In Memoriam

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Professor Jaak Panksepp. (Photograph courtesy of Bowling Green State University)

Personal Reflections on the Neuroscientific Legacy of Jaak Panksepp (1943-2017)

Douglas F. Watt, School of Medicine, Boston University

 

“Nature has placed mankind under the government of two sovereign masters, pain and pleasure . . .they govern us in all we do, in all we say, in all we think: every effort we can make to throw off our subjection will serve but to demonstrate and confirm it.” – Jeremy Bentham

July 2017 – One of the truly great voices in modern neuroscience was silenced recently when Jaak Panksepp suddenly died on April 18th of this year, following a fairly routine surgical procedure that went badly awry, generating a serious ischemic-hypoxic insult, a tragedy from which not even Jaak Panksepp could recover.  Jaak had battled long and hard in relationship to recurring bouts with various cancers, and is too often the case, it was actually the cancer therapy process that eventually killed him and not the cancer itself, although that distinction offers little solace.  Nevertheless, the terrible and final suddenness of all this, despite Jaak’s age and medical history, was shocking and deeply upsetting to all of us who were close to Jaak.  None of us is truly prepared for these departures at the end of life, even though we all know that they are coming eventually… and inevitably.  Jaak’s neuroscientific work was, as many appreciate, centered at least in part around separation distress as a prototype emotional state, and he has had much to say, over the many years of his career, about the nature of the very grief that all who knew Jaak well have been feeling since his passing.  That grief still lies heavy on many hearts, of family, friends and colleagues.

The personal and professional losses for me are almost impossible to separate, as I lost a senior colleague, a highly valued friend, and a mentor in neuroscience, who was decisively the largest and most positive single influence in the second half of my professional career.  But neuroscience lost at least as much, an irreplaceable leader, a brilliant scholar and scientist, and a most passionate advocate for an affect-centered view of the brain and mind.  I take solace in my sense of great good fortune for having had 20+ years of close colleagueship and friendship with Jaak, sharing half a dozen week-long Affective Neuroscience seminars, many book chapters, dozens of larger conferences and other fruitful and sundry projects.  I believe that over time his legacy and many gifts to the sub-fields of mind and brain science will prove increasingly resonant and prescient.  Progress in science is intrinsically slow by its very nature, but once in a while a highly creative genius comes along, whose insight and integration of large amounts of previously fragmented material, creating new paradigms and concepts, moves the field forward in sudden and deceptive leaps.  In the increasingly ‘twinned’ fields of neuroscience and psychology, Jaak was one of those people.

By any conventional standard of scientific productivity, Dr. Panksepp was a giant contributor in relationship to the neuroscience of emotion and clearly staggeringly prolific, with over 450 empirical and review publications, and literally many dozens of book chapters in various edited volumes and compilations.  His publication vitae centrally featured his 1998 textbook, Affective Neuroscience, increasingly regarded as a landmark and a classic publication in neuroscience.  This was updated with Lucy Biven in the less technical and more accessible treatment, The Archaeology of Mind.  He also solo edited the well-regarded two-volume Advances in Biological Psychiatry in 1996, the more recent Textbook of Biological Psychiatry in 2004, and the highly technical compilation Handbook of the Hypothalamus with Peter Morgane which few have read but which, despite its age, is still to this day the most comprehensive collection of work on that critically important neural system and its functional networks.  The very last of his many edited volumes, The Psychology and Neurobiology of Empathy, was completed with me in 2016.  This huge body of work has had an enduring and pervasive influence on the fields of psychology, psychiatry, and both clinical as well as experimental neuroscience, and has inspired at least two generations of young scientists eager to understand more about the mysteries of the brain and mind and the pivotal role of emotion in its neurodynamics.

