CAS behavior
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CAS behavior



Summary
The complexity of behavior is explored through Sapolsky developing scenarios of our best and worst behaviors across time spans, and scientific subjects including: anthropology, psychology, neuroscience, sociology.  The rich network of adaptive flows he outlines provides insights and highlight challenges for scientific research on behavior. 

Complex adaptive system (CAS) theory builds on Sapolsky's details highlighting the strategies that evolution has captured to successfully enter niches we now occupy. 

Behave
In Robert Sapolsky's book 'Behave' he binds his research on baboons to the complex adaptive systems (
This page introduces the complex adaptive system (CAS) theory frame.  The theory is positioned relative to the natural sciences.  It catalogs the laws and strategies which underpin the operation of systems that are based on the interaction of emergent agents. 
John Holland's framework for representing complexity is outlined.  Links to other key aspects of CAS theory discussed at the site are presented. 
CAS
) that influence their and our behavior. 

The Behavior
Sapolsky explains the approach of the book.  In the first part some good or bad behavior occurs driven by selection of a strategy.  Typically the context and meaning of the behavior are more complex than the mechanics.  The complexity can only be understood by incorporating all of the following aspects: 
In the second part of the book Sapolsky looks at realms of behavior where the strategies already described are most important.  This includes behaviors that are:
He concludes by summarizing that nothing seems to cause anything; instead everything just modulates something else.  But he says we lucky educated humans must try to cope with complexity and be our best.  We can be scientific and compassionate indicates an emotional state where resonance with someone else's distress leads one to help them. 


And he notes the challenges in developing these complex descriptions:

One Second Before
Sapolsky explores the neurobiology of the brain that commanded the muscles to act in the second before the behavior.  He leverages Maclean's Triune brain is Dr. Paul MacLean's popular but discredited 1940s theory of the brain.  He proposed a three layer structure:
  1. Reptilian inner brain containing circuits for basic survival; which is interfaced to layer 2 through the hypothalamus and together with the brain stem, spine and projections into the body make up the autonomic nervous system. 
  2. Limbic middle brain containing emotional circuits which signal layer 1 through the hypothalamus. 
  3. Rational outer brain which is uniquely human.  
because while wrong it provides a good organizing metaphor for the automaticity, emotion are low level agents distributed across the brain and body which associate, via the amygdala and rich club hubs, important environmental signals with encoded high speed sensors, and distributed programs of action to model: predict, prioritize guidance signals, select and respond effectively, coherently and rapidly to the initial signal.  The majority of emotion centered brain regions interface to the midbrain through the hypothalamus.  The most accessible signs of emotions are the hard to control and universal facial expressions.  Emotions provide prioritization for conscious access given that an animal has only one body, but possibly many cells, with which to achieve its highest level goals.  Because of this emotions clash with group goals and are disparaged by the powerful.  Evolutionary psychology argues evolution shaped human emotions during the long period of hunter-gatherer existence in the African savanna.  Human emotions are universal and include: Anger, Appreciation of natural beauty, Disgust, Fear, Gratitude, Grief, Guilt, Happiness, Honor, Jealousy, Liking, Love, Rage, Romantic love, Lust for revenge, Passion, Sadness, Self-control, Shame, Sympathy, Surprise; and the sham emotions and distrust induced by reciprocal altruism.   and cognition is the ability to orchestrate thought and action in accordance with internal goals according to Princeton's Jonathan Cohen. 
aspects of behaviors.  What were the crucial proximate
Plans are interpreted and implemented by agents.  This page discusses the properties of agents in a complex adaptive system (CAS). 
It then presents examples of agents in different CAS.  The examples include a computer program where modeling and actions are performed by software agents.  These software agents are aggregates. 
The participation of agents in flows is introduced and some implications of this are outlined. 
agents
?
Within this representation Sapolsky calls out three brain regions that are the most significant contributors to this final common pathway that funnels all the factors of the overall
This page introduces the complex adaptive system (CAS) theory frame.  The theory is positioned relative to the natural sciences.  It catalogs the laws and strategies which underpin the operation of systems that are based on the interaction of emergent agents. 
John Holland's framework for representing complexity is outlined.  Links to other key aspects of CAS theory discussed at the site are presented. 
CAS
, described in subsequent chapters of Behave, to converge and create our best and worst behaviors.  The main
Plans are interpreted and implemented by agents.  This page discusses the properties of agents in a complex adaptive system (CAS). 
It then presents examples of agents in different CAS.  The examples include a computer program where modeling and actions are performed by software agents.  These software agents are aggregates. 
The participation of agents in flows is introduced and some implications of this are outlined. 
agents
are the: Amygdala contains > 12 distinct areas: Central, Lateral.  It receives simple signals from the lower parts of the brain: pain from the PAG; and abstract complex information from the highest areas: Disgust from the insula cortex.  It sends signals to almost every other part of the brain, including to the decision-making circuitry of the frontal lobes.  It has high levels of D(1) dopamine receptors.  During extreme fear the amygdala drives the hippocampus into fear learning.  It outputs directly to subcortical reflexive motor pathways when speed is required.  Its central nucleus projects to the BNST.  It signals the locus ceruleus.  The amygdala:
  • Promotes aggression.  Stimulating the amygdala promotes rage.  It converts anger into aggression and when impaired it impacts the ability to detect angry facial expressions.  
  • Participates in disgust
  • Perceives fear promoting stimuli.  In PTSD sufferers the Amygdala overreacts to mildly fearful stimuli and is slow to calm down and the amygdala expands in size over a period of months.  Fear is processed by the lateral nucleus which serves as the input from various senses, and the central nucleus which outputs to the brain stem (central grey - freezing, lateral hypothalamus - blood pressure, activates paraventricular hypothalamus => crf -> hormone adjustments). 
  • Has lots of receptors for and is highly sensitive to glucocorticoids.  Stress inhibits the GABA interneurons in the basolateral amygdala (BLA) allowing the excitatory glutamate releasing neurons to excite more. 
  • Is sensitive to unsettling/uncertain social situations where it promotes anxiety.  It is also interested in uncertain but potentially painful situations.  The amygdala contributes to social and emotional decision making where the BLA supports rejecting an unacceptable offer, as allowed in the Ultimatum Game, by injecting implicit mistrust and vigilance, generating an anger driven rejection that is used as punishment.  The amygdala is very rapidly excited by subliminal signals from the thalamus of outgroup skin color.  The amygdala subsequently tips social emotions against outgroups unless restrained by the frontal lobe or influenced by subliminal priming to prioritize inclusion.  The fast path from the thalamus rapidly but inaccurately signals its identified a weapon. 
  • Promotes male, but not female, sexual motivation when it is an uncertain potential pleasure. 
  • Responds to the longing for uncertain potential pleasures and fear that the reward will not be worth it if it happens.  The amygdala turns off during orgasm. 
  • Uses but is not directly involved in vision. 
, Frontal cortex, mesolimbic includes projections between the tegmentum and the accumbens, amygdala, hippocampus;
/mesocortical dopamine pathways includes projections between the tegmentum and the prefrontal cortex;
; with other occasional bit players.  And their contributions alter as the agent's adapt depending on the other factors of the
This page introduces the complex adaptive system (CAS) theory frame.  The theory is positioned relative to the natural sciences.  It catalogs the laws and strategies which underpin the operation of systems that are based on the interaction of emergent agents. 
John Holland's framework for representing complexity is outlined.  Links to other key aspects of CAS theory discussed at the site are presented. 
CAS
.  This then affects the other factors: fusiform is a region of the brain which supports advanced mechanisms of shape recognition and implements the early stages of reading.  Subliminal priming with words did not depend on the shape of the word.  The fusiform gyrus was able to process the abstract identity of a word without caring if it was upper or lower case.   While high up in the cortex it can operate below the level of conscious experience.  It contributes to social emotions with:
  • Its face area being more activated by faces with in-group skin color.  
  • It activating when shown pictures of cars in automobile aficionados. 
  • It activating when shown pictures of birds in birdwatchers; since it really recognizes examples of items from an individual's emotionally salient categories. 
pays attention to what ever has become highly salient to that individual. 

