Language: English Spanish French. Cross-species affective neuroscience studies confirm that primary-process emotional feelings are organized within primitive subcortical regions of the brain that are anatomically, neurochemically, and functionally homologous in all mammals that have been studied. Emotional feelings affects are intrinsic values that inform animals how they are faring in the quest to survive. To understand why depression feels horrible, we must fathom the affective infrastructure of the mammalian brain. Advances in our understanding of the nature of primary-process emotional affects can promote the development of better preclinical models of psychiatric disorders and thereby also allow clinicians new and useful ways to understand the foundational aspects of their clients' problems.
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Abundant research into human clinical applications has supported the hypothesis that imbalances in these ancient primary emotional systems are strongly linked to psychiatric disorders such as depression. The present paper gives a concise overview of Panksepp's main ideas. It gives an historical overview of the development of Panksepp's AN thinking. It touches not only areas of neuroscience, but also shows how AN has been applied to other research fields such as personality psychology.
Finally, the present work gives a brief overview of the main ideas of AN. The scientist who coined the term affective neuroscience , Panksepp , , had the insight as a young clinical psychology student working in a mental hospital that understanding emotions was the key to developing more effective treatments for psychiatric hospital patients and all those suffering with psychopathology.
This insight led to his graduate school career change from clinical psychology into what we now call neuroscience. He had also realized that the level of understanding that was needed would require brain research that could not be conducted on human beings.
Hence, he began probing the neural constitution of emotions in the deep foundations of the mammalian brain. In the wake of the discovery in the early s of endogenous opioids in the mouse brain, Panksepp began working on another potential emotional behavior system in the brain. Scott, a senior colleague at Bowling Green State University BGSU , had studied the social behavior of dogs for many years Scott and Fuller, and was currently exploring separation distress vocalizations in puppies.
Meanwhile, Jaak had recognized similarities between opioid withdrawal in drug addicts and the social distress caused by broken relationships and had also noticed that opioid addicts frequently came from marginal family social backgrounds. Panksepp hypothesized that opioids might be related to mammalian separation distress calls, and a BGSU research group soon demonstrated that low doses of morphine would soothe the separation distress vocalizations in canine puppies Panksepp et al.
By , Panksepp was convinced that there were at least four biological brain-based emotional action systems Panksepp, , which at that time he labeled Expectancy, Rage, Fear, and Panic. The mapping of the seven primary emotional systems by means of electrical stimulation of the mammalian brain including pharmacological challenges and brain lesions represents the heart of what Panksepp named affective neuroscience.
Of note, Panksepp concluded that there was insufficient evidence to include Social Dominance as a primary emotion, and he considered it an acquired behavior [see further thoughts on this in van der Westhuizen and Solms ].
Panksepp did include a detailed treatment of the homeostatic affect HUNGER in Affective Neuroscience but did not consider it in the same category as emotional affects, which were more directly relevant to mammalian psychopathology and personality. The remainder of this review essay will outline key themes of affective neuroscience as developed in the research and writings of Jaak Panksepp.
Rather than continuing to review the development of his thinking throughout his scientific career, the focus here will be on selected affective neuroscience principles first featured in Affective Neuroscience but also elaborated in The Archaeology of Mind Panksepp and Biven, , The Emotional Foundations of Personality Davis and Panksepp, , as well as numerous theoretical review papers. All mammals are sentient beings meaning that it feels like something to be alive and dealing with the challenges in their worlds.
Primal emotions and their accompanying affects appear to have acquired the capacity to move animals to action in ways that promoted their survival. Evolution has endowed mammalian brains with at least these seven primary-process emotional action systems, which serve as survival guides.
These primary emotions arise from subcortical brain regions that are largely homologous, especially across mammals, with each emotion having a distinct brain anatomy, neuropharmacology, and physiology for details beyond the scope of this paper, see Panksepp, ; Panksepp and Biven, Jaak Panksepp felt that the key affective neuroscience question was the neural constitution of raw affects Panksepp et al. For a recent published obituary on Jaak Panksepp's life please see Davis and Montag Mother Nature aka evolution speaks to all mammals in the oldest language, the language of emotional affects.
The ancestral voices to use Ross Buck's phrase guide their choices as they navigate life. Yet, experiencing a primary affect does not necessarily mean all mammals can self-reflect on their emotional experiences. That capacity may be reserved for the more cortically endowed mammals. We know that animals experience primary affects because of empirical measures: They will work vigorously to sustain affective states by learning to turn on ESB evoking positively valenced emotions and correspondingly escape or avoid the negatively valenced emotions.
