Jaak Panksepp and Lucy Biven, The Archaeology of Mind, Neuroevolutionary Origins of Human Emotions (2012)

At its most fundamental level, the neurosensory system of every animal, including humans, is a physiological system related to monitoring our entire physical system, whether the animal is awake or not, and, when awake (including the sleep-wake transition period), reacting to stimuli.  The latter is referred to as the arousal system.  In the case of vertebrates, four neurotransmitters (chemicals) --- acetylcholine, norepinephrine, dopamine, and serotonin --- stimulate different arousal systems originating in the brain stem, evolutionarily the oldest part of the vertebrate brain, and motivate certain behaviors such as seeking food, flight or fight behavior, and sexual activity.  There are certain emotional systems that are virtually at the core of the arousal system, at least during periods of wakefulness, and for several decades now, Jaak Panksepp has been researching and advocating that these emotional systems have their origins with animals evolutionarily older than humans, located in the older subcortical areas of mammalian brains and only later connected with the cortical areas. 

The Archaeology of Mind begins and ends with vertebrate animals, yet the evolutionary story is older, and to tell the story of what is missing from Panksepp's account I excerpt heavily from Steven Rose's The Future of the Brain, which outlines the evolution of the brain from unicellular organisms, to eukaryotes, to invertebrate animals and vertebrates.  This excerpting is important to a point I wish to make.  Rose says this:

"By the time that cells capable of metabolism and faithful replication, of symbiogenesis and competition appear, all the defining features of life have emerged:  the presence of a semi-permeable boundary separating self from non-self; the ability to metabolise -- that is, to extract energy from the environment so as to maintain this self --- and to self-repair, at least to a degree when damaged; and to reproduce copies of this self more or less faithfully.  All of these features require something we may term adaptability or behavior --- the capacity to respond to and act upon the environment in such a way as to enhance survival and replication.  At its simplest, this behavior requires neither brains nor  nervous systems, albeit a sophisticated set of chemical and structural features.  What it does require is the property that some would call a program:  at its most general way of describing both the individual chemical components of the cell and the kinetics of their interactions as the cell or living system persists through time. ***

"Built into this program must also be the possibility of modifying its expression, transiently or lastingly, in response to the changing contingencies of the external environment.  *** One way of conceiving of this capacity to vary a program is as an action plan, an 'internal representation' of the desired goal-- at its minimum, that of survival at least until replication is achieved.  I will be arguing that, in multicellular organisms, such action plans are ultimately what brains are about.

"Amongst the most basic forms of adaptive behavior drawing on such action plans is goal-directed movement-- of a unicell swimming towards food for instance. [Emphasis added].  Dip a thin capillary tube containing a solution of glucose into a drop of bacteria-rich liquid, and the bacteria collect around the mouth of the capillary from which the glucose diffuses--a phenomenon first noted as long ago as the nineteenth century.  Such simple responses engage a series of necessary steps.  First, the cell needs to be able to sense the food.  In the simplest case the food is a source of desirable chemicals --- perhaps sugars or amino acids-- although it may also be the metabolic waste products excreted by another organism.  Indeed the molecule does not have to be edible itself provided it can indicate the presence of other molecules that can be metabolized-- that is, it acts as a signal.  *** But signals are only signals if there is recipient who can interpret the message they bear.  Cell membranes are studded with proteins whose structure is adapted to enable them to trap and bind specific signaling molecules floating past them, and hence read their message.  This chemical detection system is the most basic of all sensory mechanisms.

"Interpreting the message --- using it to develop a plan of action -- should make it possible for the cell to determine the direction of the gradient and finally to move up it to the source.  Moving towards a specific chemical source --- chemotaxis --- requires that the cell possess some sort of direction indicator or compass.  One way of creating such a compass, employed by bacteria, is to swim in a jerky trajectory, enabling the cell to interpret the gradient by comparing the concentration of the attractant chemical at any moment with that a moment before.***"

If Jaak Panksepp were reading this passage he would certainly connect it to his research of emotions in animal brains.  It describes the precursor to what Panksepp regards as the most central emotional system in mammals: the SEEKING system (see below).  Rose continues:

"The molecules trapped by the receptor on the surface membrane serve as signals, but very weak ones.  To produce as dramatic a cellular response as turning and moving in the right direction requires that signals are highly amplified.  The mechanism by which this is carried out, even in the seemingly simplest of unicells turns out to be the basis on which the entire complex apparatus of nervous systems and brains is subsequently built.  The receptors are large proteins, oriented across the lipid membrane, with regions sticking out into the external environment, and also 'tails' which reach into the interior of the cell (the cytoplasm).  When the signal receptor binds to the receptor protein its effect is to force a change -- a twist, if you like -- in the complex shape of the receptor.  ***

"One way of speaking of this process, favoured by neurologist Antonio Damasio, even in so limited an animal as Paramecium, is as 'expressing an emotion.'   Emotion for Damasio, is a fundamental aspect of existing and a major driver of evolution.

"*** With multicellularity, 'behaviour' becomes a property of the organism as a whole, to which 'needs' of individual cells are subordinated.  The internal representation which makes possible the action plan for organism can be delegated to specific cell ensembles.  This requires new modes of communication to be developed.  Where previously there were only two classes of signals -- those arriving from the external environment to the cell surface, and those internal to the cell --- there are now three.  Signals from the external environment are still registered by sensory cells on the surface and are transmuted by molecular cascades with them, but now the response to those cascades requires that further messages be sent from the sensory cells to other regions of the body, including of course the contractile cells.  Sometimes the sensory cells make contact with intermediaries whose task it is to synthesise and secrete the necessary 'messenger molecules.'  [Emphasis added]. The messengers can then be distributed through the body either by way of a circulatory system or by diffusion through the extracellular space between the body cells, and are detected as before by specialized receptor proteins on the surface membranes of their targets.  When molecules that served such messenger functions were first identified in mammals, they were given the generic name of hormones.  It was only later, and to some surprise, that it was discovered that many of the same molecules also serve as intercellular signals in very early multicellular organisms, another powerful example of evolutionary conservation.***

"It is easy to imagine a sequence whereby neurons evolved from secretory cells.  Instead of discharging their contents generally into the surrounding space and circulatory system, the secretory cells could have put out feelers (called 'processes') enabling them to make direct contact with their targets so as to signal rapidly to them and them alone.  Messages could be conveyed between the two either electrically or chemically --- by a depolarizing wave or by secreting a messenger molecule across the membrane at the point where the two cells touch.  In fact, both phenomena are know to occur.

"The first step towards such nervous systems can be seen among the large group of Coelenterates, believed to be amongst the earliest true multicellular animals.  The best known is perhaps the Hydra, a tiny creature that sits at the bottom of streams attached to rocks or water plants, waving its tentacles above its mouth.   When a potential source of food brushes past its tentacles, the Hydra shoots out poisonous threads, collects the paralysed victim and thrusts it into its mouth.  *** A well fed Hydra is quiescent; when hungry it waves its tentacles vigorously, or moves its location by repeatedly turning head-over-heals, seeking food-rich or oxygen-rich environments (once again, Damasio would regard these acts 'expressing emotions').***

"What distinguishes a fully-fledged nervous system --- our own for instance --- is a one-way flow of information through the system, from dendrites to axon, from sensory cell to effector.  Of course this is mediated via all the feedback loops, but none the less there is a directionality to it that the Hydra's does not possess.