The Amygdala
The amygdala is strongly associated with aggression, fear is an emotion which prepares the body for time sensitive action: Blood is sent to the muscles from the gut and skin, Adrenalin is released stimulating: Fuel to be released from the liver, Blood is encouraged to clot, and Face is wide-eyed and fearful.  The short-term high priority goal, experienced as a sense of urgency, is to flee, fight or deflect the danger.  There are both 'innate' - really high priority learning - which are mediated by the central amydala and learned fears which are mediated by the BLA which learns to fear a stimulus and then signals the central amygdala. 
, anxiety is manifested in the amygdala mediating inhibition of dopamine rewards.  Major anxiety results in elevated glucocorticoids and reduces hippocampal dendrite & spine density.  Some estrogen receptor variants are associated with anxiety in women. 
, social & emotional are emotions that are induced in response to other people's signals, are implemented by specific brain regions including: Prefrontal cortex, Insula cortex, Anterior cingulate cortex, Amygdala; receive lots of projections from interoceptive networks.  Sapolsky asserts in the moments just before we prioritize a consequential act the process is less rational and autonomous than we assume.  There are many significant signals from the prior seconds to minutes that effect social emotions:
  • Our brains respond subliminally to skin color very quickly: Amygdala activates, Fusiform face area activates; prior to the conscious stream activating the anterior cingulate and DLPFC which then inhibit the amygdala. 
  • Social dominance is culture independent and accurately subliminally assessed after a 40-millisecond exposure.  Stable status relations activate the ventromedial prefrontal cortex and DLPFC, while a dynamic situation also activates the amygdala.  
  • People who are subliminally judged attractive by the medial orbitofrontal cortex are considered kinder, smarter and more honest.  They are given more breaks. 
  • Faces and eyes in particular are most important subliminal cues.  They are monitored by the fusiform.  People respond more appropriately under the subliminal influence of eyes. 
  • Olfactory sensors send more direct projections to the limbic network than other sensory networks.  Pheromones signal fear activating the amygdala. 
  • Observing pain responses in others results in empathy even among young children. 
  • Words are important emotional signals providing unconscious priming of social responses.  Kahneman & Tversky demonstrated how the phrase '95% survival rate' is found to be a more acceptable choice than '5% death rate'.  Sapolsky notes that prosocial word priming fosters cooperation with antisocial word priming doing the opposite.  
  • Cultural objects such as visible: flags, team badges; subliminally modify in-group outgroup decisions. 
  • The presence of women in a situation alters the responses of men: Increased risk-taking, more focus on luxuries, increased aggression; in circumstances where conflict is already encouraged but not when status is achieved prosocially.  
  • Physical environment shapes behavior as demonstrated by Philip Zimbardo and leveraged in broken windows policing.  
  • Bodily adjustments to sensory structures introduce adaptive complexity, with the brain being influenced to become more sensitive and alter the sensor networks to make some more sensitive.  But these adaptations also vary culturally.  Collectivist cultures focus on a visual scene's surrounding contextual information while people from individualistic cultures focus on the focal object!   
decision making, social reflects the amygdala's particular sensitivity to unsettling social circumstances that stimulate anxiety because of uncertainty.  Being unsure of your place in a hierarchy is unsettling activating the amygdala.  Rationally going against a counter-factual but group agreed judgement stimulates the amygdala.   uncertainty is when a factor is hard to measure because it is dependent on many interconnected agents and may be affected by infrastructure and evolved amplifiers.  This is different from Risk.  , [moral] disgust is a universal human emotion.  Pinker notes it has its own facial expression and is codified in food taboos.  The mind must be associated with the proximate environment and parents minimize the risk for their omnivorous children by teaching them what foods to eat and what to avoid.  The children's minds are initially receptive to trying all foods but their brains subsequently lock in on the foods they have experienced.  These parental choices are affected by schematic influence on what has been beneficial.  Adolescent's brain developments undermine these constraints enabling intergroup transfers.  Disgust is modulated by the insula cortex which projects signals to the amygdala. 
; with lots of supporting experimental evidence.  It gets signals from all sensory systems.  It is especially signalled about: pain amplifies the aggression response of people by interoceptive signalling of brain regions providing social emotions including the PAG projecting to the amygdala; making aggressive people more so and less aggressive people less so.  Pain is the main reason people visit the ED in the US.  , disgust.  It sends alarms throughout the brain & body and has fast circuits to stimulate movement.  Sapolsky explains how learning allows signals interpreted in sub-regions (e.g. auditory in the BLA is the basolateral amygdala, a relatively recently evolved part of the amygdala which learns stimuli to fear and then signals the central amygdala.  It recieves inputs from all sensory networks.  Some are fast pathways that allow the BLA to detect and respond when the sensory cortex is unaware.  But it is far less accurate than the cortex.  The BLA's learning involves increased excitability of synapses coupling the BLA and central amygdala.  This is due to gene driven: Increased levels of growth factors promoting new connections, more receptors for excitatory neurotransmitters in dendritic spines.  The BLA also responds to signals from the frontal cortex that a stimulus no longer appears frightening.  This subset of BLA cells respond inhibiting the associating subset.  Stress and glucocorticoids increase levels of CRH and BDNF encouraging the building of new dendrites and synapses. 
) of the amygdala to be bound: synaptic, a neuron structure which provides a junction with other neurons.  It generates signal molecules, either excitatory or inhibitory, which are kept in vesicles until the synapse is stimulated when the signal molecules are released across the synaptic cleft from the neuron.  The provisioning of synapses is under genetic control and is part of long term memory formation as identified by Eric Kandel.  Modulation signals (from slow receptors) initiate the synaptic strengthening which occurs in memory. 
, growth factor are chemical signals which stimulate cellular growth.  In the brain they induce plasticity. 
changes refers to lasting changes to the brain that occur throughout the life span of the organism.  Many aspects of the brain can be altered into adulthood.  Almost anything in the nervous system can change in response to sustained stimulus.  And in a different environment the changes will often reverse.  The changes include:
  • The strength of dendritic input alters due to genetic, neural and hormonal signals
    • Hebb notes that memories require strengthening of preexisting synapses.  Glutamate responsive neurons' post synaptic dendritic spines have two types of receptor: non-NMDA and NMDA.  NMDA channels are responsible for this strengthening mechanism.  LTP then occurs to prolong the increase in excitability of the synapse. 
    • The LTP operation results in calcium diffusion which triggers new spine formation in adjacent parts of the dendrite.  Eventually that can stimulate dentrite growth enabling more neurons to connect. 
    • Short term stress promotes hippocampal LTP.  
    • Sustained stress promotes:
      • Hippocampal & frontal cortex  LTD & suppresses LTP.  Subsequent reductions in NCAM then reduce dendrite and synapse density. 
      • Amygdala LTP and suppresses LTD boosting fear conditioning.  It increases BDNF levels and expands dendrites in the BLA. 
    • Depression and anxiety reduce hippocampal dendrite and spine number by reducing BDNF. 
  • The axon's conditions for
    • Initiating an action potential. 
      • Progesterone boosts GABA-ergic neurons response to GABA decreasing the excitability of other neurons over a period of hours. 
    • Duration of a neuron's refractory period.  Testosterone shortens the refractory period of amygdala and amygdala target neurons over a period of hours. 
  • Synaptic connections being constantly removed and recreated
  • Synapses being created or destroyed.  Stimulation generates additional dendritic spines which become associated with a nearby axon terminal and within weeks a synapse forms.  The synapse then contributes calcium diffusion through LTP triggering more spine formation.  When dendritic spines recede synapses disappear. 
  • Cortical maps change to reflect alterations in the inputs and outputs from the body. 
  • Birth of brain cells in many areas of adult brains: the hippocampus (where 3% are replaced each month) and olfactory bulb and lesser amounts in the cortex. 
  • Restructuring after brain damage including axonal plasticity.  Distant rerouting of axons is observed but no mechanism has been identified yet. 
  • Vision is plastic in predators, where the eyes are moved during final development.  Dehaene argues for neuronal recycling supporting reading.  
; to the central area is a relatively ancient evolved part of the amygdala which processes 'innate' fears.  It projects to the BNST to raise the heart rate and blood pressure in preparation for fight and flight. 
linking the signals to fear.  And signals from the frontal cortex can teach the amygdala not to fear some bound signal any more: medial PFC in Buckner's fMRI based analysis is part of the brain's default mode network.  It facilitates the flexible use of information during the construction of self-relevant mental simulations.  Nauta considered its ventromedial part as participating in the limbic system.  With the medial temporal lobe it converges on important integration nodes: posterior cingulate cortex.   signals to the inhibitory circuits of the BLA that a sound is not to be feared.  Even when involved in pleasure the amygdala actually contributes awareness of the uncertainty of a potential pleasure and anxiety, fear & anger is an emotion which protects a person who has been cheated by a supposed friend.  When the exploitation of the altruism is discovered, Steven Pinker explains, the result is a drive for moralistic aggression to hurt the cheater. 
that the reward may not happen.  And feedback from the autonomic nervous system does not alter what is felt but does change the intensity.  Fear and aggression are not inevitably intertwined by the amygdala but typically become so for people who are already prone to this and when aggression is reactive and frenzied. 

