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Cross-Species Neuroaffective Parsing of Primal Emotional Desires and Aversions in Mammals

Jaak Panksepp
Emotion Review 2013 5: 235
DOI: 10.1177/1754073913477515
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477515
2013

EMR5310.1177/1754073913477515Emotion ReviewPanksepp Primal Emotions and Desires

Cross-Species Neuroaffective Parsing of Primal

Emotional Desires and Aversions in Mammals

Emotion Review
Vol. 5, No. 3 (July 2013) 235240
The Author(s) 2013
ISSN 1754-0739
DOI: 10.1177/1754073913477515
er.sagepub.com

Jaak Panksepp

College of Veterinary Medicine, Washington State University, USA

Abstract
The primal motivational systems of all mammals are constituted of the evolved affective brain networks that gauge key survival
issues. However, since progress in functional neuroscience has historically lagged behind conceptual developments in psychological
science, motivational processes have traditionally been anchored to behavioral rather than neural and affective issues. Attempts to
retrofit neuroaffective issues onto established psychological-conceptual structures are problematic, especially when fundamental
evidence for primal affective circuits, and their neural nature, comes largely from animal research. This article provides a synopsis of
our growing understanding of primary-process emotional systems of mammalian brains and minds, which provides a new empirically
based infrastructure for higher levels of human theorizing.

Keywords
affective neuroscience, approach, avoidance, motivations

Our understanding of approach and avoidance motivations, and

associated positive and negative affects, has advanced enormously in the past few decades (Elliot, 2008). However, largely
missing from human psychological analyses have been (a) empirically based evolutionary-hierarchical views of the relevant neural substrates, (b) empirical parsing of primal brain networks that
exist for diverse motivational BrainMind functions, and hence
(c) linkages of such knowledge to discussions of higher motivational processes in humans. Affective neuroscience helps fill this
vacuum with evidence for diverse primary-process emotional
systems that generate seven affectively motivated approach and
avoidance patterns (Panksepp, 1998; Panksepp & Biven, 2012).
Still, scientific approaches that acknowledge experienced
emotions and motivations in other animals are still deemed
problematic (Dawkins, 2012; LeDoux, 2012). For a classic
overview that led comparative psychologists away from the
mind, see Chapter 1 of Warden, Jenkins, and Warner (1935,
p. 136); as noted there: Mentalistic theories may be discarded
at once on the ground that it is impossible to obtain valid data
concerning the conscious life of infra-human organisms.
Accordingly, it is understandable that evidence from other animals has been neglected in psychology. Neuroscience can
change that bias since rewarding and punishing properties
of subcortical circuits can be empirically evaluated and can
illuminate human emotions and motivations.

Here I focus on empirically illuminated primary-process

emotional and motivational systems of mammalian brains
namely the unconditional (instinctual) emotional response systems that help control higher BrainMind functionsfor
example, secondary processes that mediate learning and memory, and tertiary processes that control higher mental processes,
best studied in humans. There is insufficient space here to highlight all views (see Lewis, Haviland, & Barrett, 2008; and for
controversies with constructivist approaches, see Zachar and
Ellis, (2012). Suffice it to say, brain and mind contain interlinked (nested) evolutionary layers (Figure 1), and few have
focused on primal neuro-emotional issues best illuminated by
animal brain research (Panksepp, 1998, 2011). Higher emotional and motivational structures, best elucidated by human
research (not addressed here), remain functionally linked to primal affective functions that emerged earlier in brain evolution.
So how can we scientifically evaluate the primal emotional
experiences of nonspeaking animals? A key advance has been
localization of instinctual emotional systems using deep brain
stimulations (DBS), and evaluation of raw affective experiences
(primal consciousness) empirically by determining whether
such evoked brain states serve as rewards and punishments
using traditional learning tasks (Delgado, Roberts, & Miller,
1954; Olds & Milner, 1954; Panksepp, 1971). They do, quite
uniformly! There are no rewards or punishments for

Corresponding author: Jaak Panksepp, College of Veterinary Medicine, Department of VCAPP, Washington State University, PO Box 646520, Pullman, WA 99164-6520, USA.
Email: jpanksepp@vetmed.wsu.edu

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236 Emotion Review Vol. 5 No. 3

Figure 1. A conceptual summary of evolutionary hierarchical bottomup (nested) and topdown (circular) causation that presumably operates in every
primal emotional system of the brain. This schematic summarizes the hypothesis that in order for higher MindBrain functions to operate, they have to
be integrated with the lower BrainMind functions, with primary processes being depicted as squares, secondary processing (learning and memory) as
circles, and tertiary processing (higher cognitive functions), at the top, as rectangles. Please imagine each symbol being color-coded, to better envision
the nested hierarchies that integrate the various levels of the BrainMind (adapted from Panksepp, 2011, which contains a color rendition of this
conceptualization). Bottomup control prevails in early infancy and early childhood development. Topdown control is optimized in adulthood.