"Whereas the Hydra's neurons are scattered throughout the body, the next crucial step was to concentrate them within an organized system. *** C. elegans  has a head and tail end, and as it is more important for it to know where it is going than where it has been, many of its sensory cells are clustered at its head end.  From these, nerve connections run to clusters of interneurons, pack into groups (ganglia) with short interconnecting processes between the cells within the group and longer nerve tracts leading out along its gut and ultimately to the effectors:  contractile, egg- and sperm producing cells.  These neurons use many of the neurotransmitters that are found in mammalian brains (notably the amino acid glutamate), an indication of how far back in evolutionary terms these molecules were adapted for signaling functions.***

"The evolutionary track I have been mapping," writes Rose, "has led from proto-cells to faithfully replicating eukaryotes capable of responding adaptively to patchy environments, from single-celled eukaryotes to multicellular animals with internal signaling systems, and from these to fully-fledged nervous systems capable not merely constructing action plans, but of modifying those plans, at least temporarily, in response to environmental contingencies.  But we haven't yet arrived at brains.  This must have been the next step along the evolutionary path that led to humans.  Concentrating neurons in ganglia is a way of enhancing their interactions and hence their collective power to analyze and respond to incoming stimuli.  Locating them at the front end of the organism is the beginning of establishing not merely a nervous system but a brain, though head ganglia or brains only slowly begin to exert their primacy over the other ganglia distributed through the body.*** [Turning to invertebrates] although insect (arthropod) and molluscan neurons are pretty similar to human neurons, and the biochemical motors that drive the system -- their electrically excitable membranes and the neurotransmitters --- work in the same way, the organization of the system is entirely different.  In molluscs and arthropods the central ganglion --- the nearest any amongst these huge numbers of species have to a brain --- and the principal connecting pathways between it and other ganglia lie arranged in a ring around their guts.  This is a device that can be seen even in earthworms, and it imposes a fundamental design limitation on the complexity of the nervous system.***

"The development of large brains required two major changes in the construction of nervous systems:  the separation of the nerves themselves from the gut, and the concentration of nervous power.  It also required the first step towards the development of a bony skeleton.  Amphioxus,  small sea-floor fish, is an example.  Less behaviourally sophisticated than octopus or bee, it has a flexible rod of cartilage, a notochord, running down its back --- the forerunner of the spinal column --- with the merit of providing a bracing device against which muscles can pull.  More relevantly for the present argument is that eh major nerves and central ganglion lie in a continuous tube running the length of the creature's body, thus disentangling them from the gut and giving space for growth."

We have not even discussed Panksepp's research yet, but there is much here in Steven Rose's account of the evolutionary development of the animal nervous system that indicates the system of neurotransmitters and specialized receptors found in vertebrates long preceded the development of the brain stem in vertebrates.  And there is a suggestion by Steven Rose that this system was capable of "expressing emotions," although probably not in the same sense that Panksepp intends.  But it would be fair to say that human emotional systems and those of other mammals not only have their origins in vertebrate animals older than humans, but in the earliest forms of life on earth.  This is an anthropomorphic view of human emotions as described by Frans DeWaal in The Ape and The Sushi Master (see June 17, 2010 post).  To be sure, Panksepp is careful to admonish in his discussion of similarities between the neurological systems of humans and other mammals that "similar does not mean the same."  There are similar structures and similar transmitters and receptors in the brain, but their location within the brain may be slightly different or even vastly different, and those differences may result in small or even large differences between humans and other mammals.  But in identifying these similarities, Panksepp observes, as the book's subtitle hints, the neuroevolutionary origins of human emotions.  Panksepp decries the history of human psychological research that declined to recognize emotions in animals.  There is considerable research available today that rebuts that notion.

Panksepp discusses several emotional systems, but central to nearly all of them is what he has labeled the SEEKING system.  And in beginning this discussion, we can think back to Steven Rose's reference to the "goal-directed movement-- of a unicell swimming towards food for instance." 

Panksepp is controversial within the neuropsychiatric community, challenging some of the dogmas of neuroscience and human psychotherapy.  One of the dogmas is reflected in this statement from Rita Carter's Mapping the Mind (see November 6, 2011 post):  "A huge volume of evidence suggests that consciousness emerges from the activity of the cerebral cortex that the particular type of consciousness that includes the sense of self requires activation in the frontal lobes.  Ask yourself this:  Where, precisely, do I feel that "I" am centered?  If you are like most people, you will point to a position just above the bridge of your nose.  It is right behind here that you will find the prefrontal cortex --- the area of the frontal lobe most closely associated with the generation of consciousness.  This region is also responsible for our conscious perception of emotion and our ability to attend and focus.  Most important of all, it endows the world with meaning and our lives with a sense of purpose.  The symptoms of schizophrenia, depression, mania and Attention Deficit Disorder are mainly due to frontal lobe disorder."  Carter's sentiment reflects a view that leads psychotherapists to focus on treating the executive, regulatory capacity of the human brain in the frontal cortex in order to overcome these disorders.  While Panksepp does not dismiss the role of the prefrontal cortex in the conscious life of humans, he does disagree with the directionality implicit in this statement:  for Panksepp, like Antonio Damasio (see April 8, 2011 post) "the generation of consciousness" begins with the evolutionarily older parts of the brain --- in the midbrain, where neurotransmitters are generated  --- as well as the limbic system, which together are at the foundation of the seven emotional systems he describes in Archaeology of Mind.   It is here that the "core self" of consciousness emerges, or as Panksepp calls it, the core affective self.  The symptoms of certain mental disorders, Panksepp believes, are not "mainly due to frontal lobe disorder" but may have more to do with the imbalanced (excessive or diminished) production of specific chemicals in the brain in the more ancient parts of the brain.  And as the previous post suggests, epigenetics provide some explanation in the case of stress disorders caused by early childhood abuse leading to excessive production of cortisol that overwhelms the ability of the limbic system to restore calm. 

