Neuroscience and Biobehavioral Reviews 35 (2011) 1946–1958

Contents lists available at ScienceDirect

Neuroscience and Biobehavioral Reviews

j ournal h o m ep ag e: www. e l s ev ie r . com / l o ca te / n eu bi o r ev

Review

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

Neuroscience Personality Scales
Kenneth L. Davisa, ∗, 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

ar t ic l e in f o abst r ac 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: 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
 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
labeled as ‘irritability’ and ‘anger’.
Neuroanatomical, neurochemical, pharmacological, and physi
ological brain research have helped to more precisely define 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 person
ality 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 com
plexities of primary process affective space shared by all mammals.
If so, that could also open up a common terminology for study per
sonality across species. It is a major supposition of this paper that
these basic emotions are fundamental powers of the human Brain
Mind, homologous with feelings experienced by other mammals,
and hence potentially of critical importance for understanding the
foundations of personality (we intentionally conflate ‘brain’ and
‘mind’, and double capitalized BrainMind to reflect our monistic
ontology).
There is strong evidence for these seven well defined emo
tion 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 pro
cesses. (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 behav
ioral 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 specific
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 con
trol 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 sub
cortical nature of these emotion systems. Also, each instinctual
emotional system engenders affective valence since animals termi
nate 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 SEEK
ING system is foundational for aspects of all of the other primal
emotions (e.g., during FEAR intensification, the shift from freezing
to flight may be due to recruitment of SEEKING related dopamin
ergic 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. Specifically, 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 assess
ment of human temperaments/personalities.
But this is not the first attempt to bridge basic neuroscience
and human personality evaluation. The two individuals who have
attempted to construct neurobiologically based personality assess
ment instruments in the past are Cloninger (1986) who based
his thinking on functional animal work on some of the first well
characterized neurochemical systems of all mammalian brains –
dopamine, norepinephrine, and serotonin (see below). In con
trast, Carver and White (1994) developed the Behavioral Activation
and Behavioral Inhibition Scales on the basis of Jeffrey Gray’s
(1981) early brain research recognizing the existence of behavioral
approach/arousal and withdrawal/inhibition (aka, reward and pun
ishment) 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 rele
vant for defining temperamental/personality dimensions, with the
popular Cloninger assessment tool following a brief introduction of
the five 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 Fac
tor 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 clin
ical 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 five
personality dimensions are usually labeled Extraversion or Sur
gency, Agreeableness, Conscientiousness, Emotional Stability, and
Intellect or Openness to Experience (Hofstee et al., 1992). These
five 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 first four dimensions have been more robust with
Openness to Experience typically accounting for the least variance
of the five factors (Goldberg, 1990). A comprehensive presenta
tion of the five factor “descriptive” model was provided by Hofstee
et al. (1992) and consisted of an analysis of over 500 adjectives
placed on five 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. Cloninger’s biologically based personality theory
Cloninger was the first to attempt an integration of personality
and brain systems with his biogenic amine based “biosocial” theory
of personality (Cloninger, 1987) and the Tridimensional Personal
ity 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 reflect brain serotonin
activity, the brain’s presumed punishment system (which is highly
debatable, see Panksepp, 1986, 1998), and the resulting sensitiv
ity to threatening situations, and (3) Reward Dependence featuring
behavioral maintenance and sensitivity to reinforcement contin
gencies 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) 1946–1958
Our affective neuroscience perspective would align the Novelty
Seeking dopaminergic brain incentive dimension with the SEEK
ING system. The SEEKING system not only promotes exploration,
investigation and foraging but probably ‘energizes’ all basic emo
tional 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 specific contributors to very specific
types of affects the way Cloninger proposed, even though they
figure heavily in the arousal dimensions of all personality control
systems. Norepinephrine promotes attention by amplifying signal
to 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, Coop
erativeness, 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 Tem
perament and Character Inventory (TCI; Cloninger, 1999). Although
Cloninger’s earlier TPQ factors have been related to many genetic
variables, many of Cloninger’s later theoretical claims have not
been supported. In other words, heritabilities did not differenti
ate 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 tem
perament 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 five factors (Farmer and Goldberg,
2008a,b). Others also reported considerable convergence between
Cloninger’s 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 emo
tional system foundations of personality. The scales were rationally
defined 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 definitional
circularity since comparative research has demonstrated how each
primary dimension can be manipulated in humans as indepen
dent 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 identified by cross species ESB based affective
neuroscience research. We excluded LUST, which seems less rel
evant 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 contami
nate 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 defined as anticipating new positive experiences including
being curious, liking to strive for solutions to problems, and gener
1949
ally liking to explore. CARING centered on nurturing tendencies
including liking to care for others, being drawn to young chil
dren and pets, and feeling softhearted towards animals and people
in need. The FEAR scale incorporated experiencing anxiety, wor
rying, difficulty making decisions, ruminating, feeling tense, and
losing sleep. ANGER included feeling hotheaded, being easily irri
tated 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 relation
ships 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 scien
tific 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 first 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 first study using these scales, reliabilities for the
ANPS scales, computed as Cronbach’s alpha, were reported rang
ing 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 Extraver
sion (r = .46), (3) CARE with Agreeableness (r = .50), (4) FEAR with
Emotional Stability (r = −.75), (5) SADNESS with Emotional Stabil
ity (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 five factor personality
model.