Where and what were Jaak Panksepp’s greatest contributions?  This is not an easy question to answer, because his interests spanned across many traditional disciplinary boundaries, including such disparate topics as a theory of music, the fine-grained anatomy and connectivity of the mesodiencephalon, and the nature of laughter as an affective signal.  And yet the common denominator in all of these was Jaak’s keen and unwavering focus on the nature of affect and the implications of affect for every other important psychological and neuroscientific territory.  The following list of Jaak’s major areas of scientific contribution has to be considered simply my personal scientific opinion, and therefore hardly definitive or unbiased, but I will outline three main areas of scientific contribution.  Others, indeed, many others might be added, but space considerations demand a shorter list – so here’s mine.

On a fundamental and intrinsic relationship between consciousness and emotion

As perhaps his most critical contribution, Jaak continually emphasized the centrality of emotion and affect in any potential neuro-architecture for a conscious mind, what I would call an ‘affect-centric’ view of the embodied mind.  Affects were evolutionarily conserved core routines guiding the living mind.  This strong conviction that emotion provided not just an interesting ‘coloration’ to consciousness contrasted starkly with the cognition-centered and the predominantly sensory view of consciousness at the time of his early seminal work and publications.  Instead of this conventional sensory/cognitive perspective on conscious architectures, affects were conceptualized as signaling functional integrations foundational to any version of a conscious architecture; additionally, all strong affects were thus intrinsically conscious, a shared scientific conviction that originally brought us together and created an immediate intellectual bond that was to energize our many shared ventures and projects.

Thus it was no coincidence that we met for the first time in the second biannual Tucson Series on the Science of Consciousness in 1996.  It would be an understatement to suggest that affective neuroscience perspectives might have been underrepresented at the Tucson Conference in 1996.  We shared an immediate sense of  dismay, tempered perhaps by a certain amusement, that the fields of psychology and neuroscience seemed at times so blissfully unaware that a mind without the internal ‘value compass’ provided by emotion and affect had no basis for organizing basic behavioral, or for that matter, higher attentional and cognitive priorities. “What are they smoking?” we wondered over drinks at that first conference!

We shared a sense of astonishment that affects were not more universally appreciated as the true ‘reinforcers’ referenced by behaviorism, and wondered how anyone in the many disparate fields of psychology and neuroscience could somehow miss this central ‘keystone’ fact of our psychology.  As consciousness had been for the four previous decades a mostly discredited subject under the still powerful aegis of behaviorism, we were joined in this early conviction in the mid-1990s by precious few in neuroscience, but notably, by Antonio Damasio.  Damasio, Joe LeDoux, and several other luminaries in emotion theory and consciousness studies all participated as commentators in one of my first ventures with Jaak, an ASSC (Association for the Scientific Study of Consciousness) web-seminar on Emotion and Consciousness given in the summer of 1998, which generated lively debates still resonating to this day about exactly how to conceptualize the role of emotion in consciousness.  Can consciousness and emotion be simply conceived of as largely orthogonal processes, with just a single intersection, when emotion created a conscious affective state?  We argued that this single intersection was just the tip of the iceberg, in terms of the rich interweaving of affective signals into every aspect of conscious life, from what we found interesting, to what we avoided, to what we craved and sought out, from what we learned forever to what we forgot next week, indeed, virtually every aspect of the mind.

The crux of our early shared conviction was simply that affects (both homeostatic and emotional) and their underlying brain systems, had to be foundational in some poorly understood fashion for how neural activity gives rise to an organized agency – a ‘selfhood’ in the brain, and furthermore, that such a self would be an ‘agentic self’ and not any version of a passive observer, with behavioral initiatives and basic motivation intrinsically organized around and by those emotional and homeostatic affects.  In other words, it made sense to assume that all behavior was forever hinged to the pursuit of the many and varied positives and also equally in the service of minimizing the many varied negatives, echoing Bentham’s early brilliant intuition, an insight essentially recapitulated by Freud 100 years later.  Without those ‘value signals’, there was simply no other basis for motivation or behavior.