The seven emotional systems described by Panksepp (and he does not rule out that there may be more) are these:

The Seeking System.  This does not immediately sound like it describes an emotional system, but clearly Panksepp is correct in characterizing the Seeking System.  This is the system "that allows animals to search for, find and acquire resources that are needed for survival.  Arousal of this Seeking System produces all kinds of approach behaviors, but it also feels good in a special way.  It is not the kind of pleasure we experience when eating a fine meal, or the satisfaction we feel afterwards.  Rather it provides the kind of excited, euphoric anticipation that occurs when we look forward to eating that meal . . . the anticipation of sex . . . the thrill of exploration."  Panksepp refers to the Seeking System as the primary process emotional powers that makes animals into active agents in their environments.  "Among animals in the wild, it is easy to see the Seeking system in action.  Resources are not readily available and animals must persistently seek them out in order to survive.  They must hunt or forage for food and search for water, find twigs or dig holes to fashion sheltering nests.  The Seeking system urges them to nurture their young, to search for a sexual partner, and when animals live in social communities, to also find nonsexual companions, forming friendships and social alliances. . . Although this system vigorously responds to homeostatic needs, to emotional urges and to enticing temptations, it operates more or less continuously in the background, albeit at much lower levels when people and animals are not in any particular need of resources or troubled by problems that urgently require solutions.  This system keeps animals constantly exploring their environments so they can remember where resources are."  Importantly, in Panksepp's view, it is the Seeking System that is the motivator behind the intellectual pursuits of the neocortex:  "the neocortex does not provide its own motivation; the neocortex is activated by subcortical emotional systems . . . the neocortex is the servant of our emotional systems."  It is the Seeking System that urges architects, artists, writers, politicians, and scientists to discover new and better ways to solve problems and express themselves.  It "energizes all human creativity."  Seeking arousal "is an anticipatory gift of nature that provides seemingly infinite opportunities for learning; with the developmental/epigenetic emergence of higher mental processes, it gradually fine-tunes reasonable expectations, working hypotheses, as in the conduct of science."  It is intimately connected with learning, which Panksepp describes as an "automatic, unconscious process that enhances are natural proclivity to engage with the world in ever more subtle ways as our minds mature."  In contrast, affect (behavioral outcomes connected to arousal of instinctual emotional systems) is never unconscious; it is felt.

Chemically, the Seeking System is understood to be aroused by dopamine transmitters, but glutamate, which functions in learning and memory, and neuropeptides such as orexin and neurotensin are understood to activate the Seeking System while dynorphin is believed to deactivate it.  The neurons for these transmitters are found in the midbrain:  anatomically, ventral tegmental area, the medial forebrain bundle, the lateral hypothalamus, the nucleus accumbens, and then running to the medial prefrontal cortex via the mesolimbic and mesocortical dopamine pathways.  "In all mammals," notes Panksepp, "the nucleus accumbens interacts with the medial frontal cortex to promote simple appetitive learning (and addictions).  Because the Seeking System energizes the frontal neocortical regions, especially the medial zones that focus on immediate emotional needs, we are able to devise strategies to obtain rewards and escape sanctions (pain) and other pitfalls.  We remember particularly pleasurable experiences and the possibility of addiction is created.  Dopamine transmitters are associated with drugs of abuse, and when they are overly excited there can be negative consequences from addiction.  On the other hand, when the Seeking System is underactive, depressive feelings can emerge. Humans differ from other animals here in one important respect; the dopamine pathways that energize the cortex are linked not only to the frontal cortex but to other sensory-perceptual cortices in the back of the brain.

The Rage System.  The Rage System needs little explanation:  the foundation of anger and aggression.  What it is not deserves some explanation:  it probably has little to do with war among societies (group aggression), nor is it about predatory aggression such as seeking food.  In contrast to the Seeking System, which is largely a "positive" emotion, the Rage System produces unpleasant affects.  The Rage System is connected to dominance systems in species.  The Rage System runs from the medial areas of the amygdala to the medial hypothalamus to areas of the periaqueductal gray (PAG). As with the Seeking System (and all the other emotional systems Panksepp describes), these are the ancient areas of the brain.  The chemicals that can promote rage include testosterone (known to promote physical aggression in males to a greater extent than females), Substance P (important to pain perception), norepinephrine, glutamate, acetylcholine, and nitric oxide synthases.  The Rage System can be controlled by chemical inhibitors such as gamma-aminobutyric acid (GABA) and oxytocin. 

The Fear System.  Similarly, the Fear System needs little explanation.  Like the Rage System, it is not a positive emotion; it produces anxiety, stimulates flight, fight or freezing.  The Fear System operates between the PAG and the amygdala and it is aroused by external and internal stimuli, notably pain, but some responses appear to be innate caused by hard-wired sensory inputs.  Panksepp mentions rats fear of open spaces, sudden movements and loud noises as example innate fear responses.  But fear is connected to memory as well, and memory plays a significant role in conditioning fear responses.  On memory, Panksepp explains, that learning and memory are automatic and involuntary responses (mediated by unconscious mechanisms of the brain), which in their most lasting forms are commonly tethered to emotional arousal. Emotional arousal is a necessary condition for the creation of fear-learning memories.

The Lust System. The Lust System drives basic mammalian physical impulses (sexual affects) on the one hand and social emotions on the other, which can be both positive and negative.  It can drive anti-social behavior (rape, stalking) as well as building families and promoting other forms of well-being.  In the male brain the center of primary sexual urges is in the medial regions of the anterior hypothalamus, (although Panksepp notes that "the precise brain location varies from one species to another).  Testosterone stimulates pleasure in the male, which activates neuropeptides such as vasopressin and promotes sexual ardor, courtship, intermale aggression and possibly jealousy.  Testosterone also activates nitric oxide in the brain, which promotes heightened sexual eagerness.    In females, estrogen and progesterone (the estrus cycle) controls sexual arousal, but adrenal testosterone plays a role in sexual receptivity.  The Lust System, Panksepp says, "recruits" the Seeking System "dopamine-fueled search for companionship.

The Care System.  The Care System is not universal in the animal kingdom, but nearly all mammals and birds exhibit maternal care for their young.  In fish, the job of tending to a nest of eggs is left to fathers, and the brain circuits that drive this behavior Panksepp calls the Care System.  Panksepp notes that researchers learned of the existence of the Care System in mammals when they discovered that blood transfusions from postpartum female rats to virgin rats would lead to maternal behavior in the virgin rats, including nest building, hovering over young, and gathering the young who strayed from the nest.  Panksepp concedes that researchers still do not which chemicals in the transferred blood interact in the brains of virgin rats to cause these behaviors, but given similarities between the urge to provide Care and the urges underlying the Seeking System, brain arousal from dopamine in conjunction with opioids, as well as oxytocin and prolactin are likely involved.  Panksepp hypothesizes that the evolution of the Care System might be traced back to chemicals found in the Lust circuits of reptiles, such as vasotocin, which has a calming effect and promotes nurturant moods in some birds, and neuropeptides like mesotocin that may have evolved in vasopressin and oxytocin, which is recognized as a key maternal chemical.  The maternal (and paternal) nurturing behavior must be recognized as a critical factor in the development of social brain systems.  Research shows that both oxytocin and vasopressin strengthen social memories and are believed to be promote social bonds among mammals.  (See July 16, 2010 post).

Research on the Care System in rats also reveals evidence of epigenetic changes leading to more prosocial behavior.  Female rats lick their pups during early development and this has been shown to influence the emotional abilities of young rats later in life.  Abundantly licked rats grow up to be less anxious, more resistant to stress, and more capable of exhibiting learning and other adaptive behavior later in life.  These adult rats have diminished stress hormones (corticotrophin-releasing factor (CRF)) and adrenocorticotrophic hormone (ACTH), more GABA receptor cites, which promotes reduced anxiety, and more receptors for glutamate and norepinephrine, which facilitate learning.  Emotionally, these animals are less anxious, showing more activity and fearlessness, and better learning and performance in a variety of fear-inducing situations.  This research could have been cited by Nessa Carey in The Epigenetic Revolution.  (See April 28, 2013 post). 