When factor analyzing ANPS scales with FFM scales, we reported
a five 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 specific 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 first 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 gen
erally replicates Gray’s (1970) Behavioral Inhibition and Approach
Systems. We feel this is a striking demonstration of the multifac
torial nature of any general positive and negative valence type of
measurement tool. In this vein, it is also important to note Goldberg
and colleague’s 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 substan
tive dimensions of personality.”
 K.L. Davis, J. Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 1946–1958 1951

Pahlavan et al. (2008) confirmed the psychometric properties
of the ANPS in a French population including scale reliabili
ties, factor structure, and correlations with the FFM. Berthoz
and colleagues (personal communication) replicated the Pahlavan
et al. (2008) findings on a large sample (n = 830) and also
performed an item level confirmatory factor analysis obtain
ing an acceptable fit modeling 6 factors (standardized root
mean square residual = .06, root mean square error of approxi
mation = .039, and the comparative fit index = .78). Savitz et al.
(2008c) obtained significant evidence of heritability for four of
the ANPS scales: CARE, SEEK, FEAR, and Spirituality. Through per
sonal correspondence, we know the scale has been translated into
a dozen languages, and has already proved its utility in eval
uating 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.
47321
mdanps24
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 bl ocks 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.
1950 K.L. Davis, J. Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 1946–1958

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.
47321
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) sug
gested 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 Conscientious
ness. 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 Canli’s (2008) view of
impulsiveness as a “top down” lack of behavioral inhibition
involving the inferior frontal cortex and subthalamic nucleus fits
well with King’s (2007) description of chimpanzee Conscientious
ness – characterized by predictable, not impulsive or erratic,
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
PLAY .46* .29*
SEEK .13 −.01
CARE .25 .50*
FEAR −.19 −.17
ANGER −.04 −.48*
SADNESS −.21 −.13
Spirituality .15 .26*
Student sample: n = 170 (50 males, 121 females).The Five Factor Model was measured using Goldberg (1992) adjectives, selecting 14 markers for e ach Big Five dimension
(data adapted from Davis et al., 2003).
*
p < .001, two tailed.
These results provide encouraging evidence that the FFM
reflects primary subcortical affects. Although ANPS items attempt
to address primary affects directly, since all self report assessments
must include cognitive reflection, we interpret the ANPS scales as
tertiary (thought mediated) approximations of the influence of the
various primary emotional systems in people’s lives. However, it
is our working hypothesis that the subcortical primary processes
neural systems, where the foundations of emotions reside, can gen
erate 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. Agreeable
ness 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,
human’s closest evolutionary relative. These findings support the
Darwinian concept of continuity of species that “the difference in
the mind between man and the higher animals, great as it is, cer
tainly 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 nega
tive 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 dis
tinct 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 com
bine 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, nur
turing feelings on the positive pole and hostile, demanding feelings
on the negative pole. Our view is that while the FFM is an impor
tant theory free, empirical step forward in parsing personality, it
Conscientiousness Emotional stability Openness to experience
.00 .12 .13
−.01 .01 .47*
.12 −.07 .06
−.24 −.75* −.05
−.30* −.65* −.08
−.30* −.68* −.00
.14 .09 .17
reflects a tertiary cognitive type reconfiguration 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, study
ing families with a member having bipolar disorder (BPD), Savitz
et al. (2008a) found that the ANPS SADNESS scale was significantly
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
eralized 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 diag
nosed 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 depres
sion (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 significant differences across
groups.
Savitz et al. (2008b) also studied the distribution of hypo
manic, cyclothymic, and hostile traits in BPD probands and their
families. After controlling for age, gender, depression, and mania,
BPD II patients had significantly 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 significantly correlated with higher ANGER scores and
lower SEEK and PLAY scores while the Altman Self Rating Mania
Scale significantly correlated with higher ANGER, SEEK, and PLAY
scores.
The construct validity of the ANPS ANGER scale was also sup
ported 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 dopaminer
gic 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 significantly higher ANPS ANGER scores,
with CC genotypes having significantly lower ANGER scores. MRI
data on a subsample of subjects revealed that ANGER scores cor
related negatively with gray matter volume in the left amygdala.
Reuter’s 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 significantly lower scores on
the personality dimensions FEAR and SADNESS of the ANPS, as well
as on the overall super factor of Negative Emotionality, than carri
ers of the other serotonin transporter and oxytocin gene variants
(Montag et al., 2011).
 K.L. Davis, J. Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 1946–1958
Reuter et al. (2005) also demonstrated a link between the ANPS
SEEK scale and creativity. Using figural, verbal, and numeric cre
ativity tests, subjects with high SEEK scores had higher numeric
creativity scores and were significantly superior on figural and ver
bal creativity. SEEK scores also explained more than 15% of the
variance of total creativity. Using intelligence tests as a covari
ate 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 person
ality traits in the FFM, affective neuroscience (Panksepp, 1998)
suggests an enhanced neuroscientifically premised personality
research model. By providing a putative physiological basis for
FFM traits, affective neuroscience helps five factor theory escape
circular reasoning by suggesting brain mechanisms that can be
manipulated to influence the expression of personality. However,
even more important may be helping to advance personality theory
from trait and situationist approaches to interactionist and ontoge
netic personality models (see Ballantyne, 1995 for a more detailed
discussion of personality theory levels).