The Frontal Cortex
Sapolsky stresses the frontal cortex of the cerebral cortex is at the front of the brain.  It includes the: prefrontal cortex, motor cortex.  Sapolsky asserts it makes you do the harder thing when it's the right thing to do.  The frontal cortex supports working memory to sustain focus on a task.  It also coordinates the strategic actions necessary to achieve success.  It provides impulse control, regulation of emotion, and willpower.  The prefrontal cortex maintains focus by deprioritizing currently irrelevant streams of information.  The frontal cortex tracks rules.  Over a lifetime that builds into a costly activity.  Once it tires responses become less prosocial.  But practice shifts operation of tasks to the cerebellum.  The frontal cortex signals the tegmentum and accumbens with the conclusions of its expectancy/discrepancy calculations.  The frontal lobe provides executive function, considering bits of information, assessing patterns and then prioritizing the strategies.  The frontal lobe is the most recent part of the brain to evolve and involves a disproportionate percentage of primate-unique genes in its development and operation.  It does not complete development until the mid-20s.  It includes spindle neurons.  It is easily damaged.  Sapolsky (Nauta) notes that its ventromedial prefrontal cortex is a quasi-member of the limbic system. 
makes you do the harder thing when it's the right thing to do.  It is significant that it is the last brain region to fully mature.  This only happens during the mid-twenties.  The frontal cortex is highly distinctive: evolution, size, complexity.  There are many unique, highly individuated, primate genes active in the frontal cortex.  And one unique cell type: the von Economo neuron are also called spindle neurons and are:
  • Found only in: Primates, Whales, Dolphins, Elephants;
  • Present in the: Insula, Anterior cingulate - focused on empathy; integrating and repurposing these basic facilities into high level capabilities such as moral disgust. 
  • First neurons destroyed by FTD. 
which is central to empathy is the capability to relate to another person from their perspective.  It is implemented by spindle neurons.  It is context dependently mediated by estrogen.  It develops over time: Piaget's preoperational stage includes rudimentary empathy, Theory of mind supports the development; initially feeling someone's pain as one integrated being, then for them and eventually as them.  In adults, when someone else is hurt the anterior cingulate cortex, amygdala & insula activates projecting [scapegoating] to the vmPFC.  If the pain is physical the PAG activates and motor neurons for the area where the other person was injured.  The intertwining of the ACC amygdala & insula in adults results in attribution of fault even when there is none which can make it hard to step in and actually help.  But in seven-year-olds the activation is concrete: PAG and sensory & motor cortexes with minimal coupling to the rudimentary vmPFC.  In older children the vmPFC is coupled to limbic structures.  Ten to twelve year olds abstract empathy to classes of people.  By adolescence the vmPFC is coupled to theory of mind regions and intentional harm induces disgust via the amygdala.  Sapolsky explains adolescent boys are utilitarian and tend to accept inequality more than girls do.  But both sexes accept inequality as the way it is.  Sociopaths do not develop empathy. 
(ACC is either the
  • Anterior cingulate cortex which:
    • Is a central focus of empathy supporting people relating to other's pain.  This is dependent on oxytocin. 
    • In non-human mammals it processes interoceptive signals.  The ACC focuses the internal signals into high level 'gut intuitions.'  Pain catches the ACC's attention. 
    • Performs discrepancy detection from the outcome that was predicted - at a high level.  The ACC cares about the meaning of what is predicted. 
      • If the ACC has been convinced that a pain killer placebo has inhibited pain signals, the ACC will stay silent about actual pain that is signalled from interoceptive networks.  
      • The ACC will signal: physical pain, emotional pain, metaphorical pain, anxiety, disgust, embarrassment, social exclusion especially in adolescence; as one and the same.  The ACC's abnormalities being associated with major depression. 
    • Has a bridging role between the empathetic and self-interested pain monitor.  Sapolsky notes the ACC is essential for learning fear and conditioned avoidance by observation alone through an intermediate step of shared representation of self.  He concludes "At its core the ACC is about self-interest, with caring about the other person in pain as an add-on." 
  • American College of Cardiology
) and binding the insula to moral disgust.  The frontal cortex looks for patterns in its situational models and selects from strategic actions.  It partners with the cerebellum is involved with the efficiency of fine movement. It modulates the force and range of motion and is involved in motor coordination and the learning of motor skills.  Damage to the cerebellum impairs standing, walking, or performance of coordinated movements. A virtuoso pianist or other performing musician depends on their cerebellum.  The cerebellum receives visual, auditory, vestibular, and somatosensory information.  It also receives information about individual muscular movements being directed by the brain.  The cerebellum integrates this information and modifies the motor outflow, exerting a coordinating and smoothing effect on the movements.  However, patients born without a cerebellum have survived reasonably well.  The cerebellum is part of the implicit learning mechanism.  It is required for the rabbit eye-blink to be classically conditioned to respond to a sound, and puff of air (threat to eye).  It integrates the sound and puff and outputs the response to the motor area (blink).
which takes over repeated strategies including: bravery, honesty; making them automatic and rapid.  Within the frontal cortex the prefrontal cortex (PFC) is
  • The front part of the frontal lobe of the cerebral cortex.  It evolved most recently.  During adolescence when the PFC is still deploying, older brain agents provide equivalent strategies: ventral striatum.  The PFC has been implicated in planning, working memory: dorsolateral; decision making: Orbitofrontal cortex; and social behavior.  Different PFC circuits track internal reward driven strategies and externally signalled advice.  The PFC chooses between conflicting options, letting go or restraint, especially between cognition and emotions.  It imposes an overarching strategy for managing working memory.  It is essential for thinking about multiple items with different labels.  It includes neurons that are interested in particular sub-categories: Dog, Cat.  Once it has made a decision it signals the rest of the frontal lobe just behind it.  Glucocorticoids decrease excitability of the PFC.  
(PFC) sub-region is the decider, but it gets tired easily after which decisions become less prosocial is prioritizing benefiting others through: Help, Charity, Truth; even if many are acts of restitution to balance out antisocial acts.  Due to the Insula's binding of physical and metaphorical disgust, physically washing your hands can be enough to reduce the need for a prosocial act.  
.  Social complexity expands the frontal cortex.  FTD is frontotemporal degeneration, the most common form of dementia for people under age 60.  It results from progressive loss of neurons from the frontal or temporal lobes.  The first neurons known to die are von Economo neurons.  Tau and TDP43 proteins both congregate in these areas.  People with FTD display behavioral disinhibition and socially inappropriate behaviors.  They appear apathetic and don't make decisions because the PFC is being destroyed.  There are three main types: Behavioral variant, primary progressive aphasia, Disturbances of motor function;
destroys it.  The dorsolateral PFC is dorsolateral prefrontal cortex which is:
  • At the heart of decision making - highly rational, unsentimental 
  • A major agent of working memory
  • The most recently evolved part of the prefrontal cortex.  
  • Mainly interconnected with other parts of the cortex.  
is the most socially rational part of the prefrontal cortex.  It inhibits the amygdala contains > 12 distinct areas: Central, Lateral.  It receives simple signals from the lower parts of the brain: pain from the PAG; and abstract complex information from the highest areas: Disgust from the insula cortex.  It sends signals to almost every other part of the brain, including to the decision-making circuitry of the frontal lobes.  It has high levels of D(1) dopamine receptors.  During extreme fear the amygdala drives the hippocampus into fear learning.  It outputs directly to subcortical reflexive motor pathways when speed is required.  Its central nucleus projects to the BNST.  It signals the locus ceruleus.  The amygdala:
  • Promotes aggression.  Stimulating the amygdala promotes rage.  It converts anger into aggression and when impaired it impacts the ability to detect angry facial expressions.  
  • Participates in disgust
  • Perceives fear promoting stimuli.  In PTSD sufferers the Amygdala overreacts to mildly fearful stimuli and is slow to calm down and the amygdala expands in size over a period of months.  Fear is processed by the lateral nucleus which serves as the input from various senses, and the central nucleus which outputs to the brain stem (central grey - freezing, lateral hypothalamus - blood pressure, activates paraventricular hypothalamus => crf -> hormone adjustments). 
  • Has lots of receptors for and is highly sensitive to glucocorticoids.  Stress inhibits the GABA interneurons in the basolateral amygdala (BLA) allowing the excitatory glutamate releasing neurons to excite more. 
  • Is sensitive to unsettling/uncertain social situations where it promotes anxiety.  It is also interested in uncertain but potentially painful situations.  The amygdala contributes to social and emotional decision making where the BLA supports rejecting an unacceptable offer, as allowed in the Ultimatum Game, by injecting implicit mistrust and vigilance, generating an anger driven rejection that is used as punishment.  The amygdala is very rapidly excited by subliminal signals from the thalamus of outgroup skin color.  The amygdala subsequently tips social emotions against outgroups unless restrained by the frontal lobe or influenced by subliminal priming to prioritize inclusion.  The fast path from the thalamus rapidly but inaccurately signals its identified a weapon. 
  • Promotes male, but not female, sexual motivation when it is an uncertain potential pleasure. 
  • Responds to the longing for uncertain potential pleasures and fear that the reward will not be worth it if it happens.  The amygdala turns off during orgasm. 
  • Uses but is not directly involved in vision. 
's out-grouping responses.  The ventromedial PFC (vmPFC) is:
  • Focused on the impact of emotion on decision making
  • A participant in limbic system operations 
  • Many human behaviors involve interactions between the vmPFC, the limbic system & the dorsolateral prefrontal cortex.  Part of decision making is for the limbic system to internally simulate (often with the help of the sympathetic nervous system) what alternative outcomes of a decision will feel like with the results of these somatic marker analyses being reported to the vmPFC.  
  • Damage to the vmPFC results in bad decision making: Poor judgement in choosing friends & partners, Failure to respond to negative feedback; because they can't feel the issues. 
imparts essential decision making to social/emotional challenges and tightly integrates with limbic structures supports emotional circuits: Amygdala, Hippocampus, Septum, Habenula, Mammillary bodies; all of which signals the midbrain through the hypothalamus. The broad interconnections of these regions with a part of the frontal lobe suggested to Walle Nauta that it (ventromedial prefrontal cortex) is a quasi-member of the limbic network.  .  These two subparts are inversely correlated but typically collaborate: both are needed in different situations to be prosocialStress is a multi-faceted condition reflecting high cortisol levels.  Dr. Robert Sapolsky's studies of baboons indicate that stress helps build readiness for fight or flight.  As these actions occur the levels of cortisol return to the baseline rate.  A stressor is anything that disrupts the regular homeostatic balance.  The stress response is the array of neural and endocrine changes that occur to respond effectively to the crisis and reestablish homeostasis. 
  • The short term response to the stressor
    • activates the amygdala which: Stimulates the brain stem resulting in inhibition of the parasympathetic nervous system and activation of the sympathetic nervous system with the hormones epinephrine and norepinephrine deployed around the body, Activates the PVN which generates a cascade resulting in glucocorticoid secretion to: get energy to the muscles with increased blood pressure for a powerful response.  The brain's acuity and cognition are stimulated.  The immune system is stimulated with beta-endorphin and repair activities curtail.  But when the stressor is
  • long term: loneliness, debt; and no action is necessary, or possible, long term damage ensues.  Damage from such stress may only occur in specific situations: Nuclear families coping with parents moving in.  Sustained stress provides an evolved amplifier of a position of dominance and status.  It is a strategy in female aggression used to limit reproductive competition.  Sustained stress:
    • Stops the frontal cortex from ensuring we do the harder thing, instead substituting amplification of the individual's propensity for risk-taking and impairing risk assessment! 
    • Activates the integration between the thalamus and amygdala. 
      • Acts differently on the amygdala in comparison to the frontal cortex and hippocampus: Stress strengthens the integration between the Amygdala and the hippocampus, making the hippocampus fearful. 
      • BLA & BNST respond with increased BDNF levels and expanded dendrites persistently increasing anxiety and fear conditioning. 
    • Makes it easier to learn a fear association and to consolidate it into long-term memory.  Sustained stress makes it harder to unlearn fear by making the prefrontal cortex inhibit the BLA from learning to break the fear association and weakening the prefrontal cortex's hold over the amygdala.  And glucocorticoids decrease activation of the medial prefrontal cortex during processing of emotional faces.  Accuracy of assessing emotions from faces suffers.  A terrified rat generating lots of glucocorticoids will cause dendrites in the hippocampus to atrophy but when it generates the same amount from excitement of running on a wheel the dendrites expand.  The activation of the amygdala seems to determine how the hippocampus responds. 
    • Depletes the nucleus accumbens of dopamine biasing rats toward social subordination and biasing humans toward depression. 
    • Disrupts working memory by amplifying norepinephrine signalling in the prefrontal cortex and amygdala to prefrontal cortex signalling until they become destructive.  It also desynchronizes activation in different frontal lobe regions impacting shifting of attention. 
  • During depression, stress inhibits dopamine signalling. 
  • Strategies for stress reduction include: Mindfulness. 
impacts the frontal cortex: like distraction and heavy cortical load, it can dissociate the frontocortical importance signal about some action from the Boolean do/don't do it signal, explaining why you may do exactly the wrong thing under stress. 