humans that are not affectively experienced. This correspondence allows access to positively and negatively valenced emotional arousals in animals (Panksepp, 2011), and evaluation of
discriminability of those affective states (Stutz, Rossi, Hastings,
& Brunner, 1974). Abundant evidence indicates that subcortical
emotional brain state-shifts are affectively experienced by
humans (Coenen, Schlaepfer, Maedler, & Panksepp, 2011;
Panksepp, 1985). With accruing understanding of neurochemical codes of these systems, additional predictions are made
about affective arousals in humans (Panksepp, 1998). Such
epistemological possibilities, hardly recognized in academic
psychology, facilitate neuroempirical understanding of emotional states and associated motivational processes in humans.

How Neuroscience Illuminates Emotional

Approach and Avoidance Feelings
This article will contextualize discussions of approach and avoidance motivations within the emerging affective neuroscience
strategies that seek to understand the fundamental valuative (i.e.,
affective) emotional foundations of mind (for detailed coverage,
see Panksepp, 1998; Panksepp & Biven, 2012). In this view, the
broad generic concept of motivation circumscribes many types
of fundamental BrainMind functions (primary processes) that
contribute to organisms maximizing positive affective states and

minimizing negative ones. Such foundations of human mind,

deeply subcortical, promote the construction of higher cognitive
processes (tertiary brain states) through learning and memory
(here called secondary processes). Clearly, the BrainMind has
evolutionary progressions (layerings) still evident in brain
organizationolder brain functions being more caudal and
medial in the brain, with more recent ones being more rostral and
lateral. These levels of organization, interpenetrant in nested
hierarchical ways (Figure 1), are described next.

Primary Processes
At the foundation of the mind, various unconditioned stimulus
(UCS) and unconditioned response (UCR) affect systems undergird mental life. Such fundamental tools for living, like
SEEKING (capitalized to highlight a new neurofunctional
nomenclature), remain poorly studied in psychology, partly
because deep understanding requires brain research best pursued through affectively sensitive work on other animals.

Secondary Processes
At the intermediate level of neural organization are a variety of
learning systems that interface UCSs and UCRs into learned
subroutines. These intermediate functions arise from brain areas

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Panksepp Primal Emotions and Desires 237

called the basal ganglia, which include brain regions such as

amygdala, bed-nuclei of the stria terminalis, dorsal and ventral
striatal regions, and hippocampus. They allow basic needs of
body and brain to be interfaced effectively with environments in
which organisms find themselves.

Tertiary Processes
On top of these are species-typical neocortical expansions, tissues called general computational space which allow for the
emergence of higher cognitive strategies that are commonly discussed in terms of thoughts and decision-making strategies, and
ruminations when more intensely affective.
We have optimal access to primary processes in animal
research, and thereby affective qualia that are foundational
for mind (Panksepp, 1998; Panksepp & Biven, 2012). Most
tertiary-process issues are best studied in humans. Secondary
processes are well studied in both. With primary-process brain
research, we can move beyond the law of effect, (Thorndike,
1911) toward neuromechanistic laws of affect whereby silent
synapses are converted to active ones through environmentally
precipitated brain affective shifts (Panksepp & Biven, 2012).
Distinct types of primal affects are promoted by homologous neural circuits concentrated in subcortical regions in all
mammals, ready to guide behavior and learning soon after
birth. These motivation-generating systems come in distinct
flavors, with homeostatic, sensory, and emotional affects
being meaningful global categories, each with many exemplars. Primal affects get connected to environments through
learningsecondary processing which, stated simply, coax
animals to promote good feelings and diminish bad ones
thereby allowing internal survival issues to be related to world
events. I ignore tertiary processes herehigher psychological
processes, much harder to study in animals than humansbut
simply note how intimately they are intermeshed with primal
affective values.
Although human cognitive approaches have little empirical
access to subcortical neural mechanisms of primal affects, we
can analyze evolutionarily related animal feelings by studying
the rewarding and punishing properties of subcortical emotional
UCRsaction networks that feel like something when aroused.
The sensory and homeostatic affects are harder to study neuroscientifically because they are not as intimately linked to discrete action systems. Those affects are more linked to UCSs in
traditional behavioral parlance.
The DBS-validated primal emotional affective systems
deserve a special scientific nomenclaturefor example,
SEEKING, RAGE, FEAR, LUST, PANIC/GRIEF, and PLAY
(Panksepp 1982, 1998). In labeling primal emotional processes,
semantic ambiguities were avoided, while not losing the needed
communicative clarity, through the convention of capitalizing
common vernacular terms. The neuroanatomical referents for
these emotional labels are well identified with DBS, and the
underlying networks and neurochemistries, albeit presently
only partly characterized, can be further clarified with new neuroscientific tools (e.g., neurochemically specific optogentics).