This set of early assumptions – which ran largely counter to the sensory and cognition-centric views in early consciousness studies – is being slowly but progressively validated, but we found an unlikely early ally in the global workspace perspectives of cognitive theorist Bernie Baars and colleagues, such as Jim Newman.  This central neurologic dependency of conscious architectures upon affective signals was revealed in the neurodynamic dependence of cortical systems on the integrations achieved in the mesodiencephalon to do any form of meaningful and organized cognitive work (see Merker, 2007 for the most definitive and exhaustive summary to date).  It was also revealed starkly in the underappreciated syndrome of akinetic mutism, where collapse of emotion and motivation leaves a virtually rudderless and engine-less thalamocortical system and a disorder of consciousness (see Watt and Pincus, 2004 and Watt, 2012 for reviews).  And while one can say that neuroscience still lacks its ‘keystone in the arch’ – a definitive and well-validated neural theory of mind as one of the core emergent properties and mysteries in Nature – increasing evidence argues that conscious architectures are highly distributed and non-local.  They exist as transient adaptive networks, flitting about in an ‘enchanted loom’ in Sherrington’s haunting metaphor, with new transient and highly distributed alliances being formed on a moment-to-moment basis, but bridging the entire vertical axis of the brain.

At this point in the evolution of a neuroscientific theory of mind, the critical role of the mesodiencephalon in such distributed networks, as upper brainstem and medial diencephalic territories rich in ancient homeostatic and affective systems, is simply beyond meaningful dispute.  An extensive lesion of these mesodiencephalic territories will collapse consciousness totally and permanently, even without a trace of damage to any aspect of the more telencephalic thalamocortical systems that are virtually the exclusive anatomical focus of cognitive neuroscience.  Indeed, just a full lesion of the tiny territory of midbrain PAG – at the ventral tip of the large group of brain affective systems running from paleocortex through limbic regions, to hypothalamus and down into the brainstem, leaves the person ‘gone’ in a comprehensive sense, and with a permanent and severe akinetic mutism.  This may hint at PAG’s role in laying the neuroevolutionary foundations for those more evolved affective systems, sitting at the bottom of the highly distributed affective networks extending upward from reticular-hypothalamic to limbic and paralimbic systems.

The neurodynamics of this profound ‘system integration’, particularly in terms of a developmental trajectory that bootstraps the conscious mind from core homeostatic and affective systems, clearly remain to be more fully elucidated, but the ontogenetic development of consciousness for each of us appears to clearly recapitulate our phylogeny, as the brain builds networks from the inside (ventral and medial regions) out to the more telencephalic lateral and dorsal regions.  Although the cortex still dominates cognitive neuroscience – and with good reason as it forms the neural substrate for virtually all cognitive content – I believe that future generations will strongly credit those handful of neuroscientific pioneers who kept pointing at neurodynamic integrations, still poorly understood, but achieved first in more ancient mesodiencephalic, tectal and tegmental systems, as foundational for all the cognitive activity at the top of the system.  Jaak will likely be highly regarded and credited as one of those prescient pioneers.

On understanding emotion in terms of interactive but partially discrete prototype states energized by an overarching, precedent and integrative SEEKING system

Jaak Panksepp also made pivotal contributions to unraveling one of the central mysteries of emotion, namely, that it somehow functions as a central clearinghouse in which all emotional activators must compete and interact, a fact which necessitates global and highly distributed neural system architectures for emotion (and an underappreciated direct correlate of the global network ideas just reviewed).  In other words, emotion cannot be allocated to any kind of simple ‘module’ or any version of a local network, but has to be understood as a distributed integrative system, a realization which made him rather fond of placing the affective primes in all capitals (‘FEAR’, ‘RAGE’, ‘GRIEF’, ‘PLAY’, ‘SEEKING’, ‘CARE’), as his way of denoting this distributed or ‘system’ property.  Although this might have seemed semantically ‘quirky’ to some (including even to this author at times J), it was energized by Jaak’s realization that more global system concepts were mandated, and where perhaps the various affective primes might function as ‘nodes’ within an integrated system, thus creating a ‘neural clearinghouse’ in which all the potential affective states could compete and interact in a variety of adaptively critical antagonistic and agonistic ways.  And Jaak would truly cringe when someone might extrapolate that each of these capitalized primes might be some kind of ‘module’ in the brain.