One ontogenetic model is Scarr and McCartney’s (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 influence the kind of care infants receive from their parents.
While the passive effect is hypothesized to decrease with onto
genetic development, the evoked effect continues throughout an
individual’s 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 find attractive, which accounts for adopted
siblings not resembling each other’s personalities by late adoles
cence (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 defining the attractiveness of environments. Plea
surable experiences result in finding similar situations attractive
in the future with traumatic or painful experiences having the
opposite effect. One example prediction would be that individu
als 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 find 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 rough
and 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 off
spring (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 shorter
lived species, one could also test Scarr and McCartney’s (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 primi
tive 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 emo
tional/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 reflecting basic
conditioning learning. In humans, and perhaps some other corti
cally well endowed animals, thoughts about how external events
relate to internal events yield even higher tertiary forms of con
sciousness consisting of thoughts about thoughts, awareness of
awareness, and the linguistic/symbolic transformation of experi
ence at which humans excel (for overview, see Vandekerckhove
and Panksepp, 2009 and in this issue). However, it is the primary
process 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 reflect
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 defined 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 MacLean’s (1990)
sub neocortical “limbic” and “reptilian” areas of the central ner
vous system. It is our position that in these evolutionally older
parts of the brain, much more ancient than “neomammalian”
cortex, one finds the most important evolutionary “roots” of per
sonality. 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 emo
tional 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 “sec
ondary” conditioning and “tertiary” thoughts and self reflections,
but the evolutionary origin and foundational power of these dis
crete, persistent systems is pre human.
4.3. State vs. channel functions
The influence of these basic emotion systems is pervasive, affect
ing not only our actions and reactions but also our perceptions,
thoughts, and memories. The mechanism mediating these old brain
influences may lie in the distinction between “channel” and “state”
brain functions (Mesulam, 2000). For “state” modulation, each cor
tical area receives inputs arising from limbic neurons, which can
modulate activity in the entire cerebral cortex. These influences
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
1954 K.L. Davis, J. Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 1946–1958
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 interpre
tation of experience rather than its content” (Mesulam, 2000, pp.
78–79).
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 tem
poral 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 sub
cortical “states” biased perceptions and generated greater reactive
tendencies. Such tendencies would likely result in observed indi
vidual differences in the many behavioral dimensions of individual
lives.
Canli and Lesch (2007) have analyzed the influence of the sero
tonin transporter gene (5 HTT) short variant on fMRI measured
activity of amygdala and related brain areas and found that well
adjusted 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. geo
metric shapes. Analysis of emotionally negative or neutral stimuli
vs. a fixation rest condition showed decreased amygdala activation
to neutral relative to fixation stimuli, which was “interpreted as
indicating elevated amygdala activation during the processing of
emotionally undefined 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 sup
ported with a perfusion imaging study that measured absolute
amygdala blood flow at rest. Higher baseline amygdala activity lev
els 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 specific role in modulating any of the
primary emotional responses but functions as a general modula
tor 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 difficult to inte
grate results into primary brain emotion systems, since affective
neuroscience would predict that, while the FEAR and ANGER sys
tems are closely related anatomically in lower brain regions, they
remain distinctly separate systems, and my have distinct higher
brain regions of influence where raw affects are blended with cog
nitive 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) activ
ity 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 partic
ipants. During an initial control scan, subjects just watched other
participants play the virtual ball tossing game, ostensibly because
“technical difficulties” prevented them from participating. In the
second scan, subjects were linked in with “virtual” participants sup
posedly in other scanners and “included” in the virtual ball tossing
game. To create a sense of social exclusion, during the final scan
subjects were socially excluded when the other players (actually a
computer program) stopped tossing the ball to the subject. Subjects
showed significantly more dACC activation during the social exclu
sion 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 pre
frontal 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). Con
versely, 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 can
cer pain (Zaza and Baine, 2002). Women with continuous social
support reported less labor pain (Kennell et al., 1991). Coronary
bypass patients with more social support reported less pain and
took less pain medication (Kulik and Mahler, 1989). Given these
findings, personality theorists and practitioners must distinguish
between distress reflecting 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 per
sonality 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, psy
chometric 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) 1946–1958
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 clin
ical 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 O’Connor, 1998). However, a
challenge for the FFM may be identifying facets that are congru
ent with neuroscience research rather than relying on an arbitrary
number of rationally defined facets, some of which may be best
defined 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 neu
roscience 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 defined brain emotions in ani
mals 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 regard
ing 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 nurtur
ing therapist demeanor. Shedler (2010) has summarized data on
the long term effectiveness of psychodynamic therapy and like
wise 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 out
comes to rigid insensitive implementation of therapies. In a second
point, Shedler also emphasized the importance of “labeled expe
rience” 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 beneficial.