They will also demonstrate conditioned place preferences or aversions for situations where they have experienced such stimulation in the past Panksepp, Further, animals will emit conditioned positive and negative vocalizations in places where they have experienced positively or negatively valenced ESB Knutson et al.
However, we are unable to measure feelings affective qualia directly, not even in humans. Apart from experiencing primary-process emotional affects, what is much more difficult to study in the non-human mammalian world is Tulving's autonoetic consciousness characterized by the capacity of humans to experience affective nuances often reflecting human higher-order cognitions and language including having thoughts about thoughts.
This represents a limitation of cross-species affective neuroscience. However, subtler models of affective concepts such as pessimism in dogs Mendl et al. The primary-process emotions require no learning. It is not necessary to teach a child to become angry, fearful, or to panic after having lost sight of parents in a crowd. Nor do we need to teach children how to play. These evolved foundational tools for living are somehow automatically built into our heritage.
Indeed, as introduced above, the valenced affects associated with each of the primary emotions serve as endogenous rewards and punishments for behaviors that activate emotions. For example, receiving painful stings from hornets flying out of the nest you accidently disturbed fills you with fear, and without thinking about it you immediately react by running away from the menace.
Having reached a safe distance, you feel relief that you seem to be out of danger, and likely begin examining the tiny wounds, which are beginning to swell slightly as the pain intensifies, and you may begin to clarify at a safe distance the details of the hornet nest's appearance and location. This event will be forever embedded in your memory, and you will have learned to avoid repeating this experience by remaining more vigilant when outdoors walking through unfamiliar terrain.
Each primary-process emotional command system likely encompasses a separate reward or punishment system. These learning systems can be thought of as secondary-processes integrating new experiences into the primary framework allowing for previously neutral environmental stimuli to elicit the emotion and for novel reactions to become associated with such stimuli.
For an example of novel reactions to an emotional arousal, over the ages, humans when threatened have learned to reach for their swords, and more recently their pistols instead of clenching their fists. However, more evidence is needed regarding the extent to which these different emotional systems encompass different learning and memory parameters.
While deep brain stimulation DBS allows researchers to demonstrate that brain stimulation at specific sites evokes distinct emotional behaviors that are accompanied by corresponding affects, there remains the question of whether the emotional affects elicited at these sites are similarly distinct. We do know that rats can learn to discriminate between DBS in the hypothalamus and septal regions of the brain Stutz et al.
We also know that animals can distinguish between the emotional states induced by the addictive drugs morphine and cocaine Overton, We also know that DBS in humans at homologous brain sites seems to evoke homologous affects see Panksepp, for a review. However, much more research needs to be done before there is any clear assurance regarding the number of distinct primary affects or the role of electrical or pharmacological stimulants in generating those affects.
Primary affects are constituted at the subcortical level. There is ample evidence that primary-process emotional brain systems do not require the neocortex. In rats, decortication does not block the rewarding effects of subcortical ESB Huston and Borbely, , In humans, strong emotions decrease cortical activation Damasio et al.
Further support in humans for subcortical emotions without cortex is offered by Merker who has reviewed the case of hydranencephalic children who are born without a cerebral cortex. These children clearly show that appropriate emotional responses even in humans do not require the participation of the neocortex. As further evidence of an independent subcortical brain that can function without a neocortex, Merker also reviewed Penfield and Jasper in which brain surgery under local anesthesia was performed on conscious patients with a history of severe epileptic seizures.
Along these lines, Damasio et al. Plus the left and right insular cortices were entirely destroyed including the insula itself as well as the bilateral entorhinal cortex, hippocampus proper, and amygdala. In short, Patient B. Yet, all indications including the observations of strangers, the observations of the research team, psychological evaluations, and a structured questionnaire completed by his spouse comparing his emotional behavior before and after his disease were that Patient B.
The lack of importance of the cortex for emotional behavior and displays was supported by Whishaw's review of the experimental rat decortication literature. While there were differences between whether the surgery took place neonatally or closer to maturity, the neonatal group showed few deficits.
These subjects exhibited no interruption in post-surgery sucking and grew to maturity with near normal weights. They exhibited normal posture during face washing with no deficits in grooming. They were able to reproduce with six out of eight females being able to successfully raise their litters with normal cleaning, suckling, and caring for their pups.
Panksepp's group replicated the effect of neonatal decortication on rat juvenile play Panksepp et al. Measures of play vigor and tests of play solicitation behaviors did not detect differences, which suggested that play motivation was intact in the decorticate rats. They did observe a decrease in frequency of pinning and shorter pin duration.