In his view, the negatives and positives would have to be understood as fundamentally antagonistic, but there were also more subtle versions of promotion and inhibition between systems of similar valence.  For example, while playfulness might ‘set the table’ so to speak for the activation of sexual desires, at some point excessive playfulness might actually inhibit activation of the LUST system.  Such an image of complex agonistic and antagonistic systemic interaction between the primes fits beautifully with the phenomenology of emotion and emotional behaviors, as all kinds of emotional pushes and pulls clearly do agonize and antagonize one another in various ways – part of what gives our (and general mammalian) behavior its delightful and frustrating unpredictability.

In this sense, I believe many, particularly the social constructionist school of emotion theorists, did not fully appreciate how in Panksepp’s system of the seven emotional primes, the SEEKING system was conceptualized as a “special class of one”, and that it had to be the evolutionarily precedent or ‘master’ system, a supposition recently gaining impressive validation by empirical work on dopamine in crayfish (Huber et al., 2011).  In social constructional theories of emotion, the notion of ‘core affect’ (i.e., Russell and Barrett, 1999), with an emphasis on dimensional aspects of approach/avoidance, arousal, and valence, is not actually operationally dissimilar to how Panksepp conceived of the integrative and superordinate role of the SEEKING system, although there are still important differences between Panksepp and social construction theories that cannot be glossed over.  Panksepp parted company with constructional views if they went so far as to argue that the categorical or ‘emotional prime’ systems (observing for example a principled distinction between PLAY and RAGE as neurobiologically meaningful) were simply a specious cultural convention or ‘meme’ carried over from a pre-scientific ‘folk’ nomenclature, but without any real neurologic and biological basis.

Panksepp’s theory of the emotional ‘primes’ provides real ways of immediately and directly linking the SEEKING system to all the other primes.  In other words, FEAR was the seeking of safety, PLAY the seeking of rough-and-tumble joyful engagement, RAGE the seeking of an end (sometimes ‘with prejudice’) to that toxic agent or circumstance that might be the source of one’s frustration or injury, maternal CARE the seeking of safety, restored homeostasis or good feelings in a dependent and vulnerable other (and thus indexing a proto-empathy system, as classically outlined in Panksepp, 1998, and more recently in Watt and Panksepp, 2016).  In this critical sense, I sometimes felt that students as well as some critics of Panksepp did not always clearly appreciate that all of these classic affective ‘primes’ or prototypes had to be, at least in some sense, specialized ‘resonances’ or evolved ‘nodes’ interacting within a more ancient and precedent generalized motivational arousal or SEEKING system.  Perhaps this key insight was expressed too implicitly at times, and where this precedent role of the SEEKING system was not made more explicit, although it is stated at several points in his magnum opus (Panksepp, 1998).

One underappreciated benefit – indeed one great beauty of this set of concepts from the standpoint of scientific theory – is that it provided an integrated neural substrate for the competitive interaction of the emotional primes, where the negatives and positives could easily inhibit each other, a clear functional fact of affective life that any good model had to explain.  Additionally, the SEEKING system concept provided an easy evolutionary bridge from homeostasis to emotion, as the SEEKING system network contains, as one of its core nuclei, the lateral hypothalamus which receives abundant dopaminergic enervation from the midbrain, providing an immediate neural bridge for integrating behavior (via its extensive basal ganglia and mesolimbic connections), homeostatic imbalances, and sources of homeostatic relief.  Thus, in its most ancient forms, as Jaak famously offered, “the SEEKING system gets thirsty animals to water, hungry animals to food, cold animals to warmer environments,” etc.  The SEEKING system creates neural links from rewards and punishments to reward predictors or reward cues, and binds both of those classes to basic behavioral routines, such that a hungry animal can activate motor routines to approach a source of food and then initiate consummatory routines, under the guiding influence of this master motivational system.  It is hard to imagine a more elegant set of integrative predictive tools about emotion and motivation than what this set of basic ideas affords modern neuroscience.