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 ther
apy (CBT), a finding that was replicated by Hayes et al. (1996). A
third theme, which dealt with CBT’s emphasis on cognitive distor
tions, 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 psychotherapeu
tic 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 interven
tions 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 dis
played by therapists, the PLAY system could have powerful and
underutilized therapeutic effects as well (Panksepp, 2009). Acti
vating 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 con
struction of the social brain (Panksepp, 2007b, 2008). Play deprived
human children, just like other mammalian young, develop height
ened motivations to play. In humans, this can result in receiving
psychostimulant drugs, which are effective inhibitors of physical
play urges. Animal models confirm 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, play
fulness 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 emo
tionality (see Watt and Panksepp, 2009). Play urges in adults may
be reenergized by physical activities like sports or dance accompa
nied 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 scientific 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 non
human ancestors were of “degree” and not of “kind.”
In one sense, affective neuroscience helps reconnect personal
ity 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 thera
peutic change, lies in the sensitive exploration of troubling feelings
and reactive patterns and perhaps not as much in changing dis
torted cognitions. Consequently, recognizing that our emotional
instincts are fundamentally action systems suggests the impor
tance 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 ‘embod
ied’ creatures, whose emotionality is closely linked to motor
action systems. In adult psychotherapy, the PLAY system may
have considerable untapped potential for helping patients reinte
grate troublesome emotional experiences towards more adaptive
and emotionally comfortable affective trajectories. For many
people, bringing the body and emotional actions on to the ther
apeutic field, 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 clar
ification. 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 dis
order. Furthermore, ANGER and CARING need to remain distinct
dimensions rather than fusing them onto opposite poles of a scien
tifically confusing Agreeableness factor. One advantage of distinct
emotional dimensions is their underlying unipolar nature, inter
active 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) 1946–1958
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 affec
tive 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: Parkinson’s disease has been linked with distinctive per
sonality differences that include an increased risk for depression.
Remy et al. (2005) used PET analysis to investigate the brain cor
relates of depression in Parkinson’s patients and matched controls
and found that depressed Parkinson’s subjects had lower activity in
the locus coeruleus, anterior cingulate cortex, thalamus, amygdala,
and ventral striatum–all very ancient brain areas. They hypoth
esized that depression and anxiety in Parkinson’s 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 Parkinson’s disease and found that Parkinson’s patients
with depression displayed lower Extraversion and higher Neuroti
cism 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 link
ing 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 inter
est 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, Spiri
tuality (with only 12 items), followed by a filler 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 filler 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 scor
ing 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” pro
cedures are identical.
Copyright © 2004, Kenneth L. Davis, Ph.D., Jaak Panksepp, Ph.D.,
Pegasus International, Inc. All rights reserved.
References
Allport, G.W., Odbert, H.S., 1936. Trait names, a psycho lexical study. Psychol.
Monogr. 47 (1), 171.
Altman, E., Hedecker, D., Peterson, J.L., Davis, J.M., 1997. The Altman Self Rating
Mania Scale. Biol. Psychiatry 42, 948–955.
Ashton, M.C., Lee, K., Goldberg, L.R., de Vries, R.E., 2009. Higher order factors of
personality: do they exist? Rev. Pers. Soc. Psych. 13, 79–91.
Axelrod, S.R., Widiger, T.A., Trull, T.J., Corbitt, E.M., 1997. Relations of Five Factor
Model antagonism facets with personality disorder symptomatology. J. Pers.
Assessment 69 (2), 297–313.
Ball, S.A., Tennen, H., Kranzler, H.R., 1999. Factor replicability and validity of the tem
perament and character inventory in substance dependent patients. Psychol.
Assessment 11 (4), 514–524.
Ballantyne, P.F., 1995. From initial abstractions to a concrete concept of personality.
In: Lubek, I. (Ed.), Recent Trends in Theoretical Psychology, vol. 4. Springer, New
York, pp. 151–157.
Bechara, A., Damasio, H., Damasio, A.R., Lee, G.P., 1999. Different contributions of
the human amygdala and ventromedial prefrontal cortex to decision making. J.
Neurosci. 19, 5473–5481.
Beck, A.T., Steer, R.A., 1993. Manual for the Beck Depression Inventory. The Psycho
logical Corporation, San Antonio.
Block, J., 1995. A contrarian view of the five factor approach to personality descrip
tion. Psychol. Bull. 117 (2), 187–215.
Brown, J.L., Sheffield, D., Leary, M.R., Robinson, M.E., 2003. Social support and exper
imental pain. Psychosom. Med. 65, 276–283.
Canli, T., Lesch, K.P., 2007. Long story short: the serotonin transporter in emotion
regulation and social cognition. Nat. Neurosci. 10 (9), 1103–1109.
Canli, T., Maolin, Q., Omura, K., Congdon, E., Haas, B.W., Amin, Z., Herrmann, M.J.,
Constable, R.T., Lesch, K.P., 2006. Neural correlates of epigenesis. Proc. Nat. Acad.
Sci. U.S.A. 103 (43), p16033–16038.