The Panic/Grief System.  Panic and grief intuitively seem like strange bedfellows but the common emotional/behavioral link in this "system" is separation anxiety, something that is seen across a number of species.  Grief connotes a sadness that arises from social loss; panic connotes a separation from a secure or stable environment.  Immediately, one can conjure linkages between what Panksepp labels the Panic/Grief System and the Care System, the Fear System.  The Panic System is seen in early childhood development over anxiety in separation of mother and child ("Born to Cry" is the title of this chapter), but it has also been found to be less active in adults.  The Panic/Grief circuits are found in several of the same subcortical areas identified with other systems, including the PAG and surrounding subcortical regions including the dorsomedial thalamus, the ventral septial area, the dorsal preoptic area and the bed nucleus of stria terminalis.  Previously identified stress neuropeptides such as CRF and  ACTH, and glutamate (an excitatory neurotransmitter associated with every emotional response) arouse the Grief System.  Imbalances in the Grief System are a key factor in a variety of emotional disorders because so much mental illness, Panksepp notes, is rooted in the incapacity to enjoy the security of warm interpersonal attachments.  Panic attacks, depression, autism, and a variety of other social phobias are part of the Grief pathologies.  The identification of neuropeptides that actually diminish separation distress and mediate the Care System, such as oxytocin and prolactin,and the stimulation of mu-opioid receptors in the brain may have role in treatments of these disorders.

The Play System.  Finally, but not least, something that one might not think of as an emotional system, but Panksepp clearly documents that it is, particularly in mammals: the Play System.  "Physical playfulness is a birthright of every young mammal and perhaps of many other animals as well. . . It is now certain that a genetically determined Play network that mediates positive affect exists in mammalian brains, although many details remain to be worked out."  The Play System is likewise concentrated in subcortical brain regions, intimately linked to the Seeking System: the urge to play is like a type of hunger, and is not necessarily a social need, although it is linked to social emotional systems. Play is linked to the capacity to laugh, a positive emotional affect. Laughter is not merely found in humans, but also noises made by rats, chirping of birds.  Laughter is stimulated early in children, including mimicry.  Like the Seeking System, dopamine, which is engaged during activity that entails considerable positive anticipation and euphoria, is believed to fuel the Play System because it is aroused (correlated) during play.  Play activates sensory inputs, such as touch, which go directly to older midline regions of the brain such as the parafascicular complex and the posterior dorsomedial thalamic regions. 

In the foregoing, I have catalogued for each of Panksepp's seven emotional systems of the brain the suspected chemistries and at the outset I tried to demonstrate that research documents the ancient role of chemicals in the neurological systems of species and their potential link to the development of emotional system.  My objective in this outline is to highlight a point in a previous post about social emotions, including moral emotions.  In his book Moral Origins, Christopher Boehm concludes by saying that in a few generations we "may have identified some of the genetic mechanisms that help us to behave egoistically, nepotistically, and altruistically, along with others that make for sympathetic generosity, domination and submission, and a variety of other socially significant behaviors that are relevant to morality, including our shame responses."  The earlier post (see November 21, 2012 post) observed that "Boehm may well be right that we will identify the genetic mechanisms behind moral and immoral behavior in a few generations, but the roadmap of investigation is already before us and it begins with emotions.  I say this for two reasons:  first, if anything, genes code for our body chemistry; genes may or may not code for specific behavior (moral or otherwise), although I doubt it (see November 30, 2009 post).  But emotions are driven by electro-chemical actions and reactions in our various body systems and ultimately the neurological system leading to our brains, and genes do code for these electro-chemical actions and reactions and genes code for our brain and other body organs.  If we want to understand the genetic basis for moral and immoral behavior we will look for the genes tied to these body systems and the chemistry that drives emotions."   Panksepp's aggregation of the research on these primary process emotions is a good peek into the links between genes, chemistries, and anatomical structures related to emotions.  In addition to linking genes with the chemicals and brain structures that drive these emotional systems, the inquiry contemplated by Boehm would presumably link these seven emotional systems to other more complex emotional systems not considered "primary process" systems,  including the social emotions discussed in the November 21, 2012 post such as embarrassment, shame, guilt, contempt, indignation, sympathy, compassion, awe, gratitude, and pride. 

One cannot help read The Archaeology of Mind without feeling that Panksepp believes he has been walking in the wilderness of neuroscientific research that treats emotional systems as fundamental, more fundamental than research of the neocortex.  While he now believes that Antonio Damasio has joined his crusade with the publication of Self Comes to Mind (see April 8, 2011 post), in which Damasio gave a tip of the hat to Panksepp's research, Panksepp is skeptical of Damasio's earlier somatic marker hypothesis and the assertion that core consciousness (a higher order mapping process outside the subcortical regions) generates inner emotional feelings of what is happening by synthesizing information from maps abut the body and about the environment. As stated earlier, it is the subcortical emotional system that energizes the neocortex, says Panksepp, not the other way around.  Fundamentally, Panksepp believes that mental and emotional disorders go hand in hand and are best understood as a chemical problem, and when understood in that leads to two important conclusions:  (1) that chemistry will have a key role in providing treatment, and (2) it will cause psychotherapists to recognize that treatment must deal with the emotional aspects of the older subcortical parts of the brain.  For Panksepp, the key question for all neuroscientists and biological psychiatrists is this:  "How are raw affective experiences created in the brain?"  The answer he believes will clarify the foundational nature of experience in general as well as affective disturbances.  For example, Panksepp writes, for depression he would ask:  Why does depression feel so bad?  Why does depression hurt?  Why is it so psychologically painful?  What does it mean to experience social pain?  Few neuroscientists have been willing to ask these questions.

One cannot conclude a statement about Panksepp's research without noting what he neither ignores, but nonetheless does not dwell on: the role of the cortical areas of the brain in human consciousness.  When he does acknowledge higher order BrainMind structures, he says this:  "Although arousals of the primary process emotional networks of mammalian brains are intensely experienced by humans and other animals, it is especially important to recognize that the secondary processes of the BrainMind, the basic forms of learning, memory, and habit formation are among the most unconscious 'mental' processes of them all.  Once we understand this, then many of the bizarre and faulty views from psychology's past may be rectified.  For instance, 'free will' is not a figment of our imagination as too many scientists are ready to claim these days.  Free will is a higher tertiary-level neurocognitive function that we use on a regular basis (and quite effectively when we are not too emotionally aroused) for future planning actions.  This is brought out beautifully in the concept of 'autonomy' and 'self-determination' as developed by Ryan and Deci (2006).  However, we cannot readily will ourselves out of underlying emotional turmoil that has been created through the consolidation of maladaptive affective patterns at primary and secondary levels of BrainMind organization.  At primary-process levels of emotional processing there is no free will, there is no 'controlled cognitions.' Neither do the automatic secondary processes of learning and memory functions, that are molded by our wild animal passions developmentally, exhibit free will.  That can only emerge from well-sculpted, deeply reflective, cognitive attitudes."  He adds, "It is surely our vast cerebral 'thinking cap' --- our extensive cortico-cognitive apparatus --- that distinguishes us mentally from our animal ancestors.  That adds layers of complexity that cannot be readily addressed with animal models."  Michael Gazzaniga would agree.  (See September 27, 2009 post). But "language," Panksepp says," our most unique cerebral skill, "emerges through emotional guidance.  Through language, however, we can uniquely study the extended tertiary-process cognitive affective consciousness of humans.  And this is why there continues to be enormous growth in descriptive (ie. nonneuroscientific) emotion studies in psychology (Davidson et all., 2003)." 