Emotion are low level agents distributed across the brain and body which associate, via the amygdala and rich club hubs, important environmental signals with encoded high speed sensors, and distributed programs of action to model: predict, prioritize guidance signals, select and respond effectively, coherently and rapidly to the initial signal.  The majority of emotion centered brain regions interface to the midbrain through the hypothalamus.  The most accessible signs of emotions are the hard to control and universal facial expressions.  Emotions provide prioritization for conscious access given that an animal has only one body, but possibly many cells, with which to achieve its highest level goals.  Because of this emotions clash with group goals and are disparaged by the powerful.  Evolutionary psychology argues evolution shaped human emotions during the long period of hunter-gatherer existence in the African savanna.  Human emotions are universal and include: Anger, Appreciation of natural beauty, Disgust, Fear, Gratitude, Grief, Guilt, Happiness, Honor, Jealousy, Liking, Love, Rage, Romantic love, Lust for revenge, Passion, Sadness, Self-control, Shame, Sympathy, Surprise; and the sham emotions and distrust induced by reciprocal altruism.   can be controlled to some extent by thought.  There are two strategies:

Sapolsky argues the frontal cortex gets its metaphorical motivation to do the harder thing from the brains dopaminergic reward systems. 

The Mesolimbic/Mesocortical Dopamine System
Dopamine is a synaptic signal supporting generalized goal-directed behavior & anticipation of reward.  Its significance is that the receptors that detect the signal are of the slow acting type and are used to alter (modulate) the response of fast acting dopaminergic neural circuits in which the receptors are deployed (LTP).  The signal detects significant changes including predictions of models and actual results which differ unexpectedly.  The dopamine network architecture is designed to signal the possibility of any type of reward: Norm violation punishment, Winning a lottery, & Misfortune of an envied competitor.  Dopamine signalling:
  • Rescales continuously to accommodate the range of intensity offered by different stimuli.  So dopamine's responses to any reward habituate.  GABA is released by some tegmental neurons to induce habituation. 
  • Reflects the anticipation of reward.  It supports establishment of a relationship between a signal, working for a reward and obtaining the reward, but subsequently dopamine is mainly released encouraging the work, right after the signal supporting anticipation of the reward.  Anticipation requires learning and is reflected in hippocampus activity.  That explains context dependent cravings.  And the learning architecture means reliable cues become rewarding.  The accumbens supports willpower.  And dopamine
  • Promotes goal-oriented behavior needed to obtain & likely to achieve the reward - through the dopamine projections to the prefrontal cortex.  That makes dopamine central to:
    • Motivation.  This binding fails in depression - due to stress and in anxiety - due to signals from the amygdala.  
    • The prefrontal cortex's mesocortically stimulated support for willpower to act to delay rewards.  To sustain work for delayed rewards additional dopamine is released based on the length of the delay and the rewards uncertainty (modelled in the dorsolateral prefrontal cortex - which promotes the long term and the ventromedial prefrontal cortex - which promotes the short term) and the anticipated size of the reward (modelled in the accumbens).  Impulsiveness in ADHD is reflected in abnormal dopamine processing.  Addictive drugs bias the dopamine network towards impulsiveness.  
  • Is lowered by certain gene variants which induce: less dopamine in the synapse, fewer receptors, lower responsiveness of receptors; associated with (as tiny effects in hugely varying social scenarios): sensation seeking, risk taking, attentional problems, extroversion; where:
    • The receptor D4's gene shows high variability.  The D47R form is relatively unresponsive to dopamine.  
    • Dopamine is degraded by COMT.  The COMT gene includes a variant which is highly efficient reducing dopamine signalling but with complicating gene/environment interactions.  
    • Dopamine is removed from the synapse by a reuptake transporter DAT. 
is central to
Read Montague explores how brains make decisions.  In particular he explains how:
  • Evolution can create indirect abstract models, such as the dopamine system, that allow
  • Life changing real-time decisions to be made, and how
  • Schematic structures provide encodings of computable control structures which operate through and on incomputable, schematically encoded, physically active structures and operationally associated production functions. 
motivated pursuit of goals
.  Sapolsky explains the best and the worst behaviors involve the projections from the ventral tegmentum is part of the tegmentum which contains dopamine long system cell bodies (high levels of D(2)).  Their axons branch extensively and reach many areas including: Mesolimbic to the limbic system: amygdala, hippocampus; Tegmentostriatal to the nucleus accumbens, Mesocortical to the forebrain including the prefrontal cortex.  The terminals are fairly evenly distributed through out layers 1 - 6.  So dopamine can modulate input and output excitatory and inhibitory transmissions.  If a rodent wins a fight on his home territory, there are long-lasting increases in levels of testosterone receptors enhancing pleasurable effects.  The lateral ventral tegmental area is now known to be one of two large adrenergic pathways, along with the locus ceruleus.  During child birth the ventral tegmentum deploys more oxytocin receptors increasing its sensitivity to the neuropeptide.  
to the:
Social interactions are affected by dopamine rewards:

The Anticipation of Reward
The dopamine is a synaptic signal supporting generalized goal-directed behavior & anticipation of reward.  Its significance is that the receptors that detect the signal are of the slow acting type and are used to alter (modulate) the response of fast acting dopaminergic neural circuits in which the receptors are deployed (LTP).  The signal detects significant changes including predictions of models and actual results which differ unexpectedly.  The dopamine network architecture is designed to signal the possibility of any type of reward: Norm violation punishment, Winning a lottery, & Misfortune of an envied competitor.  Dopamine signalling:
  • Rescales continuously to accommodate the range of intensity offered by different stimuli.  So dopamine's responses to any reward habituate.  GABA is released by some tegmental neurons to induce habituation. 
  • Reflects the anticipation of reward.  It supports establishment of a relationship between a signal, working for a reward and obtaining the reward, but subsequently dopamine is mainly released encouraging the work, right after the signal supporting anticipation of the reward.  Anticipation requires learning and is reflected in hippocampus activity.  That explains context dependent cravings.  And the learning architecture means reliable cues become rewarding.  The accumbens supports willpower.  And dopamine
  • Promotes goal-oriented behavior needed to obtain & likely to achieve the reward - through the dopamine projections to the prefrontal cortex.  That makes dopamine central to:
    • Motivation.  This binding fails in depression - due to stress and in anxiety - due to signals from the amygdala.  
    • The prefrontal cortex's mesocortically stimulated support for willpower to act to delay rewards.  To sustain work for delayed rewards additional dopamine is released based on the length of the delay and the rewards uncertainty (modelled in the dorsolateral prefrontal cortex - which promotes the long term and the ventromedial prefrontal cortex - which promotes the short term) and the anticipated size of the reward (modelled in the accumbens).  Impulsiveness in ADHD is reflected in abnormal dopamine processing.  Addictive drugs bias the dopamine network towards impulsiveness.  
  • Is lowered by certain gene variants which induce: less dopamine in the synapse, fewer receptors, lower responsiveness of receptors; associated with (as tiny effects in hugely varying social scenarios): sensation seeking, risk taking, attentional problems, extroversion; where:
    • The receptor D4's gene shows high variability.  The D47R form is relatively unresponsive to dopamine.  
    • Dopamine is degraded by COMT.  The COMT gene includes a variant which is highly efficient reducing dopamine signalling but with complicating gene/environment interactions.  
    • Dopamine is removed from the synapse by a reuptake transporter DAT. 
network architecture ensures once reward contingencies are learned dopamine release is associated with anticipation.  Mastery, expectation & confidence of the result are rewarded until the binding fails!  The learning involves a circuit of excited hippocampus is a part of the brain involved in the temporary storage or coding of long-term episodic memory.  Memory formation in the cells of the hippocampus uses the MAP kinase signalling network which is impacted by sleep deprivation.  The hippocampus dependent memory system is directly affected by cholinergic changes throughout the wake-sleep cycle.  Increased acetylcholine during REM sleep promotes information attained during wakefulness to be stored in the hippocampus by suppressing previous excitatory connections while facilitating encoding without interference from previously stored information.  During slow-wave sleep low levels of acetylcholine cause the release of the suppression and allow for spontaneous recovery of hippocampal neurons resulting in memory consolidation.  It was initially associated with memory formation by McGill University's Dr. Brenda Milner, via studies of 'HM' Henry Molaison, whose medial temporal lobes had been surgically destroyed leaving him unable to create new memories.  The size of neurons' dendritic trees expands and contracts over a female rat's ovulatory cycle, with the peak in size and cognitive skills at the estrogen high point.  Adult neurogenesis occurs in the hippocampus (3% of neurons are replaced each month) where the new neurons integrate into preexisting circuits.  It is enhanced by learning, exercise, estrogen, antidepressants, environmental enrichment, and brain injury and inhibited by various stressors explains Sapolsky.  Prolonged stress makes the hippocampus atrophy.  He notes the new neurons are essential for integrating new information into preexisting schemas -- learning that two things you thought were the same are actually different. 
amygdala contains > 12 distinct areas: Central, Lateral.  It receives simple signals from the lower parts of the brain: pain from the PAG; and abstract complex information from the highest areas: Disgust from the insula cortex.  It sends signals to almost every other part of the brain, including to the decision-making circuitry of the frontal lobes.  It has high levels of D(1) dopamine receptors.  During extreme fear the amygdala drives the hippocampus into fear learning.  It outputs directly to subcortical reflexive motor pathways when speed is required.  Its central nucleus projects to the BNST.  It signals the locus ceruleus.  The amygdala:
  • Promotes aggression.  Stimulating the amygdala promotes rage.  It converts anger into aggression and when impaired it impacts the ability to detect angry facial expressions.  
  • Participates in disgust
  • Perceives fear promoting stimuli.  In PTSD sufferers the Amygdala overreacts to mildly fearful stimuli and is slow to calm down and the amygdala expands in size over a period of months.  Fear is processed by the lateral nucleus which serves as the input from various senses, and the central nucleus which outputs to the brain stem (central grey - freezing, lateral hypothalamus - blood pressure, activates paraventricular hypothalamus => crf -> hormone adjustments). 
  • Has lots of receptors for and is highly sensitive to glucocorticoids.  Stress inhibits the GABA interneurons in the basolateral amygdala (BLA) allowing the excitatory glutamate releasing neurons to excite more. 
  • Is sensitive to unsettling/uncertain social situations where it promotes anxiety.  It is also interested in uncertain but potentially painful situations.  The amygdala contributes to social and emotional decision making where the BLA supports rejecting an unacceptable offer, as allowed in the Ultimatum Game, by injecting implicit mistrust and vigilance, generating an anger driven rejection that is used as punishment.  The amygdala is very rapidly excited by subliminal signals from the thalamus of outgroup skin color.  The amygdala subsequently tips social emotions against outgroups unless restrained by the frontal lobe or influenced by subliminal priming to prioritize inclusion.  The fast path from the thalamus rapidly but inaccurately signals its identified a weapon. 
  • Promotes male, but not female, sexual motivation when it is an uncertain potential pleasure. 
  • Responds to the longing for uncertain potential pleasures and fear that the reward will not be worth it if it happens.  The amygdala turns off during orgasm. 
  • Uses but is not directly involved in vision. 
& frontal cortex of the cerebral cortex is at the front of the brain.  It includes the: prefrontal cortex, motor cortex.  Sapolsky asserts it makes you do the harder thing when it's the right thing to do.  The frontal cortex supports working memory to sustain focus on a task.  It also coordinates the strategic actions necessary to achieve success.  It provides impulse control, regulation of emotion, and willpower.  The prefrontal cortex maintains focus by deprioritizing currently irrelevant streams of information.  The frontal cortex tracks rules.  Over a lifetime that builds into a costly activity.  Once it tires responses become less prosocial.  But practice shifts operation of tasks to the cerebellum.  The frontal cortex signals the tegmentum and accumbens with the conclusions of its expectancy/discrepancy calculations.  The frontal lobe provides executive function, considering bits of information, assessing patterns and then prioritizing the strategies.  The frontal lobe is the most recent part of the brain to evolve and involves a disproportionate percentage of primate-unique genes in its development and operation.  It does not complete development until the mid-20s.  It includes spindle neurons.  It is easily damaged.  Sapolsky (Nauta) notes that its ventromedial prefrontal cortex is a quasi-member of the limbic system. 
neurons, specialized eukaryotic cells include channels which control flows of sodium and potassium ions across the massively extended cell membrane supporting an electro-chemical wave which is then converted into an outgoing chemical signal transmission from synapses which target nearby neuron or muscle cell receptors.  Neurons are supported by glial cells.  Neurons include a:
  • Receptive element - dendrites
  • Transmitting element - axon and synaptic terminals
  • Highly variable DNA schema using transposons. 
projecting to dopamine neurons.  This ensures the return of context-dependent cravings in addiction.  And it means more dopamine will be generated when the result becomes uncertain. 