Another underappreciated benefit of Jaak’s emotional system concepts was that they provided a highly plausible theoretical bridge for how homeostasis proper (likely the antecedent evolutionary process) might have given rise to the emotional primes, as they emerged from basic evolutionary-predictive extensions of core homeostatic mandates.  This was no coincidence, as Jaak cut his teeth on energy balance research.  In other words, perhaps fear reflects an anticipatory forward-looking extension of pain and the basic aversion to tissue damage.  In fear, we are not damaged yet by an approaching predator or more powerful rival, but we might be, if we do not seek safety, and either freeze to avoid detection, or flee, if this less metabolically costly solution is not possible.  In separation distress, a young infant mammal is not yet metabolically compromised but suddenly finds itself separated from both its source of metabolic supply (caretakers) as well as its primary protection from predation, suggesting that separation distress may have emerged as an anticipatory forward-looking extension of these basic homeostatic routines to ensure the safety of vulnerable offspring.

While this way of linking more ancient homeostatic processes with classic emotional primes remains still mostly untested, and in need of much further probing and investigation, there is no meaningful evidence (that I am aware of anyway) against it.  It is also deeply consistent with how evolution functions to extend the brain as an increasingly sophisticated prediction engine, stacking new predictions on top of older successful behavioral routines.  Evolution operates from the highly conservative principle of having no need to “re-invent the wheel” in every new context, just tweak and enhance the existing operations a bit here and there to manage the new adaptive context.  Encephalization is, in this real sense, nothing more than the creation of more and more sophisticated prediction engines (and consistent for example with evidence that Wernicke’s area is specialized for the prediction of phonemic sequences).  Jaak’s early work on the mysteries of feeding and energy balance exposed him at the beginning of his career to the challenge of finding integration across highly varied and different classes of motivational ‘vectors’ including evolutionarily more ancient and more recent classes of motivations, and his SEEKING system concepts were the brilliant solution.

This basic set of theoretical notions finds strong validation in the fact that opioids not only modulate pain and reward satiety (two very ancient homeostatic operations presumably evolutionarily preceding the classic emotional primes), but in more complex and extended mammalian brains they have come to modulate social bonding, play and separation distress (all critical emotional operations of a highly social brain).  Play and quieter forms of social comfort are high opioidergic states, while separation distress is organized in good part by the inverse condition of low opioidergic signals (validated in humans by Zubieta et al., 2003), coupled perhaps with high CRF signals.  In this sense, separation distress might reflect an evolutionary extension of pain, while social comfort in a secure attachment perhaps is an evolutionary extension of a basic homeostatic wellness and satiety.  Thus, the SEEKING system concept provides a ready and plausible evolutionary and functional bridge between homeostasis and emotion that no other neural system concept currently affords to my knowledge.

Panksepp was able to successfully interpret the confusing results of the electrical brain stimulation (EBS) experiments on the ‘self-stimulation system’ (as originally labeled by Milner and Olds), and was not seduced by the simplistic behaviorist concept that this was “the reward system.” Jaak realized from many experiments that this system had to be conceptualized differently from these seductive oversimplifications, as the data suggested that this system supported both the seeking of rewards but also the avoidance of punishments.  This SEEKING system concept allowed Jaak to achieve a theoretical integration and coherence where there had been previously only confusion and apparent contradiction, particularly around the way in which electrical brain stimulation of SEEKING system trajectories (such as VTA, accumbens or medial forebrain bundle) could generate such apparently disparate behavioral results.