And with that paragraph, I pull the next book off of The Bookshelf. 

Christopher Boehm, Moral Origins, The Evolution of Virtue, Altruism, and Shame (2012)

Anthropologist Christopher Boehm is interested in this question:  Why did humans evolve a conscience?  Before exploring this question and what Boehm believes may be the answer to his question, I want to develop some facts and opinions of others on issues that I believe bear on this question, some of which Boehm does discuss and some of which he does not discuss in his book Moral Origins.

The first issue I want to address is the role of emotions, and in particular the social emotions in the origins of human morality.  Boehm concludes Moral Origins by saying that in a few generations we "may have identified some of the genetic mechanisms that help us to behave egoistically, nepotistically, and altruistically, along with others that make for sympathetic generosity, domination and submission, and a variety of other socially significant behaviors that are relevant to morality, including our shame responses."  Boehm may well be right that we will identify the genetic mechanisms behind moral and immoral behavior in a few generations, but the roadmap of investigation is already before us and it begins with emotions.  I say this for two reasons:  first, if anything, genes code for our body chemistry; genes may or may not code for specific behavior (moral or otherwise), although I doubt it (see November 30, 2009 post).  But emotions are driven by electro-chemical actions and reactions in our various body systems and ultimately the neurological system leading to our brains, and genes do code for these electro-chemical actions and reactions and genes code for our brain and other body organs.  If we want to understand the genetic basis for moral and immoral behavior we will look for the genes tied to these body systems and the chemistry that drives emotions.  The second reason is not biological, but an observation that philosophers have made since the 18th century: that emotions, and in particular certain social emotions shape our "moral sentiments." (See April 8, 2011 post).  I am thinking particularly of David Hume (see February 27, 2011 post)  and Adam Smith (see January 11, 2011 post).  Research is beginning to show that emotions trigger moral behavior. 

The second issue is the growth of the brain in the evolution of hominids leading to homo sapiens.  Antonio Damasio's works confirm that the evolution of consciousness in humans is tied to the growth and development of the brain, particularly the cortical regions, and without "extended" consciousness apparently enabled by the larger human brain we very likely have no conscience.  (See April 8, 2011 post and October 25, 2011 post).  Damasio hits a theme in these two paragraphs from his book, Looking for Spinoza, that I quote at length below, and which I don't believe Boehm would disagree with:

"The construction of what we call ethics in humans may have begun as part of an overall program of bioregulation.  The embryo of ethical behaviors would have been another step in a progression that includes all the nonconscious, automated mechanisms that provide metabolic regulation; drives and motivations; emotions of diverse kinds; and feelings.   Most importantly, the situations that evoke these emotions and feelings call for solutions that include cooperation.  It is not difficult to imagine the emergence of justice and honor out of the practices of cooperation.  Yet another layer of social emotions, expressed in the form of dominant or submissive behaviors within the group, would have played an important role in the active give and take that define cooperation.

"It is reasonable to believe that humans equipped with this repertoire of emotions and whose personality traits include cooperative strategies would be more likely to survive longer and leave more descendants.  That would have been the way to establish a genomic basis for brains capable of producing cooperative behavior.  This is not to suggest that there is a gene for cooperative behavior, let alone ethical behavior in general.  All that would be necessary would be a consistent presence of the many genes likely to endow brains with certain regions of circuitry and with the attendant wiring --- for example, regions such as the ventromedial frontal lobe that can interrelate certain categories of perceived events with certain emotional feeling responses.  In other words, some genes working in concert would promote the construction of certain brain components, and the regular operation of those components, which, in turn, given the appropriate environmental exposures, would make certain kinds of cognitive strategy and behavior more probable under certain circumstances.  In essence, evolution would have endowed brains with the apparatus necessary to recognize certain cognitive configurations and trigger certain emotions related to the management of the problems or opportunities posed by those configurations.  The fine tuning of that remarkable apparatus would depend on the history and habitat of the developing organism." (emphasis added).

The social emotions: Shame, Sympathy and Empathy. The so-called "social emotions" --- embarrassment, guilt, shame, and pride --- are linked to moral behavior in humans, and therefore no discussion of moral origins can ignore social emotions. "It is highly probable that the availability of such social emotions has played a role in the development of complex cultural mechanisms of social regulation," writes Antonio Damasio in Looking for Spinoza. "It is also apparent that some social emotional reactions are elicited in human social situations without the stimulus for the reaction being apparent to the reactor and to observers." Damasio believes that the social emotions are buried deep in the organism's brain, innate rather than taught. (See April 8, 2011 post). Researchers have concluded that social emotions are not unique to humans. As Damasio observed in Looking for Spinoza, "Because the term 'social' inevitably conjures up the notion of human society and of culture, it is important to note that social emotions are by no means confined to humans. Look around and you will find examples of social emotions in chimpanzees, baboons, and plain monkeys; in dolphins and lions; in wolves; and, of course, in your dog and cat. The examples abound --- the proud ambulations of a dominant monkey; the literally regal deportment of a dominant great ape or wolf that commands the respect of the group; the humiliated behavior of the animal that does not dominate an must yield space and precedence at mealtimes; the sympathy an elephant shows toward another that is injured and ailing; or the embarrassment the dog shows after doing what he should not." (See also June 17, 2010 post).

Psychologists distinguish between basic emotions and social emotions.  Moral emotions are treated as a subset of social emotions.  Which emotions constitute the "basic" emotions is a matter of debate, but they at least include happiness, sadness, disgust, anger, and fear --- emotions that require an awareness of one's own somatic state.  These are ancient emotions that are tied to survival, driving us away from harm and directing us toward some reward.  Social emotions require the existence of a group, a theory of mind (TOM) and an awareness of the mental states of others.  The social emotions include embarrassment, guilt, shame, contempt, indignation, sympathy, compassion, gratitude and pride.  Following Jonathan Haidt's paper on The Moral Emotions, Antonio Damasio in Looking for Spinoza has described the basic emotions underlying the social emotions and what he calls the emotionally competent stimulus (ECS) for and consequences of the social emotions as follows:

Social emotion	ECS	Consequences	Basic emotion
Embarrassment, shame, guilt	Weakness or failure or violation in individual’s own behavior	Prevent or avoid punishment by others; restore balance to self or group	Fear, sadness
Contempt, indignation	Other individual’s violation of norms	Punishment or violation, enforcing of social norms	Disgust, anger
Sympathy, compassion	Another individual suffering, in need	Comfort, restoration of balance in other or group	Attachment, sadness
Awe, gratitude, pride	Recognition in others or self of contribution to cooperation	Reward for cooperation, reinforcing tendency to cooperate	Happiness

These social emotions, the ECS, and consequences become central to Boehm's discussion of moral origins.  Damasio's point:  "In a society deprived of such emotions and feelings, there would have been no spontaneous exhibition of the innate social response that foreshadow a simple ethical system---no budding altruism, no kindness when kindness is due, no censure when censure is appropriate, no automatic sense of one's own failings.  In he absence of the feelings or such emotions, humans would not have engaged in a negotiation aimed at finding solutions for problems faced by the group, e.g., identification and sharing of food resources, defense against threats or disputes among its members."  Boehm at least implicitly recognizes this in the subtitle of his book --- The Evolution of Virtue, Altruism and Shame --- but the genetic foundations of our emotions is not clearly called out.