University of Michigan's Huda Akil showed reliable cues to an impending reward eventually become rewarding.  Rats eventually enjoy the cue.  This is the foundation of fetishes. 

Crucially dopamine binds the value of a reward to the strategic action that attains the result.  This is implemented in the dopaminergic projections to the PFC is prefrontal cortex which is:
  • The front part of the frontal lobe of the cerebral cortex.  It evolved most recently.  During adolescence when the PFC is still deploying, older brain agents provide equivalent strategies: ventral striatum.  The PFC has been implicated in planning, working memory: dorsolateral; decision making: Orbitofrontal cortex; and social behavior.  Different PFC circuits track internal reward driven strategies and externally signalled advice.  The PFC chooses between conflicting options, letting go or restraint, especially between cognition and emotions.  It imposes an overarching strategy for managing working memory.  It is essential for thinking about multiple items with different labels.  It includes neurons that are interested in particular sub-categories: Dog, Cat.  Once it has made a decision it signals the rest of the frontal lobe just behind it.  Glucocorticoids decrease excitability of the PFC.  
which decides on the strategy to execute. 

Dopamine still operates effectively when the work is prolonged and the reward is delayed.  To maintain the action a second gradual increase in dopamine occurs.  The increase is a function of the length of the delay and the anticipated size of the reward.  The level of the increase in dopamine is controlled by the accumbens is a region of the basal forebrain rostral to the preoptic area and immediately adjacent to the septum.  The nucleus accumbens was closely associated with the limbic system and plays an important role in reinforcement.  If a rodent wins a fight on his home territory, there are long-lasting increases in levels of testosterone receptors enhancing pleasurable effects.  When prairie voles first mate, epi-genetic state changes are induced in the accumbens to support pair-bonding.  The accumbens projects to brain regions associated with movement.  The major pathways of dopaminergic neurons begin in the substantia nigra and the ventral tegmental area.  The amygdala projects back to the accumbens.  The tegmentostriatal system begins in the ventral tegmental area and projects to the nucleus accumbens.  The nucleus accumbens includes high levels of D1, D2 and D3 dopamine receptors located on the spine & shafts of dendrites of excitatory cells reduce the transfer of excitation from the dendrites to the cell bodies, so only especially strong excitatory inputs get through to the cell body to elicit excitation.  It also has D4 dopamine receptors which are highly variable.  The accumbens responds differently to rewards depending on maturity: In juveniles all reward levels result in the same response activity level, During adolescence the accumbens responds to small rewards negatively, and large rewards hugely, In adults the rewards result in measured scaled positive responses.  Chronic stress depletes dopamine from the nucleus accumbens biasing humans towards depression.   while dlPFC is dorsolateral prefrontal cortex which is:
  • At the heart of decision making - highly rational, unsentimental 
  • A major agent of working memory
  • The most recently evolved part of the prefrontal cortex.  
  • Mainly interconnected with other parts of the cortex.  
& vmPFC is ventromedial prefrontal cortex which is:
  • Focused on the impact of emotion on decision making
  • A participant in limbic system operations 
  • Many human behaviors involve interactions between the vmPFC, the limbic system & the dorsolateral prefrontal cortex.  Part of decision making is for the limbic system to internally simulate (often with the help of the sympathetic nervous system) what alternative outcomes of a decision will feel like with the results of these somatic marker analyses being reported to the vmPFC.  
  • Damage to the vmPFC results in bad decision making: Poor judgement in choosing friends & partners, Failure to respond to negative feedback; because they can't feel the issues. 
neurons assess the time delay.  A flexible arrangement that supports delayed gratification.  Differences in the volume of these different agents in different people results in the variations in effectiveness of performing delayed gratification.  Sapalsky adds that humans can delay gratification for huge amounts of time in pursuit of abstract goals.  It has not yet been determined exactly how this is done. 