Jaak realized that the behavioral and affective trajectories off of EBS should be variable, given the SEEKING system’s precedent and overarching role, depending on many variables, including species and animal personality phenotypes, adaptive context and even the current ‘mind set’ or mood of the animal, which would all influence which affective ‘prime’ might become emergent from EBS of the mesolimbic mesocortical DA system.  Thus, predatory animals like cats might be much more likely to go into a stalking mode, while forager species like rats might start sniffing around and exploring the environment, while restrained animals might actually just try to escape more vigorously.  Unfortunately, this behavioral phenotypic variability resulting from EBS “self stimulation” has been misconstrued by constructional theorists of emotion as hard evidence against the ‘categorical’ or ‘affective prime’ view.  But it’s not; it is simply more evidence that not everyone has clearly understood the full implications of Jaak’s complex ‘affective prime’ theory and the precedent, superordinate role of the SEEKING system.

Then there were the many discrete contributions (too many to specifically enumerate here) Jaak made to understanding each of these affective prototype system, aside from the overarching role of a SEEKING system, most particularly empirical contributions in relationship to separation distress/GRIEF, maternal CARE, the organism ‘defense states’ of FEAR and RAGE and perhaps his favorite system, PLAY.  The table below, first published in the ASSC Web-seminar on emotion and consciousness (from Watt, 1998) but then was featured in many of Jaak’s reviews.  This table summarizes many of his basic empirical findings (table extracted largely from Panksepp, 1998).

On the likely centrality of mammalian play in the construction of a social brain

Jaak Panksepp’s third area of critical contribution might be around placing the PLAY system more clearly front and center as a critical pro-social system in the mammalian and human brain.  There are several indices suggesting that we are perhaps the most socially dependent creature on the planet, with this a hidden corollary to the developmental costs of a big cortex and the attendant and intrinsic helplessness of human infancy.  The importance of play and its critical role in a variety of prosocial operations of the human mind and brain is still widely underappreciated.  For all the contributions of various branches of psychoanalysis to an affect-centric view of the mind, even psychoanalysis (with some notable exceptions) neglected this prototype emotional system, and sometimes treated play as simply a proxy for aggression and/or dominance under the aegis of its original ‘dual instinct’ theory.  There is now so much accumulating evidence that this system is critical in a profound way for humans – it promotes attachment, signals trustworthiness (we trust playful people far more readily and much more immediately than dour, irritable or guarded people), and interacts powerfully in a supportive and facilitating way with LUST and CARE (proto-empathy) systems.  Its cognitive extensions in humor ripple through much of our social discourse.  We can’t seem to get enough of it in fact.

Despite all the endless mass media and pop psychology chatter and confusion around what might create the basis for a good relationship, both men and women when surveyed about this question want first and foremost someone who is playful and has a good sense of humor (Chick et al., 2012).  And if the PLAY system in the brain is responsible for all forms of laughter, and if spontaneous smiling (as opposed to ‘social smiling’) is part of the PLAY affect continuum, and ‘on the way’ to laughter so to speak (amplification and strengthening of the ‘pro-smiling’ stimulus leads to outright laughter), this might well be the most underappreciated pro-social neural system in the human brain.  If spontaneous smiling and laughter are essential social signals which indicate that “things are indeed okay,” if playful interactions are a powerful augmenter of attachment, a predictor of who we are going to like and want to be with, and also potently inhibit the stress axis of the brain, Jaak’s advocacy for the critical importance of this highly prosocial system, at a time when almost no one else in neuroscience was paying much systematic attention to play, playfulness, or humor, looks more and more prescient.

What might my overall conclusion be from this body of work?  Certainly many different individuals might generate many different take-home messages from Panksepp’s work, but for me, as I stated at the beginning, it is virtually impossible for me to separate the personal from the professional in relationship to Jaak and his work.  The most enduring aspect of Jaak’s legacy for me simply is something rarely made explicit in his work, but which we talked about on a number of occasions as being more implicit – a deep reverence for an animate Nature, which may both energize the best science and may be an emotional endpoint of the best science as well.  Spirit appears to be matter profoundly organized into truly dizzying levels of complexity – that such wondrous properties as minds with affects emerge from a physical Nature over a long and still in many ways mysterious evolutionary course deepens a sense of awe and wonder.  As Panksepp himself stated many times, affects are deep and profound evolutionary gifts, ‘voices of the genes’ in his phrase, conserved value signals that provide our only real compasses keeping us on track in life, if we can understand, regulate and integrate them.  Although I feel a keen sense of loss at his passing, I am also very grateful for having encountered Jaak Panksepp in my life’s serendipitous travels, and for the many personal and intellectual treasures that have emerged from that relationship.

