (dACC) activity in response to social pain. The SADNESS system during infancy
is characterized by distress calls in response to separation from
parents and group members. In older individuals, it is thought to

be elicited by social rejection by peers and general social isolation.


Eisenberger et al. (2003) did an fMRI analysis of brain changes as she
manipulated painful social exclusion experienced by participants
supposedly playing a virtual ball-tossing game with other participants. During an initial control scan, subjects just watched other
participants play the virtual ball-tossing game, ostensibly because
technical difculties prevented them from participating. In the
second scan, subjects were linked in with virtual participants supposedly in other scanners and included in the virtual ball-tossing
game. To create a sense of social exclusion, during the nal scan
subjects were socially excluded when the other players (actually a
computer program) stopped tossing the ball to the subject. Subjects
showed signicantly more dACC activation during the social exclusion condition compared to inclusion. Measures of self-reported
distress also correlated positively with dACC activity supporting
the relationship between dACC activity and felt emotional distress
during the social exclusion condition. In addition, right ventral prefrontal cortex (RVPFC) was also more active during social exclusion
compared to inclusion conditions but negatively associated with
self-reported emotional distress suggesting that the RVPFC may
play a role in regulating the dACC.
Eisenberger and Lieberman (2005) have also linked social and
physical pain arguing that increasing or decreasing social harm will
correspond to increased or decreased sensitivity to physical pain.
For example, college students who were told they had performed
poorly on important exams also reported higher pain ratings to a
cold pressor task (Levine et al., 1993; van den Hout et al., 2000). Conversely, Brown et al. (2003) found that undergraduates reported
less pain performing the cold pressor task in the presence of a
non-evaluative supportive friend or stranger than when alone or
just with an interactive control participant. In medical settings,
social support and social activities were associated with less cancer pain (Zaza and Baine, 2002). Women with continuous social
support reported less labor pain (Kennell et al., 1991). Coronarybypass patients with more social support reported less pain and
took less pain medication (Kulik and Mahler, 1989). Given these
ndings, personality theorists and practitioners must distinguish
between distress reecting social harm avoidance and physical
harm avoidance likely linked with the SADNESS or FEAR systems
respectively. For extensive discussions of the social-pain construct,
see MacDonald and Jensen-Campbell (2011).
4.3.3. Evidence summary
The above studies link primary emotions to limbic brain activity.
Furthermore, individuals whose brain systems have higher medial
cortical resting-state levels of activity (see Northoff et al., 2011)
are more reactive to neutral stimuli and respond differently to
events. It is our position that individual differences in such higher
affective as well as lower primary-process aversive affective brain
systems (RAGE, FEAR, and SADNESS) along with the positive affect
systems of PLAY, CARING, and SEEKING are foundational for personality expression as well as the emergence of mental anguish and
pathology. Individuals with different levels of responsiveness in
these primary brain systems not only react differently to the same
stimuli, they will experience these stimuli differently and develop
different conditioned response tendencies and ongoing personal
preferences.
5. Clinical assessments from affective neuroscience
trajectories
The FFM is a factor analytic approach to personality, which has
been derived from studying normal populations. However, psychometric approaches to psychopathology (Livesley, 1986, 1987;
Clark, 1990) have demonstrated that personality disorder (PD)

K.L. Davis, J. Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 19461958

diagnosis can be conceptualized in terms of the FFM (Clark and


Livesley, 2002). Widiger and Costa (2002) reviewed 56 studies that
related the FFM to PD symptomology. Of these studies, 30 used clinical populations. The authors concluded that much of this research
indicates strong support for understanding PD symptomatology as
maladaptive variants of the personality traits included with the
FFM (Widiger and Costa, 2002, p. 80), which suggests that the FFM
does represent a meaningful continuum of behavior extending into
the realm of clinical pathology.
The NEO PI-Revised (Costa and McCrae, 1992) includes six facets
for each of the FFM dimensions. Some feel that facet level analyses
are required for optimal discrimination between PDs (Axelrod et al.,
1997; De Fruyt et al., 2006; Dyce and OConnor, 1998). However, a
challenge for the FFM may be identifying facets that are congruent with neuroscience research rather than relying on an arbitrary
number of rationally dened facets, some of which may be best
dened as factor blends, e.g. Hostility. We may be at a point in
which software capabilities now exceed both the quality of data
and the scope of conceptualization (Wiggins and Pincus, 2002, p.
109) and in which it may be more productive to use functional neuroscience rather than factor analysis to inform and identify new
trait dimensions. For example empathy (Chen et al., 2009) and
jealousy (Panksepp, 2010), and the other socially constructed
emotions, could emerge as newly dened brain emotions in animals that can be used to guide higher-order affective personality
scale construction. In our estimation, it is most important to be very
clear about the primary-process issues that undergird personality
development.