Carver, C.S., White, T.L., 1994. Behavioral inhibition, behavioral activation, and affec
tive responses to impending reward and punishment: the BIS/BAS scales. J. Pers.
Soc. Psych. 67 (2), 319–333.
Castonguay, L.G., Goldfried, M.R., Wiser, S.L., Raue, P.L., Hayes, A.M., 1996. Predicting
the effect of cognitive therapy for depression: a study of unique and common
factors. J. Cons. Clin. Psych. 64, 497–504.
Cattell, R.B., 1947. Confirmation and clarification of primary personality factors.
Psychometrika 12 (3), 197–220.
Chen, Q., Panksepp, J.B., Lahvis, G.P., 2009. Empathy is moderated by genetic back
ground in mice. PlosOne 4 (2), e4387, doi:10.1371/journal.pone.0004 387.
Clark, L.A., 1990. Towards a consensual set of symptom clusters for assessment of
personality disorder. In: Butcher, J., Spielberger, C.D. (Eds.), Advances in Person
ality Assessment, vol. 8, pp. 243–266.
Clark, L.A., Livesley, W.J., 2002. Two approaches to identifying the dimensions of per
sonality disorder: convergence on the five factor model. In: Costa, P.T., Widiger,
T.A. (Eds.), Personality Disorders and the Five Factor Model of Personality. , sec
ond ed. American Psychological Association, Washington, DC, USA, pp. 161–176.
Cloninger, C.R., 1986. A unified biosocial theory of personality and its role in the
development of anxiety states. Psychiatric Dev. 3, 167–226.
 K.L. Davis, J. Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 1946–1958
Cloninger, C.R., 1987. A systematic method for clinical description and classification
of personality variants. Arch. Gen. Psychiatry 44, 573–588.
Cloninger, C.R., 1999. The Temperament and Character Inventory—Revised. Wash
ington University, Center for Psychobiology of Personality, St. Louis, MO
(available from C.R. Cloninger, Washington University School of Medicine,
Department of Psychiatry, P.O. Box 8134, St. Louis, MO 63110).
Cloninger, C.R., 2004. Feeling Good: The Science of Well Being. Oxford University
Press, Inc., New York, NY.
Coenen, V.A., Schlaepfer, T.E., Maedler, B., Panksepp, J., 2011. Cross species affec
tive functions of the medial forebrain bundle Implications for the treatment of
affective pain and depression in humans. Neurosci. Biobehav. Rev. 2010 Dec 22
[Epub ahead of print].
Congdon, E., Canli, T., 2008. A neurogenetic approach to implusivity. J. Pers. 76 (6),
1447–1483.
Costa, P.T., McCrae, R.R., 1992. Revised NEO Personality Inventory (NEO PI R) and
NEO Five Factor Inventory (NEO FFI) Professional Manual. Psychological Assess
ment Resources, Odessa, FL.
Costa, P.T., Widiger, T.A., 2002. Introduction: Personality Disorders and the Five
Factor Model of Personality. Personality Disorders and the Five Factor Model
of Personality, second ed. American Psychological Association, Washington, DC,
USA, pp. 3 14.
Creswell, J.D., Way, B.M., Eisenberger, N.I., Lieberman, M.D., 2007. Neural corre
lates of dispositional mindfulness during affect labeling. Psychosom. Med. 69
(6), 560–565.
Damholdt, M.F., Ostergaard, K., Borghammer, P., Larsen, L., 2011. The parkinsonian
personality and concomitant depression. J. Neuropersonality Clin. Neurosci. 23,
48–55.
Darwin, C., 1872/1967. The Expression of the Emotions in Man and Animals. Oxford
University Press, Oxford.
Davis, K., Panksepp, J., Normansell, L., 2003. The affective neuroscience personality
scales: normative data and implications. Neuropsychoanalysis 5 (1), 57–69.
De Fruyt, F., De Clercq, B.J., Van De Wiele, L., Van Heeringen, K., 2006. The validity
of cloninger’s psychobiological model versus the five factor model to predict
DSM IV personality disorders in a heterogeneous psychiatric sample: domain
facet and residualized facet descriptions. J. Pers. 74, 479–510.
De Fruyt, F., Van De Wiele, L., Van Heeringen, C., 2000. Cloninger’s psychobiological
model of temperament and character and the five factor model of personality.
Pers. Ind. Diff. 29, 441–452.
Deyo, R.A., Panksepp, J., Abbott, B., 1990. Perinatal decortication impairs perfor
mance on an 8 arm radial maze task. Physiol. Behav. 48 (1), 55–60.
Digman, J.M., 1990. Personality structure: emergence of the five factor model. Ann.
Rev. Psych. 41, 417–440.
Dyce, J.A., O’Connor, B.P., 1998. Personality disorders and the five factor model: a
test of facet level predictions. J. Pers. Disord. 12 (1), 31–45.
Eisenberger, N.I., Lieberman, M.D., 2005. Why it hurts to be left out: the neurocog
nitive overlap between physical and social pain. In: Williams, K.D., Forgas, J.P.,
von Hippel, W. (Eds.), The Social Outcast: Ostracism, Social Exclusion, Rejection,
and Bullying. Psychology Press, Taylor & Francis Group, LLC, New York, NY, pp.