Before proceeding to the emotional primes, a few additional

reflections on the other major categories of affectsthe sensorially
based ones. The primal UCS-linked affective systems come in two
general categories: homeostatic and sensory. The homeostatic
affects operate via various interoceptive systems that monitor bodily states such as energy and water balances (Denton, 2006); in the
vernacular these are called hunger and thirst. Many other general
bodily states (feelings such as nausea/disgust, fatigue, and thermoregulatory coldness and warmth). When discussing such issues
as primary homeostatic feelings, we may again wish to use the
same capitalized terminological convention as employed for
emotional primes.
The molecular underpinnings of sensory affective processes
are being deciphered more rapidly (see Kringelbach & Berridge,
2010; Rolls, 2005). Taste responsivity has been well analyzed
by studying instinctual orofacial action patterns when different
taste qualities are infused into mouths of animals (Berridge,
1996), leading to clarification of brain regions and neurochemistries controlling behavioral liking reactions. The affective
implications remain to be clarified by direct evaluations of
whether identified subcortical taste regions mediate rewarding
and punishing effects with DBS.
I will now focus on the within-brain emotional action systems
that currently provide the most well-developed approach to
understanding the neural nature of affective qualia so important
for understanding motivational states. DBS evidence indicates
that the dynamics of emotional-instinctual action schema are the
fundamental substrates for the corresponding raw emotional
qualia. As noted, wherever in the brain we can evoke coherent
emotional action patterns, there exist corresponding affective
states as monitored with DBS evoked rewarding and punishing
within-brain phenomenal experiences. So far the aroused
valenced states appear to be homologous in all mammals, allowing falsifiable neuroscientific predictions across species. Thus,
the study of these systems in animals currently provides the optimal scientific strategy for decoding the constitution of affective
consciousness, even for humans. Such cross-species knowledge
may also help us fathom the motivational changes that accompany psychiatrically significant affective imbalances, providing
strategies for development of new mind medicines such as antidepressants (Burgdorf, Panksepp, & Moskal, 2011).

The Basic Emotional-Affective Circuits of

Mammalian Brains
Extensive descriptions of primary-process emotional-motivational
networks are in Panksepp (1998) and Panksepp and Biven
(2012), so only thumbnail sketches will be shared here.

The SEEKING/Desire System

SEEKING is a general-purpose appetitive motivational system
that allows many other emotional systems to operate effectively
(e.g., LUST, CARE, and PLAY). It seems to be a major source
of appetitive coherence that has at times been called libido or

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238 Emotion Review Vol. 5 No. 3

euphoria. It provokes animals to become intensely interested

in exploring their world, allowing them to find and eagerly
anticipate all kinds of resources they need for survival, from
water, food, sex, and warmth to addictive drugs and social relationships, via shared neurochemistries. Thus, even though sensory affects probably have hedonic qualia of their own, these
systems also impact emotional responses, adding motivational
intensity, at a distance, to resource acquisition. For instance, all
affectively positive sensory stimuli impact functions of the
general-purpose SEEKING system that promotes unconditional
appetitive emotional actions (e.g., increased exploration) and
learning (anticipatory wanting of specific resources).
When fully aroused, the SEEKING urge fills the mind with
eagerness and interest, and motivates organisms to move their
bodies seemingly effortlessly in search of resources they need,
crave, and desire. It sustains both mundane curiosity and sublime intellectual pursuits. When underactive for various reasonsfrom social defeat, promoting chronic helplessness/stress,
or neural deficits of old agedepression can ensue. When overactive, which can arise from drug abuse, behavior can become
excessive, guided often by psychotic delusions and manic
thoughts (Coenen et al., 2011).
Neuroanatomically, SEEKING arises from the major selfstimulation system (the medial forebrain bundle) that runs from
the midbrain ventral tegmental area (VTA) up to the medial prefrontal cortex. Animals self-stimulate this system readily,
whether via abuse of drugs or direct electrical or chemical stimulation of the brain (depending on availability, of course).
Although still poorly conceptualized as a reward, pleasure, or
reinforcement system, it actually coaxes animals and humans to
move energetically from where they are presently situated to the
places where they can find and consume resources needed for
survival. In rats, simplified objective measures of this system in
action are positive affect indicating 50 kHz ultrasonic vocalizations and invigorated sniffing (Burgdorf, Wood, Kroes, Moskal,
& Panksepp, 2007). SEEKING promotes appetitive learning by
computing reward relationships in the world, called reward prediction errors in one scheme (Schultz & Dickinson, 2000). A
key neurochemical that promotes SEEKING is dopamine, which
innervates higher brain frontal neocortices that mediate planning
and foresight, while concurrently engendering anticipatory
eagerness, both at behavioral and affective psychological levels.