6. Affective neuroscience and therapeutic effectiveness


Affective Neuroscience also has important implications regarding therapy for affective imbalances (see Coenen et al., 2011).
Panksepp (2009) has emphasized the critical nature of the CARE
system for effective therapy and has even considered how oxytocin
administration might be used to enhance a supportive nurturing therapist demeanor. Shedler (2010) has summarized data on
the long-term effectiveness of psychodynamic therapy and likewise emphasized the importance of a nurturing working alliance
between the patient and the therapist. He also discussed research
(see Castonguay et al., 1996) that linked poor therapeutic outcomes to rigid insensitive implementation of therapies. In a second
point, Shedler also emphasized the importance of labeled experience structured so the client gains awareness of previously
implicit feelings and meaning (Shedler, 2010, p. 104; quoted
from Castonguay et al., 1996, p. 499). This suggests that a better
understanding of primary-process affective feelings at a thoughtful,
tertiary-process awareness level may be therapeutically benecial.
In support of this approach, Lieberman showed that affect labeling
decreased amygdala response that was inversely correlated with
the right ventrolateral prefrontal cortex activity, which has been
previously associated with emotional regulation (Lieberman et al.,
2007), and that those more skilled in mindfulness enhanced the
effect (Creswell et al., 2007). These two themes have also been
shown to be important for the success of cognitive behavior therapy (CBT), a nding that was replicated by Hayes et al. (1996). A
third theme, which dealt with CBTs emphasis on cognitive distortions, was not supported in these studies. In fact the therapeutic
aim of cognitive change predicted poorer therapeutic outcomes.
In a review of evidence-based explanations for psychotherapeutic interventions, Kazdin (2007) concluded whatever may be the
basis of changes with CBT, it does not seem to be the cognitions
as originally proposed. (p. 8). Thus, effective therapeutic interventions may typically impact the affective dimensions and levels of
the BrainMind more than cognitive, information-processing ones.

1955

In addition to the importance of the CARE system being displayed by therapists, the PLAY system could have powerful and
underutilized therapeutic effects as well (Panksepp, 2009). Activating the PLAY system requires the bodily vigor, spontaneity, and
creativity of real rough-and-tumble PLAY, which may be most
easily applicable to children. There is good reason to believe that
one of the main functions of the PLAY system is the epigenetic construction of the social brain (Panksepp, 2007b, 2008). Play deprived
human children, just like other mammalian young, develop heightened motivations to play. In humans, this can result in receiving
psychostimulant drugs, which are effective inhibitors of physical
play urges. Animal models conrm that ADHD impulsivity can be
reduced with rough-and-tumble play during early development
(Panksepp et al., 2003). Ample physical play time for children may
be one of the best ways to protect them against ADHD as well as
depression.
While adult brains are not as plastic as those of children, playfulness can be a valuable resource for redirecting adults onto a
more adaptive life track. Robust physical activity by itself may be as
effective an antidepressant as the medications that dampen emotionality (see Watt and Panksepp, 2009). Play urges in adults may
be reenergized by physical activities like sports or dance accompanied by music that stimulates the rhythmic motor impulses of the
body (Panksepp and Trevarthen, 2009).
7. Summary of affective neuroscience perspectives on the
foundations of personality
Our scientic understanding of personality began with Darwin
and McDougal with their focus on emotion and instinct and the
recognition that the differences between humans and their nonhuman ancestors were of degree and not of kind.
In one sense, affective neuroscience helps reconnect personality theory back to our evolutionary roots by showing how closely
our personalities are linked to the ancestral affective forces we
share with other mammals and by relating psychopathology to
disturbances in the primary emotional subcortical brain systems.
Effective treatment of behavioral disorders, and long-term therapeutic change, lies in the sensitive exploration of troubling feelings
and reactive patterns and perhaps not as much in changing distorted cognitions. Consequently, recognizing that our emotional
instincts are fundamentally action systems suggests the importance of incorporating bodily activity into therapy. If a person
is frozen in an emotional state, getting them to move around
and physically interact with the therapists may help promote a
shift in affective balance. After all we are dynamically embodied creatures, whose emotionality is closely linked to motor
action systems. In adult psychotherapy, the PLAY system may
have considerable untapped potential for helping patients reintegrate troublesome emotional experiences towards more adaptive
and emotionally comfortable affective trajectories. For many
people, bringing the body and emotional actions on to the therapeutic eld, may open up many opportunities for change that
would not exist as long as therapy consists largely of talking
heads.
An exploration of cross-species, primary-process emotional
systems of the brain suggests that the FFM itself needs further clarication. Combining the key affective dimensions of FEAR, ANGER,
and SADNESS into a general Neuroticism factor detracts from their
unique contributions to personality expression and mental disorder. Furthermore, ANGER and CARING need to remain distinct
dimensions rather than fusing them onto opposite poles of a scientically confusing Agreeableness factor. One advantage of distinct
emotional dimensions is their underlying unipolar nature, interactive though they are with other emotions, and having separate
dimensions for the ANGER and CARE systems might contribute