109–127.
Eisenberger, N., Lieberman, M., Williams, K., 2003. Does rejection hurt? An fMRI
study of social exclusion. Science 302, 290–292.
Evans, L., Akiskal, H.S., Keck Jr., P.E., McElroy, S.L., Sadovnick, A.D., Remick, R.A., Kel
soe, J.R., 2005. Familiarity of temperament in bipolar disorder: support for a
genetic spectrum. J. Affective Disord. 85, 153–168.
Farmer, R.F., Goldberg, L.R., 2008a. A Psychometric Evaluation of the Revised Tem
perament and Character Inventory (TCI R) and the TCI 140. Psychol. Assessment
20 (3), 281–291.
Farmer, R.F., Goldberg, L.R., 2008b. Brain modules, personality layers, planes of
being, spiral structures, and the equally implausible distinction between TCI
R “Temperament” and “Character” Scales: reply to Cloninger (2008). Psychol.
Assessment 20 (3), 300–304.
Francis, D., Diorio, J., Liu, D., Meaney, M.J., 1999. Nongenomic transmission across
generations of maternal behavior and stress responses in the rat. Science 286,
1155–1158.
Gillespie, N.A., Cloninger, C.R., Heath, A.C., Martin, N.G., 2003. The genetic and envi
ronmental relationship between Cloninger’s dimensions of temperament and
character. Pers. Ind. Diff. 35, 1931–1946.
Goldberg, L.R., 1990. An alternative description of personality: the big five factor
structure. J. Pers. Soc. Psych. 59 (6), 1216–1229.
Gosling, S.D., 2008. Personality in non human animals. Soc. Pers. Psych. Compass 2,
985–1001.
Gosling, S.D., John, O.P., 1999. Personality dimension in non human animals: a cross
species review. Curr. Direct. Psychol. Sci. 8 (3), 69–75.
Gray, J.A., 1970. The psychophysiological basis of introversion extraversion. Behav.
Res. Therapy 8, 249–266.
Gray, J.A., 1981. A critique of Eysenck’s theory of personality. In: Eysenck, H.J. (Ed.),
A Model for Personality. Springer Verlag, Berlin, pp. 246–276.
Hayes, A.M., Castonguay, L.M., Goldfried, M.R., 1996. Effectiveness of targeting the
vulnerability factors of depression in cognitive therapy. J. Consulting Clin. Psych.
64 (3), 623–627.
Hess, W.R., 1957. The Functional Organization of the Diencephalons. Grune and
Statton, New York.
Hofstee, W., de Raad, B., Goldberg, L., 1992. Integration of the big five and circumplex
approaches to trait structure. J. Pers. Soc. Psych. 63 (1), 146–163.
Huber, R., Panksepp, J.B., Nathaniel, T., Alcaro, A., Panksepp, J., 2011. Drug sensitive
reward in crayfish: An invertebrate model system for the study of SEEKING,
1957
reward, addiction, and withdrawal. Neurosci. Biobehav. Rev. 2010 Dec 21 [Epub
ahead of print].
Kahneman, D., 2003. A perspective on judgment and choice: mapping bounded
rationality. Am. Psychologist 58 (9), 697–720.
Kazdin, A.E., 2007. Mediators and mechanisms of change in psychotherapy research.
Ann. Rev. Clin. Psych. 3, 1–27.
Kennell, J., Klaus, M., McGrath, S., Robertson, S., Hinkley, C., 1991. Continuous emo
tional support during labor in U.S. hospital: a randomized control trial. J. Am.
Med. Assn. 265, 2197–2201.
King, J.E., 2007. Dimensions of the Ape Mind: adding personality to behavior and cog
nition. In: Washburn, D.A. (Ed.), Primate Perspectives on Behavior and Cognition.
American Psychological Association, Washington, DC, USA, pp. 47–60.
Kulik, J.A., Mahler, H.I.M., 1989. Social support and recovery from surgery. Health
Psych. 8 (2), 221–238.
Levine, F.M., Krass, S.M., Padawer, W.J., 1993. Failure hurts: the effects of stress
due to difficult tasks and failure feedback on pain report. Pain 54, 335–
340.
Lieberman, M.D., Eisenberger, N.I., Crockett, M.J., Tom, S.M., Pfeifer, J.H., Way, B.M.,
2007. Putting feelings into words: affect labeling disrupts amygdala activity in
response to affective stimuli. Psychol. Sci. 18 (5), 421–428.
Livesley, W.J., 1986. Traits and behavioral prototypes of personality disorder. Am. J.
Psychiatry 143, 728–732.
Livesley, W.J., 1987. A systematic approach to the delineation of personality disor
ders. Am. J. Psychiatry 144, 772–777.
MacLean, P.D., 1990. The Triune Brain in Evolution. Plenum Press, New York.
MacDonald, G., Jensen Campbell, L.A., 2011. Social Pain: Neuropsychological and
Health Implications of Loss and Exclusion. American Psychological Association,
Washington, DC.