A Final Small Topic: Serotonin
Sapolsky notes that serotonin is a neurotransmitter.  it is:
  • Inversely associated with: human impulsive, cricket, mollusk, crustacean; aggression.  Low levels of serotonin are associated with impulsive aggression ranging from psychological measures of hostility to overt violence and cognitive impulsivity and impulsive suicide. 
  • Nearly all synthesized in the Raphe nucleus.  Tryptophan hydroxylase makes serotonin from the amino-acid tryptophan.  Monoamine oxidase degrades serotonin.  The serotonin receptor binds serotonin to initiate cross membrane signalling.  The serotonin transporter actively removes serotonin from synapses.  Reuptake is inhibited by SSRIs.  Variants of the genes coding for these various enzymes alter the strength of their effects. 
  • Increasing serotonin signalling does not lessen impulsiveness in normal subjects but did in those prone to impulsivity.  However, such experiments are fraught with complexity:
    • Transient changes induced by drugs may adjust the immediate levels of serotonin but may not demonstrate structural effects. 
    • Gene variants likely produce structural changes in the developing brain.  
    • Effects monitored in experiments are often tiny.  
    • Behavioral changes: Violence, Arson, Exhibitionism; seen in different test subjects may be difficult to compare.  
    • Monoamine oxidase has high gene/environment interactions undermining heritability estimates.  Its gene promotor is regulated by stress and glucocorticoids.  So non genetic factors such as childhood adversity and adult provocation appear to be significant.  
effects impulsivity.  The raphe nucleus consists of a group of nuclei running through the core of the brain stem from the medulla to the back of the midbrain.  It projects to the tegmentum, accumbens, prefrontal cortex, & amygdala where serotonin enhances dopamine's effects on goal-directed behavior.  The raphe is associated with:
  • Neural mechanisms of sleep and waking.  Jouvet and Renault produced large lesions that destroyed 80 - 90 % of the raphe in cats inducing complete insomnia for 3-4 days.  Slow-wave, but not REM sleep, gradually returned, but never exceeded 2.5 hours a day.  Sleep amount correlates to the 5-HT concentration.  The neurons of the raphe nuclei are rich in serotonin (5-HT).  It is suspected that the dorsal raphe nuclei inhibit the phasic components of REM sleep and thus prevent them from occurring at inappropriate times.  The activity of serotonergic neurons is low during REM sleep and higher at other times.  The dorsal raphe nucleus has been associated with 
  • Depression.  
  • Impulsive aggression which is associated with relatively low serotonin levels.  
projects serotonin signals, is an emergent capability which is used by cooperating agents to support coordination & rival agents to support control and dominance.  In eukaryotic cells signalling is used extensively.  A signal interacts with the exposed region of a receptor molecule inducing it to change shape to an activated form.  Chains of enzymes interact with the activated receptor relaying, amplifying and responding to the signal to change the state of the cell.  Many of the signalling pathways pass through the nuclear membrane and interact with the DNA to change its state.  Enzymes sensitive to the changes induced in the DNA then start to operate generating actions including sending further signals.  Cell signalling is reviewed by Helmreich.  Signalling is a fundamental aspect of CAS theory and is discussed from the abstract CAS perspective in signals and sensors.  In AWF the eukaryotic signalling architecture has been abstracted in a codelet based implementation.  To be credible signals must be hard to fake.  To be effective they must be easily detected by the target recipient.  To be efficient they are low cost to produce and destroy. 
to the tegmentum is involved in homeostatic and reflex pathways in the brain.  It includes: ventral; areas.  It forms the base of the midbrain.  It projects via the: mesolimbic dopamine pathway to limbic structures, mesocortical dopamine pathway to the prefrontal cortex.  Sapolsky explains that Wolfram Schultz demonstrated that following a reward, the dopamine system codes for discrepancy from expectation--get what you expected, and there's a steady-state dribble of dopamine.  Get more reward and/or get it sooner than expected, and there's a big burst; less and or later a decrease.  Some tegmental neurons respond to positive discrepancy from expectation, others to negative; appropriately, the latter are local neurons that release inhibitory neurotransmitter GABA.  Those same neurons participate in habituation, where the reward that once elicited a big dopamine response becomes less exciting.  These coding neurons get projections from the frontal cortex where the expectancy/discrepancy calculations occur. 
, accumbens is a region of the basal forebrain rostral to the preoptic area and immediately adjacent to the septum.  The nucleus accumbens was closely associated with the limbic system and plays an important role in reinforcement.  If a rodent wins a fight on his home territory, there are long-lasting increases in levels of testosterone receptors enhancing pleasurable effects.  When prairie voles first mate, epi-genetic state changes are induced in the accumbens to support pair-bonding.  The accumbens projects to brain regions associated with movement.  The major pathways of dopaminergic neurons begin in the substantia nigra and the ventral tegmental area.  The amygdala projects back to the accumbens.  The tegmentostriatal system begins in the ventral tegmental area and projects to the nucleus accumbens.  The nucleus accumbens includes high levels of D1, D2 and D3 dopamine receptors located on the spine & shafts of dendrites of excitatory cells reduce the transfer of excitation from the dendrites to the cell bodies, so only especially strong excitatory inputs get through to the cell body to elicit excitation.  It also has D4 dopamine receptors which are highly variable.  The accumbens responds differently to rewards depending on maturity: In juveniles all reward levels result in the same response activity level, During adolescence the accumbens responds to small rewards negatively, and large rewards hugely, In adults the rewards result in measured scaled positive responses.  Chronic stress depletes dopamine from the nucleus accumbens biasing humans towards depression.  , prefrontal cortex (PFC) is
  • The front part of the frontal lobe of the cerebral cortex.  It evolved most recently.  During adolescence when the PFC is still deploying, older brain agents provide equivalent strategies: ventral striatum.  The PFC has been implicated in planning, working memory: dorsolateral; decision making: Orbitofrontal cortex; and social behavior.  Different PFC circuits track internal reward driven strategies and externally signalled advice.  The PFC chooses between conflicting options, letting go or restraint, especially between cognition and emotions.  It imposes an overarching strategy for managing working memory.  It is essential for thinking about multiple items with different labels.  It includes neurons that are interested in particular sub-categories: Dog, Cat.  Once it has made a decision it signals the rest of the frontal lobe just behind it.  Glucocorticoids decrease excitability of the PFC.  
and amygdala contains > 12 distinct areas: Central, Lateral.  It receives simple signals from the lower parts of the brain: pain from the PAG; and abstract complex information from the highest areas: Disgust from the insula cortex.  It sends signals to almost every other part of the brain, including to the decision-making circuitry of the frontal lobes.  It has high levels of D(1) dopamine receptors.  During extreme fear the amygdala drives the hippocampus into fear learning.  It outputs directly to subcortical reflexive motor pathways when speed is required.  Its central nucleus projects to the BNST.  It signals the locus ceruleus.  The amygdala:
  • Promotes aggression.  Stimulating the amygdala promotes rage.  It converts anger into aggression and when impaired it impacts the ability to detect angry facial expressions.  
  • Participates in disgust
  • Perceives fear promoting stimuli.  In PTSD sufferers the Amygdala overreacts to mildly fearful stimuli and is slow to calm down and the amygdala expands in size over a period of months.  Fear is processed by the lateral nucleus which serves as the input from various senses, and the central nucleus which outputs to the brain stem (central grey - freezing, lateral hypothalamus - blood pressure, activates paraventricular hypothalamus => crf -> hormone adjustments). 
  • Has lots of receptors for and is highly sensitive to glucocorticoids.  Stress inhibits the GABA interneurons in the basolateral amygdala (BLA) allowing the excitatory glutamate releasing neurons to excite more. 
  • Is sensitive to unsettling/uncertain social situations where it promotes anxiety.  It is also interested in uncertain but potentially painful situations.  The amygdala contributes to social and emotional decision making where the BLA supports rejecting an unacceptable offer, as allowed in the Ultimatum Game, by injecting implicit mistrust and vigilance, generating an anger driven rejection that is used as punishment.  The amygdala is very rapidly excited by subliminal signals from the thalamus of outgroup skin color.  The amygdala subsequently tips social emotions against outgroups unless restrained by the frontal lobe or influenced by subliminal priming to prioritize inclusion.  The fast path from the thalamus rapidly but inaccurately signals its identified a weapon. 
  • Promotes male, but not female, sexual motivation when it is an uncertain potential pleasure. 
  • Responds to the longing for uncertain potential pleasures and fear that the reward will not be worth it if it happens.  The amygdala turns off during orgasm. 
  • Uses but is not directly involved in vision. 
where the serotonin enhances dopamine is a synaptic signal supporting generalized goal-directed behavior & anticipation of reward.  Its significance is that the receptors that detect the signal are of the slow acting type and are used to alter (modulate) the response of fast acting dopaminergic neural circuits in which the receptors are deployed (LTP).  The signal detects significant changes including predictions of models and actual results which differ unexpectedly.  The dopamine network architecture is designed to signal the possibility of any type of reward: Norm violation punishment, Winning a lottery, & Misfortune of an envied competitor.  Dopamine signalling:
  • Rescales continuously to accommodate the range of intensity offered by different stimuli.  So dopamine's responses to any reward habituate.  GABA is released by some tegmental neurons to induce habituation. 
  • Reflects the anticipation of reward.  It supports establishment of a relationship between a signal, working for a reward and obtaining the reward, but subsequently dopamine is mainly released encouraging the work, right after the signal supporting anticipation of the reward.  Anticipation requires learning and is reflected in hippocampus activity.  That explains context dependent cravings.  And the learning architecture means reliable cues become rewarding.  The accumbens supports willpower.  And dopamine
  • Promotes goal-oriented behavior needed to obtain & likely to achieve the reward - through the dopamine projections to the prefrontal cortex.  That makes dopamine central to:
    • Motivation.  This binding fails in depression - due to stress and in anxiety - due to signals from the amygdala.  
    • The prefrontal cortex's mesocortically stimulated support for willpower to act to delay rewards.  To sustain work for delayed rewards additional dopamine is released based on the length of the delay and the rewards uncertainty (modelled in the dorsolateral prefrontal cortex - which promotes the long term and the ventromedial prefrontal cortex - which promotes the short term) and the anticipated size of the reward (modelled in the accumbens).  Impulsiveness in ADHD is reflected in abnormal dopamine processing.  Addictive drugs bias the dopamine network towards impulsiveness.  
  • Is lowered by certain gene variants which induce: less dopamine in the synapse, fewer receptors, lower responsiveness of receptors; associated with (as tiny effects in hugely varying social scenarios): sensation seeking, risk taking, attentional problems, extroversion; where:
    • The receptor D4's gene shows high variability.  The D47R form is relatively unresponsive to dopamine.  
    • Dopamine is degraded by COMT.  The COMT gene includes a variant which is highly efficient reducing dopamine signalling but with complicating gene/environment interactions.  
    • Dopamine is removed from the synapse by a reuptake transporter DAT. 
's
Read Montague explores how brains make decisions.  In particular he explains how:
  • Evolution can create indirect abstract models, such as the dopamine system, that allow
  • Life changing real-time decisions to be made, and how
  • Schematic structures provide encodings of computable control structures which operate through and on incomputable, schematically encoded, physically active structures and operationally associated production functions. 
effects on goal-directed behavior


Seconds to minutes before
Having reviewed what the nervous system was doing to generate our best and worst behaviors Sapolsky now aims to explore:
  • What outside stimulus, acting through what sensory channel and targeting which parts of the brain, prompted this behavior?
  • Were you aware of that environmental stimulus?
  • What stimuli had your brain made you particularly sensitive to?
  • What does this tell us about our best and worst behaviors?
Huge variety of external and internal cues impact behavior and many of these signals are subliminal.  This makes us less rational and autonomous decision makers than we typically assume. 

Sapolsky repeats that an animal's dominant sensory channel has the most direct access to the limbic system supports emotional circuits: Amygdala, Hippocampus, Septum, Habenula, Mammillary bodies; all of which signals the midbrain through the hypothalamus. The broad interconnections of these regions with a part of the frontal lobe suggested to Walle Nauta that it (ventromedial prefrontal cortex) is a quasi-member of the limbic network.  .  And he stresses that subliminal sensory cues are hugely significant. 