1956

K.L. Davis, J. Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 19461958

to clearer primary-process personality thinking. Above all, it may


be helpful to have more precise tools to systematically assess the
location of experimental subjects and therapeutic clients in affective space. And that was our original motivation for developing
the ANPS. We hope it will be useful for many other research and
practical endeavors.
Note: Parkinsons disease has been linked with distinctive personality differences that include an increased risk for depression.
Remy et al. (2005) used PET analysis to investigate the brain correlates of depression in Parkinsons patients and matched controls
and found that depressed Parkinsons subjects had lower activity in
the locus coeruleus, anterior cingulate cortex, thalamus, amygdala,
and ventral striatumall very ancient brain areas. They hypothesized that depression and anxiety in Parkinsons patients were
associated with loss of dopamine and noradrenaline activity in
the limbic system. Damholdt et al. (2011) collected FFM data in
a study of Parkinsons disease and found that Parkinsons patients
with depression displayed lower Extraversion and higher Neuroticism scores. It would have been interesting to have supplemented
the Neuroticism measure with measures of SADNESS, FEAR, and
ANGER to assess likely links to discrete brain systems. Perhaps linking human personality to our ancestral emotional urges will bring
further clarity to the understanding of human personality.
Acknowledgements
We appreciate all the investigators who have taken an interest in the Affective Neuroscience Personality Scales. This work was
supported by a grant to J.P. from the Hope for Depression Research
Foundation.
Appendix A. Scoring the ANPS 2.4, copyright 2004 version
The ANPS items are arranged in 14 blocks using the following
item sequence: SEEK, FEAR, CARE, ANGER, PLAY, SADNESS, Spirituality (with only 12 items), followed by a ller research question.
The items in the even blocks are reverse scored. The easiest way
to hand score the ANPS 2.4 (and compare scores to the original
norms published in the 2003 paper) is to use a 3 for the highest
responses and a 0 for the lowest responses. This procedure allows
for the possibility of a low score of zero on each scale. In effect, the
items are being scored on a scale running from 0 to 3 rather than 1
to 4.
For example, each of the positively worded SEEK scale items
(numbers 1, 17, 33, 49, 65, 81, and 97) would be scored as
follows: Strongly Agree = 3, Agree = 2, Disagree = 1, and Strongly
Disagree = 0. Correspondingly, each of the negatively worded SEEK
scale items in the even blocks (numbers 9, 25, 41, 57, 73, 89,
and 105) would be reverse scored as follows: Strongly Agree = 0,
Agree = 1, Disagree = 2, and Strongly Disagree = 3.
There are 16 ller items in the ANPS 2.4: 7 were designed as
Dominance items (+ans8 +ans40 ans56 ans72 +ans88 +ans103
ans111), 6 were designed as social desirability or unlikely
virtue items, which can cautiously be used as an indication of
deceptive responding (ans16 +ans32 ans48 +ans64 ans80
+ans96), and 3 were written to measure social anxiety (+ans24
+ans104 +ans112).
Alternatively, the ANPS 2.4 was originally set up for scoring with
a scanner that by default numbered the bubbles from left to right
1, 2, 3, and 4. The columns were correspondingly labeled Strongly
Agree, Agree, Disagree, and Strongly Disagree. Thus, if a person
responded to an item with Strongly Agree, the scanner gave them
value of 1, and an Agree response was given a value of 2. Likewise,
if the person responded with Disagree, the scanner gave them a
value of 3, and a Strongly Disagree response was given a value of 4.

Using this 4-point scale with 1 for Strongly Agree and 4 for
Strongly Disagree (but computing scores equivalent to hand scoring above with a base score of 0) the following formulas were used
for computer scoring the 6 primary ANPS scales plus Spirituality:
SEEK score = (+21 ans1 ans17 ans33 ans49 ans65 ans81
ans97 +ans9 +ans25 +ans41 +ans57 +ans73 +ans89 +ans105).
FEAR score = (+21 ans2 ans18 ans34 ans50 ans66 ans82
ans98 +ans10 +ans26 +ans42 +ans58 +ans74 +ans90 +ans106).
CARE score = (+21 ans3 ans19 ans35 ans51 ans67 ans83
ans99 +ans11 +ans27 +ans43 +ans59 +ans75 +ans91 +ans107).
ANGER score = (+21 ans4 ans20 ans36 ans52 ans68
ans84 ans100 +ans12 +ans28 +ans44 +ans60 +ans76 +ans92
+ans108).
PLAY score = (+21 ans5 ans21 ans37 ans53 ans69 ans85
ans101+ans13 +ans29 +ans45 +ans61 +ans77 +ans93 +ans109).
SADNESS score = (+21 ans6 ans22 ans38 ans54 ans70
ans86 ans102 +ans14 +ans30 +ans46 +ans62 +ans78 +ans94
+ans110).
Spirituality score = (+18 ans7 ans23 ans39 ans55 ans71
ans87 +ans15 +ans31 +ans47 +ans63 +ans79 +ans95).
Algebraically, the hand scoring and computer scoring procedures are identical.
Copyright 2004, Kenneth L. Davis, Ph.D., Jaak Panksepp, Ph.D.,
Pegasus International, Inc. All rights reserved.
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