However, additional control studies suggested that these changes were likely due to motor changes and reduced somatosensory sensitivity. In short, they found that the play of decorticate rats appeared normal. Indeed, 16 graduate students were asked to observe a pair of juvenile rats for 30 min, one of which had its cortex surgically removed neonatally and the other, a control subject, that only had received sham surgery, and to decide which one had been decorticated.
Most chose the control rat pup with an intact cortex Panksepp, By contrast this group reported that much smaller thalamic lesions had greater influence on play in rats. The theme of small subcortical lesions having dramatic influences on emotional behavior such as losing virtually all spontaneous activity after ablating the periaqueductal gray Bailey and Davis, was convincingly addressed by Fernandez de Molina and Hunsperger They showed that the rage responses of cats could be evoked by ESB along the basic subcortical mammalian RAGE system running from the periaqueductal gray PAG , at the lowest level up to the medial hypothalamus and on up to the medial amygdala at the highest level in decreasing levels of importance.
As such, they found that aggressive responses evoked by ESB of the amygdala were abolished by lesions at the level of the hypothalamus or PAG. Aggressive responses from the hypothalamus were dependent on the PAG but not on the amygdala. And, at the lowest level, aggressive responses evoked at the PAG level were not dependent on either of the higher two levels. Clearly, our primary-process emotions and their powerful affective messages are deeply embedded in our mammalian brains.
Humans and other mammals still experience these emotions without a neocortex, and the subcortical regions are organized in an evolutionary hierarchy of importance. Understanding these cross-species emotional systems may represent the greatest challenge to neuroscience.
But, is any function in neocortex genetically determined? It might seem to many that the visual cortex is a possible candidate. Yet, Sadato et al. Specifically, positron emission topography PET showed that when individuals who were blind from an early age were performing a braille task, the tactile processing that would normally occur in conventional somatosensory cortex had been shifted to areas in the occipital cortex that are normally assumed to process visual stimuli.
Another PET study provided evidence from blind subjects for the use of visual cortex for auditory processing. Weeks et al. While sighted and blind subjects both showed activity in the posterior parietal cortex, only blind subjects also showed activity in the occipital cortex, a cortical area that is normally associated with visual processing. Thus, cortical plasticity seems to allow blind individuals to develop enhanced tactile and auditory capabilities by redirecting neocortical regions typically thought of as visual processing regions to enhance other sensory functions.
Using ferrets a species born in an exceptionally immature stage , visual input was surgically redirected to auditory cortex shortly after being born, and as predicted the auditory cortex was developmentally programmed to process vision: A follow-up study using similar procedures with mice showed that rewired mice could learn visually-cued conditioned fear Newton et al.
Both sets of animals with visual input redirected to auditory cortex developed fine cortical visual abilities even though the neocortical processing was developmentally constructed rather than genetically dictated. That we should not expect to find evolved specializations in the neocortex was further supported by the report in Science that a single gene, called ARHGAP11B , was responsible for much of the massive expansion of the human neocortex.
The researchers further determined that this newly identified gene was found only in humans, Neanderthals, and Denisovans—another extinct hominid line in southern Siberia. The gene was not found in our closest living evolutionary relative, chimpanzees Florio et al. This finding makes it increasingly unlikely that any of our neocortical higher mental abilities represent evolutionary genetically-determined specializations like are found in subcortical brain regions.
Jaak Panksepp. Some investigators have argued that emotions, especially animal emotions, are illusory concepts outside the realm of scientific inquiry. However, with advances in neurobiology and neuroscience, researchers are demonstrating that this position is wrong as they move closer to a lasting understanding of the biology and psychology of emotion. In Affective Neuroscience, Jaak Panksepp provides the most up-to-date information about the brain-operating systems that organize the fundamental emotional tendencies of all mammals.
As a global organisation, we, like many others, recognize the significant threat posed by the coronavirus. During this time, we have made some of our learning resources freely accessible. Our distribution centres are open and orders can be placed online. Do be advised that shipments may be delayed due to extra safety precautions implemented at our centres and delays with local shipping carriers. Some investigators have argued that emotions, especially animal emotions, are illusory concepts outside the realm of scientific inquiry. However, with advances in neurobiology and neuroscience, researchers are demonstrating that this position is wrong as they move closer to a lasting understanding of the biology and psychology of emotion. In Affective Neuroscience , Jaak Panksepp provides the most up-to-date information about the brain-operating systems that organize the fundamental emotional tendencies of all mammals.
Jaak Panksepp: Pioneer of Affective Neuroscience