Pertinent to our best and worst behaviors our brains are highly attuned to skin color, noting within a hundred milliseconds:
  1. Amygdala contains > 12 distinct areas: Central, Lateral.  It receives simple signals from the lower parts of the brain: pain from the PAG; and abstract complex information from the highest areas: Disgust from the insula cortex.  It sends signals to almost every other part of the brain, including to the decision-making circuitry of the frontal lobes.  It has high levels of D(1) dopamine receptors.  During extreme fear the amygdala drives the hippocampus into fear learning.  It outputs directly to subcortical reflexive motor pathways when speed is required.  Its central nucleus projects to the BNST.  It signals the locus ceruleus.  The amygdala:
    • Promotes aggression.  Stimulating the amygdala promotes rage.  It converts anger into aggression and when impaired it impacts the ability to detect angry facial expressions.  
    • Participates in disgust
    • Perceives fear promoting stimuli.  In PTSD sufferers the Amygdala overreacts to mildly fearful stimuli and is slow to calm down and the amygdala expands in size over a period of months.  Fear is processed by the lateral nucleus which serves as the input from various senses, and the central nucleus which outputs to the brain stem (central grey - freezing, lateral hypothalamus - blood pressure, activates paraventricular hypothalamus => crf -> hormone adjustments). 
    • Has lots of receptors for and is highly sensitive to glucocorticoids.  Stress inhibits the GABA interneurons in the basolateral amygdala (BLA) allowing the excitatory glutamate releasing neurons to excite more. 
    • Is sensitive to unsettling/uncertain social situations where it promotes anxiety.  It is also interested in uncertain but potentially painful situations.  The amygdala contributes to social and emotional decision making where the BLA supports rejecting an unacceptable offer, as allowed in the Ultimatum Game, by injecting implicit mistrust and vigilance, generating an anger driven rejection that is used as punishment.  The amygdala is very rapidly excited by subliminal signals from the thalamus of outgroup skin color.  The amygdala subsequently tips social emotions against outgroups unless restrained by the frontal lobe or influenced by subliminal priming to prioritize inclusion.  The fast path from the thalamus rapidly but inaccurately signals its identified a weapon. 
    • Promotes male, but not female, sexual motivation when it is an uncertain potential pleasure. 
    • Responds to the longing for uncertain potential pleasures and fear that the reward will not be worth it if it happens.  The amygdala turns off during orgasm. 
    • Uses but is not directly involved in vision. 
    activation and more so in someone who tests positive for race bias
  2. Fusiform is a region of the brain which supports advanced mechanisms of shape recognition and implements the early stages of reading.  Subliminal priming with words did not depend on the shape of the word.  The fusiform gyrus was able to process the abstract identity of a word without caring if it was upper or lower case.   While high up in the cortex it can operate below the level of conscious experience.  It contributes to social emotions with:
    • Its face area being more activated by faces with in-group skin color.  
    • It activating when shown pictures of cars in automobile aficionados. 
    • It activating when shown pictures of birds in birdwatchers; since it really recognizes examples of items from an individual's emotionally salient categories. 
    face area is less activated by other-race faces. 
The medial prefrontal cortex in Buckner's fMRI based analysis is part of the brain's default mode network.  It facilitates the flexible use of information during the construction of self-relevant mental simulations.  Nauta considered its ventromedial part as participating in the limbic system.  With the medial temporal lobe it converges on important integration nodes: posterior cingulate cortex.   differentially activates when considering misfortune to members of one's own race. 

Eyes are major sensors in primates, based on opsins deployed in the retina & especially fovea, signalling the visual system: Superior colliculi, Thalamus (LGN), Primary visual cortex; and indirectly the amygdala.  They also signal [social] emotional state to other people.  And they have implicit censorious power with pictures of eyes encouraging people within their view to act more honorably.  Eyes are poor scanners and use a saccade to present detail slowly to the fovea.  The eye's optical structures and retina are supported by RPE.  Eyes do not connect to the brain through the brain stem and so still operate in locked-in syndrome.  Evo-devo shows eyes have deep homology.  High pressure within the eye can result in glaucoma.  Genetic inheritance can result in retinoblastoma.  Age is associated with AMD. 
and then faces present the most significant subliminal signals for watching humans.  Emotion are low level agents distributed across the brain and body which associate, via the amygdala and rich club hubs, important environmental signals with encoded high speed sensors, and distributed programs of action to model: predict, prioritize guidance signals, select and respond effectively, coherently and rapidly to the initial signal.  The majority of emotion centered brain regions interface to the midbrain through the hypothalamus.  The most accessible signs of emotions are the hard to control and universal facial expressions.  Emotions provide prioritization for conscious access given that an animal has only one body, but possibly many cells, with which to achieve its highest level goals.  Because of this emotions clash with group goals and are disparaged by the powerful.  Evolutionary psychology argues evolution shaped human emotions during the long period of hunter-gatherer existence in the African savanna.  Human emotions are universal and include: Anger, Appreciation of natural beauty, Disgust, Fear, Gratitude, Grief, Guilt, Happiness, Honor, Jealousy, Liking, Love, Rage, Romantic love, Lust for revenge, Passion, Sadness, Self-control, Shame, Sympathy, Surprise; and the sham emotions and distrust induced by reciprocal altruism.   is discerned primarily from the eyes. 

Subliminal auditory cues also alter behavior in a racially biased way. 

Subliminal smell is still significant in humans.  The olfactory network directly projects to the limbic system: 

Beauty is accepted cross-culturally is how we do and think about things, transmitted by non-genetic means as defined by Frans de Waal.  CAS theory views cultures as operating via memetic schemata evolved by memetic operators to support a cultural superorganism.  Evolutionary psychology asserts that human culture reflects adaptations generated while hunting and gathering.  Dehaene views culture as essentially human, shaped by exaptations and reading, transmitted with support of the neuronal workspace and stabilized by neuronal recycling.  Sapolsky argues that parents must show children how to transform their genetically derived capabilities into a culturally effective toolset.  He is interested in the broad differences across cultures of: Life expectancy, GDP, Death in childbirth, Violence, Chronic bullying, Gender equality, Happiness, Response to cheating, Individualist or collectivist, Enforcing honor, Approach to hierarchy; illustrating how different a person's life will be depending on the culture where they are raised.  Culture:
  • Is deployed during pregnancy & childhood, with parental mediation.  Nutrients, immune messages and hormones all affect the prenatal brain.  Hormones: Testosterone with anti-Mullerian hormone masculinizes the brain by entering target cells and after conversion to estrogen binding to intracellular estrogen receptors; have organizational effects producing lifelong changes.  Parenting style typically produces adults who adopt the same approach.  And mothering style can alter gene regulation in the fetus in ways that transfer epigenetically to future generations!  PMS symptoms vary by culture. 
  • Is also significantly transmitted to children by their peers during play.  So parents try to control their children's peer group.  
  • Is transmitted to children by their neighborhoods, tribes, nations etc. 
  • Influences the parenting style that is considered appropriate. 
  • Can transform dominance into honor.  There are ecological correlates of adopting honor cultures.  Parents in honor cultures are typically authoritarian. 
  • Is strongly adapted across a meta-ethnic frontier according to Turchin.  
  • Across Europe was shaped by the Carolingian empire. 
  • Can provide varying levels of support for innovation.  
  • Produces consciousness according to Dennet. 
, and from an early age, as indicating, through the medial orbitofrontal cortex is a rich club hub region of the prefrontal cortex, involved in representing emotion & reward in decision-making.  It is positioned immediately above the orbits where the eyes sit.  The medial part judges the beauty of faces, minds and acts and will conflate them in social emotion evaluations.  The orbitofrontal cortex receives projections from the: hippocampus & associated areas of the cingulate, retrosplenial & entorhinal cortices, anterior thalamus, amygdala, midline thalamus, non-isocortical insula, mediodorsal thalamus.  , smarter, kinder & more honest people. 

Body posture is rapidly analyzed, signalling, is an emergent capability which is used by cooperating agents to support coordination & rival agents to support control and dominance.  In eukaryotic cells signalling is used extensively.  A signal interacts with the exposed region of a receptor molecule inducing it to change shape to an activated form.  Chains of enzymes interact with the activated receptor relaying, amplifying and responding to the signal to change the state of the cell.  Many of the signalling pathways pass through the nuclear membrane and interact with the DNA to change its state.  Enzymes sensitive to the changes induced in the DNA then start to operate generating actions including sending further signals.  Cell signalling is reviewed by Helmreich.  Signalling is a fundamental aspect of CAS theory and is discussed from the abstract CAS perspective in signals and sensors.  In AWF the eukaryotic signalling architecture has been abstracted in a codelet based implementation.  To be credible signals must be hard to fake.  To be effective they must be easily detected by the target recipient.  To be efficient they are low cost to produce and destroy. 
social status is a publically accepted, signal that one possesses assets: wealth, beauty, talent, expertise, access & trust of powerful people; to be able to help others. 
& dominance signals the power to hurt a rival.  Maynard Smith & Parker explain that in group situations females compete for food and males compete for females.  Maleness is a huge factor for violence.  Fighting to the death is costly for all participants so instead they indicate:
  • Size and weapons to demonstrate who will win.  Males who are, or look like, better fighters: Large heads, Big men, Height; gain in dominance. 
  • Political acumen to demonstrate they won't be pushed around and have the support of other powerful groups.  Dominant males push other rivals aside and gain interest of females, enabling themselves to replicate more.  Being a signal its authenticity can be challenged and so must be defended to remain credible.  Hotheads leverage the doomsday machine to constrain rational challenges.  Bands and cultures leverage honor.  Youth and lack of resources reduce the power of rivals' political constraints. 
helping support stable social hierarchies

Interoceptive signals indicate the body's internal state: Pain, Fatigue; seconds to minutes before.  The interoceptive 'networks' project to brain regions that implement social emotions. 
also influence our behaviors.  Sapolsky notes evidence supporting the James-Lange theory developed by William James & Carl Lange who, independently proposed that humans decide how they feel based on interoceptive signals, rather than from external ones.  Sapolsky notes the theory reflects these signals influence on behavior where social emotion processing areas receive lots of projections from interoceptive networks.  The most notable is pain.  But he writes it fails to support specificity or speed of reactions since autonomic responses are too slow to precede awareness of an emotion. 
, that these internal feelings shape your interpretation of external signals and your emotions.  But he notes issues of specificity and speed that make these influences rather than determinants of emotions. 

Words have significant behavioral influence.  Subliminal prime flashes a subliminal word or picture, the prime, immediately before another visible item, the target.  Experiments demonstrate that the presence of the subliminal prime speeds up conscious processing when the prime is represented consciously within a second of the initial showing. 
words shift behavior to be more or less prosocial is prioritizing benefiting others through: Help, Charity, Truth; even if many are acts of restitution to balance out antisocial acts.  Due to the Insula's binding of physical and metaphorical disgust, physically washing your hands can be enough to reduce the need for a prosocial act.  
Kahneman & Tversky showed word framing related to gain or loss is processed differently in decision making.  Kahneman & Tversky asked participants in an experiment, including doctors, to decide whether to administer a hypothetical drug.  When to the drug:
  • Has a 95 percent survival rate, the participants were more likely to approve its use than if it
  • Has a 5 percent death rate. 
can alter decision making. 