McDougal, W., 1908/1963. An Introduction to Social Psychology. Morrison & Gibb,
London, Edinborough, Great Britain.
Merker, B., 2007. Consciousness without a cerebral cortex: a challenge for neuro
science and medicine. Behav. Brain Sci. 30, 63–81.
Mesulam, M.M., 2000. Principles of Behavioral and Cognitive Neurology. Oxford
University Press, Oxford.
Montag, C., Fiebach, C.J., Kirsch, P., Reuter, M., 2011. Interaction of 5 HTTLPR and a
variation on the oxytocin receptor gene influences negative emotionality. Biol.
Psychiatry 69, 601–603.
Northoff, G., Wiebking, C., Feinberg, T., Panksepp, J., 2011. The ‘resting state hypothe
sis’ of major depressive disorder A translational subcortical cortical framework
for a system disorder. Neurosci. Biobehav. Rev. 2010 Dec 28 [Epub ahead of
print].
Pahlavan, F., Mouchiroud, C., Zenasni, F., Panksepp, J., 2008. French validation of
the Affective Neuroscience Personality Scales (ANPS). Revue européenne de
psychologie appliquée 58, 155–163.
Panksepp, J., 1971. Aggression elicited by electrical stimulation of the hypothalamus
in albino rats. Physiol. Behav. 6, 311–316.
Panksepp, J., 1982. Toward a general psychobiological theory of emotions. Behav.
Brain Sci. Vol5 (3), 407–467.
Panksepp, J., 1985. Mood changes. In: Vinken, P.J., Bruyn, G.W., Klawans, H.L. (Eds.),
Handbook of Clinical Neurology (Revised Series). Vol. 1 (45): Clinical Neuropsy
chology. Elsevier Science Publishers, Amsterdam, pp. 271–285.
Panksepp, J., 1986. The neurochemistry of behavior. Ann. Rev. Psych. 37, 77–
107.
Panksepp, J., 1998. Affective Neuroscience: The Foundations of Human and Animal
Emotions. Oxford University Press, Oxford.
Panksepp, J., 2005. Affective consciousness: core emotional feelings in animals and
humans. Consciousness Cogn. 14, 30–80.
Panksepp, J., 2007a. Affective consciousness. In: Velmans, M., Schneider, S. (Eds.),
The Blackwell Companion to Consciousness. Blackwell Publishing, Ltd., Malden,
MA, pp. 114–129.
Panksepp, J., 2007b. Neuroevolutionary sources of laughter and social joy: modeling
primal human laughter in laboratory rats. Behav. Brain Res. 182, 231–244.
Panksepp, J., 2008. Play, ADHD, and the construction of the social brain: should the
first class each day be recess? Am. J. Play 1, 55–79.
Panksepp, J., 2009. Brain emotional systems and qualities of mental life. In: Fosha,
D., Siegel, D.J., Solomon, M.F. (Eds.), The Healing Power of Emotion. W.W. Norton
& Company, New York, pp. 1–26.
Panksepp, J., 2010. The evolutionary sources of jealousy: cross species approaches
to fundamental issues. In: Hart, S.L., Legerstee, M. (Eds.), Handbook of Jealousy:
Theory, Research, and Multidisciplinary Approaches. Wiley Blackwell, Chich
ester, West Sussex, UK, pp. 101–120.
Panksepp, J., 2011. The neurobiology of social loss in animals: Some keys to the puz
zle of psychic pain in humans. In: Jensen Campbell, L.A., MacDonald, G. (Eds.),
Social Pain: Neuropsychological and Health Implications of Loss and Exclusion.
American Psychological Association, Washington, DC, pp. 11–51.
Panksepp, J., Biven, L., 2011. The Archaeology of Mind: Neuroevolutionary Origins
of Human Emotions. Norton, New York.
Panksepp, J., Burgdorf, J., Turner, C., Gordon, N., 2003. Modeling ADHD type arousal
with unilateral frontal cortex damage in rats and beneficial effects of play ther
apy. Brain Cogn. 52, 97–105.
Panksepp, J., Nocjar, C., Burgdorf, J., Panksepp, J.B., Huber, R., 2004. The role of emo
tional systems in addiction: a neuroethological perspective. In: Bevins, R.A.,
Bardo, M.T. (Eds.), 50th Nebraska Symposium on Motivation: Motivational Fac
tors in the Etiology of Drug Abuse. Nebraska, Lincoln, pp. 85–126.
Panksepp, J., Normansell, L.A., Cox, J.F., Siviy, S., 1994. Effects of neonatal decortica
tion on the social play of juvenile rats. Physiol. Behav. 56, 429–443.
1958 K.L. Davis, J. Panksepp / Neuroscience and Biobehavioral Reviews 35 (2011) 1946–1958
Panksepp, J., Trevarthen, C., 2009. The neuroscience of emotion in music. In: Malloch,
S., Trevarthen, C. (Eds.), Communicative Musicality. Oxford University Press,
Oxford, UK, pp. 105–146.
Peirson, A.R., Heuchert, J.W., Thomala, L., Berk, M., Plein, H., Cloninger, C.R., 1999.
Relationship between serotonin and the Temperament and Character Inventory.