Shame is particularly called out by Boehm as an emotion central to moral origins in humans, but I will address that more specifically later.  Boehm repeatedly treats sympathy and empathy as synonyms; they are not and I want to comment on that here.  But in discussing moral origins, Boehm is correct to emphasize "sympathy."  Sympathy, as Damasio's categories above suggest, is proactive: the consequence is for one person to provide comfort to another person.  As Frans DeWaal says in The Age of Empathy (see November 9, 2010 post), "Sympathy, in contrast, reflects concern about the other and a desire to improve the other's situation."  In contrast, "Empathy is the process by which we gather information about someone else."  Empathy is more of a feeling rather than an emotion.  As Jonathan Haidt writes in his article The Moral Emotions,  "Empathy is not an emotion at all; it is a tendency to feel whatever another person is feeling, including happiness, anger, or boredom."  With respect to empathy, we can recall the discussion of mirror neurons in monkeys, apes, and humans, that would facilitate these feelings.  (See October 25, 2011 post and July 16, 2010 post)  "Empathy is easily aroused," says DeWaal, but "sympathy is anything but automatic."  Sympathy is common in both humans, chimpanzees and bonobos who can be consoling.  Sympathy, however, would not occur without a capacity for empathy. 

This brings us to shame, embarrassment and guilt. Shame attracted the attention of Charles Darwin. He noted that humans universally blushed with shame that appeared to be associated with a moral conscience. This behavior is seen only in humans. By definition, shame is provoked when an individual recognizes that his or her own conduct (or perhaps the conduct of another group member, for example shame due to the conduct of one's own child or spouse) has transgressed the social rules or expectations of others.  Shame implicitly, if not explicitly, acknowledges the applicability of those social rules to one's own conduct.  It is similar to guilt, but guilt could apply to an individual's own rules and not merely social rules.  Boehm says that during his research in the African forest "the apes never appeared to me as though they were upset over their own behavior, let alone ashamed of it or remorseful. I did notice that between individuals there were postures and gestures that seemed to ask for or grant forgiveness, and in fact chimpanzees often make up after conflicts. . . However, this seems to be aimed merely at reducing tension or restoring positive relations, so reading a morally-based element of remorse into such behavior would be patently anthropocentric. Nothing I observed ever convinced me that there was something like morally based self-recrimination in the wild, for aggressors never appeared to be troubled by their actions afterward." How did humans come to blush with shame so predictably, asks Boehm? Much of the answer has to lie in biology, he says, but in considering actual selection process that might have supported the existence of a conscience Boehm's social selection' theory described above "explains the rather unusual set of agencies that created this moral faculty for us." Once shame feelings emerged "there would have been no question about [humans] moral status, reflecting that humans had strongly internalized group values and a sense of right and wrong," concluded Boehm. The expression of shame, guilt, and embarrassment enables the deviant to avoid punishment by others and restores one's place in the group.  But was the environmental trigger that induced the expression of the shame emotion in humans the behavior of other humans?  Although he does not expressly say this, Boehm's "social selection" theory seems to suggest this.

What we are ultimately interested in here is the development of altruistic behavior: direct and indirect reciprocal behavior (see September 27, 2012 post and September 12, 2012 post) in terms of the consequences described in Damasio's categories above, cooperation. What were the environmental drivers that triggered the development of moral emotions and ultimately the capacity to engage in altruistic reciprocal exchange, particularly that exchange that does not require direct reciprocal exchange?  As Martin Nowak observed in Supercooperators, "We are the only species that "can summon the full power of indirect reciprocity, thanks to our rich and flexible language."  (See September 17, 2012 post).

Human brain size and structure.  While the social insects demonstrate that cooperation is not a function of brain size (see November 4, 2009 post) , in hominids, I submit, it is highly relevant. Also relevant, according to recent research, are variations between species in the relative size of certain areas of this larger brain and the extent of interconnections between areas that can explain differences in temperament and behavior that translates into greater empathy and less aggression, which may or may not have co-evolved with the enlargement of the brain.  Also relevant may be differences in neurons between species.  (See November 9, 2010 post).  Boehm also believes this subject is relevant to his inquiry about the evolution of conscience, but he is not certain just how much weight to assign to his answer. According to Lynch and Granger in their book Big Brain, the chimpanzee has a brain size roughly 350-400 cubic centimeters (cc); the extinct Australopithecus, a brain size of about 440-450cc; extinct homo habilis, a brain size of about 600cc; the extinct homo erectus, a brain size of about 800cc-1000cc; the extinct homo heidelbergensis, a brain size of about 1200cc; the extinct neanderthal (homo neanderthalensis), a brain size of about 1500cc; and homo sapiens, a brain size of about 1350cc. In the evolutionary trail from the genus pan to the genus homo and the species within the genus homo, the brain enlarged over time (although the human brain has apparently shrunk in more modern times). The pan/homo divergence is believed to have occurred about 6 million years ago, and an emergent characteristic of the new species was bipedalism --- upright walking - which evolved with austalopithecines, perhaps as early as 3.6 million years ago, long before larger brain size. According to Lynch and Granger, changes to body that support walking increased lower trunk, and a side effect of a longer, lower trunk is to increase the space into which a uterus can expand during pregnancy. In women, this is accompanied by a wider pelvic girdle and the result is bigger babies. Bigger brains follows bigger babies. As the brain becomes bigger, most of the increase is not dedicated to sensory and motor needs, but to new neocortical areas. The biggest brains have much more cortex than smaller brains, and moreover, a bigger cortex has within it much more association cortex than sensory regions. This results in different relative intelligence for different brain size. Natural selection, conclude Lynch and Granger, did not select for intelligence. Intelligence was enabled by larger brains. Intelligence has been enabled by an expanded neocortex, which in humans is involved in higher functions such as sensory perception, generation of motor commands, spatial reasoning, conscious thought and language. The neanderthal evolved roughly 200-250,000 years ago in Europe and Asia; the homo sapien approximately 150-200,000 years ago in Africa. These capabilities of larger brains in homo sapiens did not emerge immediately. Language, for example, is not believed to have emerged until roughly 100,000 years ago. Language, I believe, is crucial for the evolution of a conscience, and Boehm appears to agree ("Morality is a group affair . . . Critical to this process is talking . . .").  Proto-languages, however, may have existed with earlier species such as homo erectus, but we will never know.  Equally critical to cooperative behavior in my view, particularly because it must be tied to indirect reciprocity, is the development of long-term memory, which is said to be encoded in the medial temporal lobe. 