References

Chick, G., Yarnal, C., & Purrington, A. (2012). Play and Mate Preference: Testing the Signal Theory of Adult Playfulness. American Journal of Play, 4, 407–40.

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, reward, addiction, and withdrawal. Neuroscience & Biobehavioral Reviews35, 1847-1853.

Merker, B. (2007). Consciousness without a cerebral cortex: A challenge for neuroscience and medicine. Target article, commentaries and author’s response. The Behavioral and Brain Sciences, 30, 63-134.

Morgane, J. P., & Panksepp, J. (1981). Handbook of the Hypothalamus: Vol. 4: Part B. Behavioral Studies of the Hypothalamus. New York: Marcel Dekker, Inc.

Morgane, J. P., & Panksepp, J. (1980). Handbook of the Hypothalamus: Vol. 3: Part A. Behavioral Studies of the Hypothalamus. New York: Marcel Dekker, Inc.

Morgane, J. P., & Panksepp, J. (1980). Handbook of the Hypothalamus: Vol. 2: Physiology of the Hypothalamus. New York, NY: Marcel Dekker, Inc.

Morgane, J. P., & Panksepp, J. (1979). Handbook of the Hypothalamus: Vol. 1: Anatomy of the Hypothalamus. New York, NY: Marcel Dekker, Inc.

Panksepp, J. (2004). A Textbook of Biological Psychiatry. New York, NY: Wiley Press.

Panksepp, J., & Biven, L. (2012). The Archaeology of Mind: Neuroevolutionary Origins of Human Emotion. New York, NY: W. W. Norton & Company.

Panksepp, J. (1998). Affective Neuroscience: The Foundations of Human and Animal Emotions. New York, NY: Oxford University Press.

Panksepp, J (1996). Advances in Biological Psychiatry, Vol. 2. Greenwich, CT: JAI Press.

Panksepp, J (1995). Advances in Biological Psychiatry, Vol. 1. Greenwich, CT: JAI Press.

Russell, J. A., & Barrett, L. F. (1999). Core affect, prototypical emotional episodes, and other things called emotion: Dissecting the elephant. Journal of Personality and Social Psychology, 76, 805–819.

Watt, D. F. (2012). Theoretical challenges in the conceptualization of motivation in neuroscience: Implications for the bridging of neuroscience and psychoanalysis. In A. Fotopoulou, D. W. Pfaff, & M. A. Conway (Eds.), From the Couch to the Lab: Trends in Psychoanalysis (pp. 85-108). New York, NY: Oxford University Press.

Watt, D. F. (1998) Emotion and consciousness: Implications of affective neuroscience for ERTAS theories of consciousness. Internet Electronic Seminar for August–September 1998.  Association for the Scientific Study of Consciousness. http://server.phil.vt.edu/assc/esem.html.  Available from author by request.

Watt, D. F., & Panksepp, J. (2016). The Neurobiology and Psychology of Empathy.  New York, NY: Nova Science Publications.

Watt, D. F., & Pincus, D.  (2004). Neural substrates of consciousness: Implications for clinical psychiatry. In J. Panksepp (Ed.), Textbook of Biological Psychiatry (pp. 75-110). New York, NY: Wiley Press.

Zubieta, J. K., Ketter, T. A., Bueller, J. A., Xu, Y., Kilbourn, M. R., Young, E. A., et al. (2003). Regulation of human affective responses by anterior cingulate and limbic mu-opioid neurotransmission. Archives of General Psychiatry, 60, 1145–1153.

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