The RAGE/Anger System

When SEEKING is thwarted, RAGE is aroused. Ideal stimuli are
frustrations and attempts to curtail animals freedom of action.
RAGE can be provoked by DBS along a longitudinal transhypothalamic pathway coursing from the periaqueductal gray
(PAG) to amygdala (Panksepp, 1971), near circuits that promote
FEAR (leading to the misleading fight or flight system moniker). RAGE invigorates affective attack and helps animals
defend themselves and their possessions, and is commonly effective in arousing FEAR in their opponents. Human anger may
derive much of its psychic energy from this brain system, since
intense, unprovoked anger attack can be instigated by DBS along

the RAGE system which runs from medial regions of amygdala,

through ventral hypothalamus and dorsal PAG. Key neurochemical activators of this system are Substance P and glutamate,
while endogenous opioids quell RAGE.

The FEAR/Anxiety System

FEAR circuitry, paralleling RAGE, helps animals avoid pain
and destruction. Mild arousal of FEAR promotes freezing and
hiding; intense arousal precipitates flight. Humans stimulated in
these same brain regions report being engulfed by an intense
free-floating anxiety with no environmental cause. Key chemistries that regulate this system are glutamate and corticotrophin
releasing factor (CRF), and specific antianxiety agentsfor
example, benzodiazepinesinhibit this system.

The LUST/Sexual Systems

Male and female sexual urges are mediated by LUST brain circuits, both overlapping and distinct. LUSTs are aroused by male
and female sex hormones, which regulate brain neuropeptides
such as vasopressin and oxytocin, respectively. Such brain
chemistries promote gender-specific sexual tendencies, including gentle prosocial acceptance by oxytocin, and assertiveness,
perhaps even jealous behaviors, by vasopressin. Distinct male
and female LUST circuits, differentiated in the hypothalamus
during infancy, become differentially aroused by gonadal hormonal tides at sexual maturity. Because brain and bodily sexuality are independently organized, mammals that are externally
male can have female-specific sexual urges. Individuals with
female-type appearances can have male-typical sexual urges.

The CARE/Maternal Nurturance System

Brain evolution has provided safeguards to assure that parents
(usually mothers) devote CARE to offspring. The massive hormonal changes at the end of pregnancy (declining progesterone,
and increasing estrogen, prolactin, and oxytocin) set the stage
for the activation of maternal urges a few days before parturition. This symphony of hormonal and neurochemical changes,
especially the heightened secretions of oxytocin and prolactin,
facilitate maternal moods which assure strong social bonding
with the offspring. Similar neurochemicals, especially oxytocin
and endogenous opioids, promote infant bonding to mothers.
These changes are foundational for one variant of love. The fact
that chemistries for female sexuality (e.g., oxytocin) have been
redeployed in nearby circuits to promote maternal care (nurturance and social bonding), suggests an intimate relationship
between female sexual rewards and maternal motivations
(Panksepp & Biven, 2012).

The PANIC/GRIEF SeparationDistress System

Young mammals are all dependent on maternal CARE for survival. In turn, they have powerful emotional voices to solicit
careintense crying when left alone in strange places. Such

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Panksepp Primal Emotions and Desires 239

separation cries alert caretakers, especially mothers, to SEEK

troubled infants and attend to their needs. The underlying neural
system has been mapped with DBS in several species (including
dorsal PAG, dorsomedial thalamus, and bed nucleus of the stria
terminalis), and the resulting PANIC response, engendering the
psychic pain of aloneness/sadness/grief, is robustly inhibited
by endogenous opioids, oxytocin, and prolactinthe major
social-attachment/social-bonding chemistries of the mammalian brain. This PANIC network is also aroused during human
sadness and depression states accompanied by low brain opioid
activity. Sudden arousal of this system may contribute to panic
attacks in humans.