As Antonio Damasio described above, parts of the prefrontal cortex are significant to social emotional associations, decision-making and moderating social behavior. Studies show that damage to a part of the ventromedial prefrontal cortex can impair empathetic responses in humans.  While other pan species have a prefrontal cortex, the prefrontal cortex of the human brain is significantly larger than that of chimpanzees, bonobos, and other apes, it also has more substantial folding (hence greater surface area) in this area than chimps and apes, and this has significant implications for behavioral differences that distinguish chimpanzees from humans.  Since we have seen that sympathy and empathy have been observed in the pan species, chimpanzees and bonobos, and that these species have prefrontal cortex structures similar, but not identical to humans, the human capacity for empathy and sympathy has its origins long ago in our non-human ancestors.  The larger brain capacity in homo, it is my belief, together with relative enlargement of certain areas of the brain and the connections between them, has probably contributed to their greater ability to engage in self-control over their egotistical impulses and greater empathy and compassion for other humans and perhaps other animals.  What we don't possess at this time is much in the way of research on the neural correlates of shame and guilt.  An Internet search came across only a couple of recent papers that acknowledged there had not been much research in this area, and that research might advance this discussion a bit.

Selfish or self-control?  In a prior post discussing Richard Wrangham's Demonic Males (see July 1, 2010 post), I noted, "If this was the only book one read on the subject of the origins of human behavior, you would conclude that male humans are born to kill, but empirical observation tells us that is not the only truism one can declare about male humans. Demonic Males is very interesting for what it documents: the apparent origins of human violence can be found in other primate species closest to humans, which indicates a genetic and evolutionary foundation for violent behavior among human males. But there is more to the story that primatologists are telling us about the commonalities among primate behaviors, including the origins of social cohesion, cooperation, and even morality and altruism. Michael Gazzaniga's Human (September 27, 2009 post) contributes much to this broader understanding of human evolution. Wrangham barely mentions these other common attributes of primate social systems. Violence is only one behavioral aspect of our social nature."  Boehm would agree that humans are very different than chimpanzees, and what he endeavors to explain in Moral Origins is that notwithstanding the fact that selfish, egoistic impulses are stronger in humans than altruistic impulses, humans have evolved a capacity for self-control that suppresses selfishness and campaigns for generosity and the Golden Rule. Boehm falls in the line of thinkers that includes Frans DeWaal and repeatedly emphasizes our dual nature:  we are selfish egoists and we are selfless altruists.  As I described in the post discussing DeWaal's The Age Empathy (see November 9, 2010 post),   "DeWaal takes aim at three myths: (1) the myth that our ancestors --- 4 foot bipedal apes --- ruled the savanna in Africa; (2) that human society is a voluntary creation of autonomous men; and (3) that our species has been waging war for as long as it has been around. Our ancestors were likely both prey and predator and survival favored genes that encouraged collaboration and companionship. The idea that humans were autonomous falsely presumes they had no need for anybody else and could voluntarily choose to live apart, uncommitted to anyone else or any place. A warlike initial state of nature that philosophers like Rousseau imagined that was overcome by social compacts is actually the reverse of human evolution: war on a grand scale, like we have known for centuries, came only after social hierarchies were formed and wealth was created. The early human species was probably defined more by social commitments and small scale collaboration that promoted primitive economic exchange and division of labor."  Moral Origins is an attempt to provide a historical account of how these social commitments and small scale collaboration evolved. 

Fire and group behavior.  Recall E.O. Wilson's view that a key event (but certainly not the only event) in the development of human eusociality was the mastery of fire.  (See September 12, 2012 post).  The nest, or its equivalent, is viewed by Wilson as a key to eusociality, and the campfire, according to Wilson, is the human equivalent of a nest found in other eusocial species.  For genus homo, mastery of fire occurred first during the era of homo erectus, perhaps as early as one million years ago but certainly 600-800,000 years ago, and long before the emergence of homo sapiens and homo neanderthalensis 200,000 years ago.  The development and use of early tools is also associated with homo erectus, and homo erectus may have lived in small bands of 30-50 men and women.  What we do not know is whether homo erectus' ability to control fire was merely the use of fire otherwise started naturally, or resulted in the development of a campfire created by homo erectus --- the equivalent of the nest in other eusocial species.  There is a debate as to whether or not homo erectus had vocal communications ability.   But clearly we can see in this species some of the precursors of human eusociality that both Wilson (September 12, 2012 post) and Martin Nowak (see September 17, 2012 post) addressed. 

The campfire is not merely a nest but it is a location where food, particularly protein-rich meat is cooked and consumed.  This is significant for the larger-brain hominid described above.  Recent studies indicate that increased consumption of cooked vegetation and meat supports larger brains.  The larger brain is not, as described above, merely defined in terms of cranial capacity, but greater neuronal capacity ("neuron rich").  If larger hominids had larger brains, as Lynch and Granger posit, it is plausible that the more successful ("fit") of these larger hominids would put these larger brains to better use than other hominids.  The brain with greater neuronal connections to other parts of the brain and increased neuronal capacity is more likely to do that, and to support those connections and neurons requires increased consumption of cooked vegetation and meat to meet the energy demands of that brain.  It is also plausible that the larger hominid with the larger brain would have to kill larger animals and consume them more frequently than the chimpanzee and other apes and hominids, and to accomplish that feat it is plausible that hunting, killing, and cooking meat in groups is a more successful ("fit") strategy.  To incent that group behavior and longer-term group cohesion for capturing and killing large animals, it is plausible that sharing meat in the vicinity of the campfire would enhance success/survival ("fitness").  Thus, the campfire and brain size are plausibly linked to survival of the hominid phenotype. 

Christopher Boehm notes that most other researchers have taken an ahistorical approach to moral origins among humans, focusing as does E.O. Wilson (September 12, 2012 post) on evolutionary theory as applied to kin selection and group selection or, as does Martin Nowak,(September 17, 2012 post) on mathematical fitness estimations for conditions favoring reciprocal behavior. Boehm offers a historical approach, but to his own credit he, like Charles Darwin, admits to the difficulties in this approach:  we have no written record from the pan troglodytes of 3-6 million years ago, or the austalopethicines of 3.5 million years or the homo habilis or the homo erectus, or even the more recent homo neanderthalensis that followed, about how these extinct species interacted socially (the extent of their within group or other group competition or cooperation), how they communicated.  Furthermore, we have a sketchy, but growing fossil record of these now extinct species, but the fossil record tells us no more about the organization of the brains of these species; all we know is their cranial capacity.  What we do have is evidence of fires, markings on bones, shaped rocks that indicates their use as tools or hunting devices found in the vicinity of the remains of these extinct species and capable of carbon dating, as well as the bones of animals nearby indicating that humans or their predecessors consumed the meat of these animals.  These are true challenges for developing a historical record of what these extinct species were actually like, and Boehm, like Darwin, is forced to describe a history based on its "general plausibility" by providing a working hypothesis, some of which are no more than "glorified hunches, while others may seem them as highly worthwhile leads for future research." 