Psychiatry Res. 89, 29–37.
Remy, P., Doder, M., Lees, A., Turjanski, N., Brooks, D., 2005. Depression in Parkinson’s
disease: loss of dopamine and noradrenaline innervation in the limbic system.
Brain 128, 1314–1322.
Reuter, M., Panksepp, J., Schnabel, N., Kellerhoff, N., Kempel, P., Hennig, J.,
2005. Personality and biological markers of creativity. Eur. J. Pers. 19, 83–
95.
Reuter, M., Weber, B., Fiebach, C.J., Elger, C., Montag, C., 2009. The biological basis
of anger: associations with the gene coding for DARPP 32 (PPP1R1B) and with
amygdala volume. Behav. Brain Res. 202, 179–183.
Ross, M.D., Owen, M.J., Zimmermann, E., 2009. Reconstructing the evolution of
laughter in great apes and humans. Curr. Biol. 19, 1106–1111.
Saucier, G., Goldberg, L.R., 2006. Personnalité, caractère et tempérament: La struc
ture translinguistique des traits./Personality, character and temperament: The
cross language structure of traits. Psychologie Franc¸ aise 51 (3), 265–284.
Savitz, J., van der Merwe, L., Ramesar, R., 2008a. Dysthymic and anxiety related
personality traits in bipolar spectrum illness. J. Affective Disord. 109, 305–311.
Savitz, J., van der Merwe, L., Ramesar, R., 2008b. Hypomanic, cyclothymic and hostile
personality traits in bipolar spectrum illness: a family based study. J. Psychiatric
Res. 42, 920–929.
Savitz, J., van der Merwe, L., Ramesar, R., 2008c. Personality endophenotypes for
bipolar affective disorder: a family based genetic association analysis. Genes
Brain Behav. 7 (8), 869–876.
Scarr, S., McCartney, K., 1983. How people make their own environments; a theory
of genotype, environment effects. Child Dev. 54, 424–435.
Scarr, S., Webber, P.L., Weinberg, R.A., Wittig, M.A., 1981. Personality resemblance
among adolescents and their parents in biologically related and adoptive fami
lies. J. Pers. Soc. Psych. 40 (5), 885–898.
Shedler, J., 2010. The efficacy of psychodynamic psychotherapy. Am. Psychologist
65 (2), 98–109.
Shewmon, D.A., Holmes, G.L., Byrne, P.A., 1999. Consciousness in congenitally decor
ticate children: developmental vegetative state as self fulfilling prophecy. Dev.
Child Neurol. 41, 364–374.
Vandekerckhove, M., Panksepp, J., 2009. The flow of anoetic to noetic and autonoetic
consciousness: a vision of unknowing (anoetic) and knowing (noetic) conscious
ness in the remembrance of things past and imagined futures. Consciousness
Cogn. 18, 1018–1028.
van den Hout, J.H.C., Vlaeyen, J.W.S., Peters, M.L., Engelhard, I.M., van den Hout,
M.A., 2000. Does failure hurt? The effects of failure feedback on pain report,
pain tolerance and pain avoidance. Eur. J. Pain 4, 335–346.
Vytal, K., Hamann, S., 2010. Neuroimaging support for discrete neural correlates of
basic emotions: a voxel based meta analysis. J. Cogn. Neurosci. 22, 1–22.
Wai, S.T., Bond, A.J., 2001. Serotonergic involvement in the psychosocial dimension
of personality. J. Pharmacol. 15 (3), 195–198.
Watson, D., Clark, L.A., Tellegen, A., 1988. Development and validation of brief mea
sures of positive and negative affect: the PANAS scales. J. Pers. Soc. Psych. 54,
1063–1070.
Watt, D.F., Panksepp, J., 2009. Depression: An evolutionarily conserved mechanism
to terminate separation distress?: a review of aminergic, peptidergic, and neural
network perspectives. Neuropsychoanalysis 11, 5–104.
Way, B.M., Creswell, J.D., Eisenberger, N.I., Lieberman, M.D., 2010. Dispositional
mindfulness and depressive symptomalogy: correlations with limbic and self
referential neural activity during rest. Emotion 10 (1), 12–24.
Widiger, T.A., Costa, P.T., 2002. Five factor model personality disorder research. In:
Costa, P.T., Widiger, T.A. (Eds.), Personality Disorders and the Five Factor Model
of Personality. , second ed. American Psychological Association, Washington, DC,
USA, pp. 59–87.
Wiggins, J.S., Pincus, A.L., 2002. Personality structure and the structure of personality
disorders. In: Costa, P.T., Widiger, T.A. (Eds.), Personality Disorders and the Five
Factor Model of Personality. , second ed. American Psychological Association,
Washington, DC, USA, pp. 103–124.
Wu, J.C., Buchsbaum, M.S., Hershey, T.G., Hazlett, E., Sicotte, N., Johnson, J.C., 1991.
PET in generalized anxiety disorder. Biol. Psychiatry 29, 1181–1199.
Zaza, C., Baine, N., 2002. Cancer pain and psychosocial factors: a critical review of
the literature. J. Pain Symptom Manage. 24 (5), 526–542.