The PLAY/Rough-and-Tumble, Physical

Social-Engagement System
Young animals have strong urges for physical play, arising from
specific subcortical brain regions (e.g., the SEEKING system
along with parafascicular thalamus) and consisting of pouncing
on each other, chasing, and wrestling. Although outwardly
aggressive, PLAY arouses positive affectsocial joy accompanied, in rats, by abundant high frequency (~50 kHz) chirping
sounds that resemble human laughter (Panksepp, 2007b).
Tickling young rats and human children maximizes their happy
chirping/laughter, promoting social bonds and friendships.
While social play can promote social dominance, PLAY presumably helps young animals learn about their social potentials,
all those complex social processes that need to be learned.
PLAY urge may promote the epigenetic construction of higher
social brains. There are many reasons to believe that children
deprived of social play are more likely to have impulse-control
problems, namely attention deficit-hyperactivity disorder
(ADHD), which can be effectively treated with psychostimulants, all of which reduce playfulness in young rats. Abundant
juvenile play may help reduce the incidence of ADHD
(Panksepp, 2007a).

Conclusion
Abundant work supports the existence of at least seven distinct
rewarding or punishing emotional systems identified in diverse
species with subcortical DBS. These systems consist of large
transverse networks, able to engender affective consciousness,
concentrated in the extended reticular activating systems (ERAS)
of mammalian brainstem, brain regions long known to be essential for consciousness. ERAS circuits interdigitate with emotiongenerating regions such as the PAG and VTA and various learning
regions such as basal ganglia, including amygdala and nucleus
accumbens as well as tertiary-processing regions such as cingulate and medial frontal cortices. All emotional systems are well
represented in the PAG and connected to higher limbic regions
through the hypothalamic and medial thalamic corridors.
Each system has abundant descending and ascending components that work together in a coordinated fashion to generate
various instinctual emotional behaviors as well as the raw feelings normally associated with those emotional behaviors. These

systems most likely interact with higher brain mechanisms, to

create higher motivational structures. More work is needed to
understand the interactions of these systems in detail, and how
they interact with homeostatic and sensory affects, all relating,
in part, to the general-purpose SEEKING urge, which is the
most prolific emotional-motivational system of them all. This
may allow us to understand apparent paradoxes, such as how
anger can be an approach motivation (Harmon-Jones, 2004)
while also generating an aversive (withdrawal) state. It is
becoming ever clearer that the SEEKING system also helps deal
with negative affective states, and that FEAR promotes the
seeking of safety. Perhaps many past conceptual conundrums
the rejection of emotional feelings in animalsarose from the
need to explore behavior from the outside, as opposed to the
internal neurodynamics of the brain.
In sum, the existence of emotional feeling states is affirmed
by animals never being neutral about DBS-evoked emotional
arousals. They find arousals of SEEKING, LUST, CARE and
PLAY systems to be rewarding; they escape arousals of
RAGE, FEAR, and PANIC systems, indicating those diverse
states are punishing. How different they are from each other can
be evaluated, even in animals, with DBS discrimination procedures, and in humans with verbal self-reports of shifting
experiences. These brain motivational systems may provide
endophenotypes for psychiatry, and affective psychoassays for
psychiatric medicinal development (Panksepp, 2012). By taking emotional feelings of other animals seriously, we have
already identified new prohedonic neurochemicals and thereby
potential new antidepressant medications (Burgdorf et al.,
2011), including new treatments for autism and ADHD
(Panksepp & Biven, 2012).
The affective internal motivational experiences of animals,
long debated and still often denied by behaviorists and behavioral neuroscientists (LeDoux, 2012) and ethologists (Dawkins,
2012), can finally be addressed empirically through affective
neuroscience strategies (Panksepp, 1998, 2011). Of course, one
must also specify reasonable evolutionary scenarios for their
emergence by considering the adaptive function of such experiential capacities. That is conceptually simple: Primal affects
intrinsic neural valuesinform animals about bodily survival
issues that promote and guide learning. Positive affects indicate
trajectories of well-being, and negative ones, paths of destruction. Affective feelings intrinsically anticipate survival, with a
wider event horizon than behavior alone, which can refine
learning. Affects are the immediate psychological currency of
our motivations.
Primal affects may have been the first experiences (qualia)
that existed on earth. Without them, capacities for conscious
awareness may not have emerged in higher brain evolution
(Solms & Panksepp, 2012). Whether other animals also have
more subtle higher order feelings, so abundant in humans who
have diverse cognitive experiences, remains currently scientifically unknown. As we solve foundational affective problems,
we may be better positioned to empirically address the more
difficult higher issues that can only be well studied in human
beings.

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240 Emotion Review Vol. 5 No. 3

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