In addition to the sketchy fossil record described above, Boehm also has at his disposal a body of evidence that can be observed today that is relevant to developing his history:  chimpanzee and bonobo behavior that is presumably similar to their behavior 6 million years ago when the ancestors of genus homo split from pan troglodytes, and the recorded observations of scholars about these species are growing; furthermore, we have a record of observations about a dwindling number of  homo sapiens hunter-gatherer groups developed over the past century to the current period, which behavior is presumably not much different than homo sapiens hunter gatherers who existed 35,000 to perhaps 200,000 years ago.  Based on these observations, Boehm concludes that human hunter gatherers, both now and then, exhibit a behavior that is distinctly different than the behavior of our closest relative, the chimpanzee.  Whereas, the male chimpanzee, is obsessed with dominance and rank and lives in social groups organized hierarchically led by an "alpha male," often displaying in-group aggression against one another and instability in rank, and female chimpanzees less obsessed with rank and dominance, less social, and less aggressive than their male counterparts, the human hunter gatherer of the late Pleistocene, based on modern observations, is characterized by egalitarian relations among persons within small groups, says Boehm.  Something happened in the evolution of the genus pan, to the australopethicines, and genus homo over the 5.8 million year period from the time of split and the emergence of homo sapiens about 200,000 years ago that transformed one species from a hierarchical dominance obsessed social group marked by within group aggression to another species characterized by within-group egalitarianism.  That is the historical trail Boehm seeks to describe in discovering how humans developed a conscience, whereby humans internalize group rules. 

Boehm believes there is a special type of natural selection that he calls "social selection" that involves "the effects of human preferences in choosing others in useful partnerships or in coming down hard on disliked deviants [cheaters]."  His working hypothesis is that at some point in the course of human history, group punishment of cheaters became severe and frequent and affected the human gene pool and ultimately favored human individuals with greater self-control in order to avoid group punishment. Group punishment could have been lethal, Boehm surmises, but it could have involved ostracism or deprivation of what was treated as community property, notably meat.  Whatever the form of punishment invites submissiveness to the group.  The "instrument" of self-control, believes Boehm, is the conscience.  What triggered this development, he submits, is humans embarking on a "new kind of subsistence pattern based on hunting" large animals for food that could only be met by groups.  For these groups to have any kind cohesion required an efficient, equitable sharing of the meat of these large animals.  The obstacle to setting up this egalitarian scenario for a small band of human hunters, Boehm recognizes, is the nature of the ancestral alpha male prone to appropriating the meat of others, which has not altogether disappeared from human nature today.  To achieve egalitarian relations among a small group of human hunters required the threat of force, enforced by the small group.  He believes that this evolutionary trend began around 200-250,000 years ago and culminated approximately 45,000 years ago.  He says this is a "tentative hypothesis," and new archaeological finds and future developments in behavioral genetics could lead to alternative hypotheses. 

Boehm suspects that the step from hierarchically-organized rank-oriented aggressive chimpanzees to egalitarian human hunter-gatherers was motivated by "rank-and-file envy over the perks of alpha bullies, which related to power, food, and sex."  In other words, at some point in time a few million years ago along the trail to homo sapiens, the submissive gradually said we have had enough, we are tired of being intimidated.  The submissive "would have developed some systematic type of collectivized and potentially lethal social control . . . to prevent high-ranking bullies from just naturally monopolizing large carcasses killed by group members and acting as free-riders, when it was the undernourished others in the band who were doing the hard work of hunting."  This type of social sanctioning is observed in modern human hunter gatherer groups in the modern era.  The alpha males are compelled to adapt to the group's will by achieving greater self-control over their egoistic impulses in order to gain greater fitness.  Boehm admits that it is possible that the evolution to egalitarianism began earlier than 200-250,000 years ago when homo sapiens first appeared on the scene (e.g. during the era of homo erectus as early as 1.8 million years ago), but at this point this is even more speculative than the plausible scenario found in the human fossil record.  Boehm points to evidence of carcasses butchered by archaic humans 400,000 years ago that show cut marks on bones that are "chaotic and varied" suggesting that several people did the butchering, consistent with what is observed in chimpanzee and bonobo meat eating scenes.  More recently, about 200,000 years ago there is evidence of cut marks on the bones of animal carcasses that are consistent with a single individual acting as butcher "reminiscent of what takes place with modern hunter-gatherers, where in effect the meat becomes a vigilant band's common property, to be widely shared in a systematic, culturally routinized fashion that averts serious conflict" where the hunters turn the carcass over to a "neutral meat distributor who is uninvolved with the kill," preventing a successful hunter from egoistically controlling the meat.  In contrast, while chimpanzees do share meat, it not a significant part of their diet and the meat does not come from large animals. The strategic motivation for meat sharing among chimpanzees appears to be for the support alliances with other chimps to maintain alpha male dominance within a group rather than support an egalitarian social environment between them. 

We can note several things about Boehm's hypothesis.  The timetable is consistent with the evolution of a larger brain in humans that we know is organized to be more socially responsive to others, both emotionally but also in terms of exercising self-control.  Boehm estimates that human moral origins appeared 35-45,000 years ago.  Interestingly, this suggests that it took about 150,000 - 160,000 years for the larger-brained homo sapiens to evolve their group structures and emotions to support collaborative hunter-gatherer groups. Did the structure of the human brain subtly evolve during this period so that certain homo sapiens with a particular brain structure and types of neurons were selected over other homo sapiens who did not have the same organization? The timetable is also consistent with what we believe occurred with the development of spoken language in humans, which has to be a key to the development of moral rules among humans. Boehm cites evidence of "preaching behavior" among egalitarian hunter-gatherer societies that may have been important to "behaviorally amplify the sympathetic generous tendencies of group members," a behavior that carried over to later hierarchical societies among humans that evolved when settled agrarianism later took root.

Finally, "social selection" is not merely the effect of the suppression of free-riding egoists; rather the intimidation and punishment of deviants acts in combination with "reputational selection."  This is a reference to what we have referred to in the previous posts (see September 17, 2012 post and September 12, 2002 post) as indirect reciprocity.  Simply put, for Boehm, there must be an explanation for why humans extend altruism to non-kin and developed a sense of virtue.  I wonder if we are not over-labeling "selection."  I think Boehm is getting close to saying that there is a gene for altruism, and in fact in his chapter titled "Testing the Selection-by-Reputation Hypothesis" he begins to refer to "altruistic genes" and "genes made for altruism."  I am skeptical of this line of thinking.  Clearly, indirect reciprocity among persons who do not know each other well or know each other at all is based on reputation.  It is a real phenomenon. We often join groups based on reputation. But what nature selects for is emotions and feelings, like attachment and sympathy and empathy, and these emotions in combination with other biological attributes selected by nature end up promoting reciprocal altruism, directly and indirectly.

Finally, what the research described by Christopher Boehm reveals is that the origins of human morality appeared tens of thousands of years before humans created social institutions and hierarchies, including religious and governmental institutions.  Importantly, morality precedes religion and the genetic origins (at least some of the genetic origins) of morality can be linked back to hundreds of thousands if not millions of years ago in other species.  Moral behavior is thus part of nature; it is not owned by religion, nor is it religiously inspired, and it predates the human mind's invention of deities.  Boehm suspects that the origins of religious belief in the brain's capacity for patternicity and agenticity, as described by Michael Shermer (see June 12, 2011 post) probably co-evolved with the origins of morality during the late Pleistocene, but gods, religious institutions, and the co-opting of morality by religious institutions came later, when humans began creating permanent communities as they transformed from hunter-gatherers to agrarian life and domestication of animals.