Symbols: National flag, Team shirt; also influence behavior.  Cues about group identity demonstrate the complex nature of the signal processing.  Asian American women primed to think about racial identity performed better in a mathematics test than those primed to think about gender.  Sapolsky notes that their assumptions about females struggling at mathematics depend on cultural influences

When women are present, or men are prompted to think about women, men increase their risk-taking, spend more on luxuries that signal status, and act more aggressively.  Except when men's status has been achieved prosocially in which case the presence of women makes the men more prosocial. 

The physical environment also signals changes in behavior: Broken windows is a 1982 crime theory of James Wilson and George Kelling, that was inspired by the results of vandalism experiments by Philip Zimbardo, which assumes urban disorder and vandalism undermine social constraints and encourage additional criminal and anti-social behavior.  By detecting and responding to petty vandalism rapidly it is assumed that more serious crime will be discouraged.  In New York Broken Windows was used to justify 'stop, question and frisk'.  However, Donohue & Levitt's analysis suggests an observed precipitous drop in crime rates depended on legal access to abortion. 
; which encouraged zero-tolerance policing in New York City resulting is reductions in crime. 

Sapolsky raises a complicating detail: the brain can
Agents use sensors to detect events in their environment.  This page reviews how these events become signals associated with beneficial responses in a complex adaptive system (CAS).  CAS signals emerge from the Darwinian information model.  Signals can indicate decision summaries and level of uncertainty. 
alter the sensitivity of specific sensory modalities
making some signals more influential.  Examples include: Directional sensing, Stress is a multi-faceted condition reflecting high cortisol levels.  Dr. Robert Sapolsky's studies of baboons indicate that stress helps build readiness for fight or flight.  As these actions occur the levels of cortisol return to the baseline rate.  A stressor is anything that disrupts the regular homeostatic balance.  The stress response is the array of neural and endocrine changes that occur to respond effectively to the crisis and reestablish homeostasis. 
  • The short term response to the stressor
    • activates the amygdala which: Stimulates the brain stem resulting in inhibition of the parasympathetic nervous system and activation of the sympathetic nervous system with the hormones epinephrine and norepinephrine deployed around the body, Activates the PVN which generates a cascade resulting in glucocorticoid secretion to: get energy to the muscles with increased blood pressure for a powerful response.  The brain's acuity and cognition are stimulated.  The immune system is stimulated with beta-endorphin and repair activities curtail.  But when the stressor is
  • long term: loneliness, debt; and no action is necessary, or possible, long term damage ensues.  Damage from such stress may only occur in specific situations: Nuclear families coping with parents moving in.  Sustained stress provides an evolved amplifier of a position of dominance and status.  It is a strategy in female aggression used to limit reproductive competition.  Sustained stress:
    • Stops the frontal cortex from ensuring we do the harder thing, instead substituting amplification of the individual's propensity for risk-taking and impairing risk assessment! 
    • Activates the integration between the thalamus and amygdala. 
      • Acts differently on the amygdala in comparison to the frontal cortex and hippocampus: Stress strengthens the integration between the Amygdala and the hippocampus, making the hippocampus fearful. 
      • BLA & BNST respond with increased BDNF levels and expanded dendrites persistently increasing anxiety and fear conditioning. 
    • Makes it easier to learn a fear association and to consolidate it into long-term memory.  Sustained stress makes it harder to unlearn fear by making the prefrontal cortex inhibit the BLA from learning to break the fear association and weakening the prefrontal cortex's hold over the amygdala.  And glucocorticoids decrease activation of the medial prefrontal cortex during processing of emotional faces.  Accuracy of assessing emotions from faces suffers.  A terrified rat generating lots of glucocorticoids will cause dendrites in the hippocampus to atrophy but when it generates the same amount from excitement of running on a wheel the dendrites expand.  The activation of the amygdala seems to determine how the hippocampus responds. 
    • Depletes the nucleus accumbens of dopamine biasing rats toward social subordination and biasing humans toward depression. 
    • Disrupts working memory by amplifying norepinephrine signalling in the prefrontal cortex and amygdala to prefrontal cortex signalling until they become destructive.  It also desynchronizes activation in different frontal lobe regions impacting shifting of attention. 
  • During depression, stress inhibits dopamine signalling. 
  • Strategies for stress reduction include: Mindfulness. 
driven alterations, Hunger based increases in sensitivity to smell of food.  There are more projections from the higher centers of the brain back to the sensory networks allowing modulation of the sensors operations.  Such complex interactions result in individuals from collectivist cultures looking at a complicated picture with foreground and background details differently to those from individualist cultures. 

Hours to days before
This is the timescale where hormones are signalling molecules: ACTH, TRH, Melanocyte stimulating hormone, Testosterone, Oxytocin, Vasopressin, Insulin, Growth hormone, Estrogen, Progesterone, Angiotensin II, Asprosin, EPO, Irisin, Leptin, FGF21 hormone, Prostaglandins, TSH, Thyroxine, Glococorticoids; that are transported by the circulatory system to interact with target organs having appropriate receptors.  The levels of hormones can fluctuate massively, as in pregnancy. 
influence the brain regions and sensory networks discussed previously.  The effects are mainly contingent and facilitative increasing sensitivity to preexisting tendencies setup over the longer term via adult neuroplasticity and adolescence and childhood and genetics and proximate environment including culture is how we do and think about things, transmitted by non-genetic means as defined by Frans de Waal.  CAS theory views cultures as operating via memetic schemata evolved by memetic operators to support a cultural superorganism.  Evolutionary psychology asserts that human culture reflects adaptations generated while hunting and gathering.  Dehaene views culture as essentially human, shaped by exaptations and reading, transmitted with support of the neuronal workspace and stabilized by neuronal recycling.  Sapolsky argues that parents must show children how to transform their genetically derived capabilities into a culturally effective toolset.  He is interested in the broad differences across cultures of: Life expectancy, GDP, Death in childbirth, Violence, Chronic bullying, Gender equality, Happiness, Response to cheating, Individualist or collectivist, Enforcing honor, Approach to hierarchy; illustrating how different a person's life will be depending on the culture where they are raised.  Culture:
  • Is deployed during pregnancy & childhood, with parental mediation.  Nutrients, immune messages and hormones all affect the prenatal brain.  Hormones: Testosterone with anti-Mullerian hormone masculinizes the brain by entering target cells and after conversion to estrogen binding to intracellular estrogen receptors; have organizational effects producing lifelong changes.  Parenting style typically produces adults who adopt the same approach.  And mothering style can alter gene regulation in the fetus in ways that transfer epigenetically to future generations!  PMS symptoms vary by culture. 
  • Is also significantly transmitted to children by their peers during play.  So parents try to control their children's peer group.  
  • Is transmitted to children by their neighborhoods, tribes, nations etc. 
  • Influences the parenting style that is considered appropriate. 
  • Can transform dominance into honor.  There are ecological correlates of adopting honor cultures.  Parents in honor cultures are typically authoritarian. 
  • Is strongly adapted across a meta-ethnic frontier according to Turchin.  
  • Across Europe was shaped by the Carolingian empire. 
  • Can provide varying levels of support for innovation.  
  • Produces consciousness according to Dennet. 


Sapolsky's discusses testosterone is a hormone secreted by the testes in response to stimulation from the hypothalamic/pituitary/testicular cascade, that makes humans more willing to do what it takes to attain and maintain status, according to Sapolsky.  That means players of the Ultimatum Game, if previously given testosterone can become more generous.  Testosterones effect is highly socially contextual so it may encourage acts of kindness or aggression (when challenged).  The level of testosterone does not predict which individuals will be aggressive in: Birds, Fish, Mammals including primates.  Genes impact the potency of testosterone by altering the enzymes that: Construct it, Convert it to estrogen, code the androgen receptor.   This androgen receptor includes a variable polyglutamine repeat which alters the sensitivity to the testosterone signal.  The more potent form is associated with boys showing more dramatic 'masculinization' of the cortex.  But the detected genetic influences are small.  Testosterone decreases activity in the prefrontal cortex and its functional coupling to the amygdala while increasing the coupling between the amygdala & the thalamus.  Testosterone shortens the refactory period of amygdaloid & amygdaloid target neurons.  This results in impulsive risk taking and more focus on unfamiliar faces and distrust of them.  Testosterone increases activity in the ventral tegmentum projecting dopamine to enhance place preference.  Winners of fights become more willing to fight in part due to testosterone increasing confidence and optimism and reducing fear and anxiety.  And winning at: Chess, Athletics, Stock trades; induces the BNST to add testosterone receptors increasing its sensitivity to the hormone.  People become overconfident and overly optimistic. 
which he asserts is far less relevant to aggression than has been assumed and oxytocin is a peptide hormone which makes humans more prosocial to and socially competent in their in-group and more antisocial to everyone else.  The effects are contingent; changing during stress and in the presence of a threatening outgroup.  Oxytocin makes people look at eyes longer, encouraging improved accuracy at perceiving emotions.  It enhances activity in the TPJ supporting modeling of other people's thinking.  Dogs and their owners secrete oxytocin increasing the amount of eye contact between them.  It is associated with pair bonding.  It is central to female mammals wanting to nurse, nursing, and remembering their child.  Its effects are context dependent and so is the regulation of the genes that control oxytocin.  Variants of a gene CD38 which facilitates oxytocin secretion from neurons are associated with differing levels of activation of the fusiform face area when looking at faces.  Sapolsky describes an oxytocin receptor gene variant that is associated with children showing: Extreme aggression, A callous unemotional style; foreshadowing adult psychopathy.  And another receptor gene variant which is associated with childhood social disconnection and unstable adult relationships.  Gene/environment interactions complicate the interpretation of the presence of particular gene variants.  Hypothalamic neurons send projections to: ventral tegmentum which also becomes more receptive during child birth, nucleus accumbens, hippocampus, amygdala where it inhibits the central amygdala suppressing fear & anxiety consistently in men while still allowing women to respond to threats to their infants, frontal cortex, olfactory network where it helps new rat mums to learn the smell of their offspring; where oxytocin prepares the brain for in-group bonding, out-grouping, birth and maternal behavior.  Outside the brain hypothalamic neurons in females send oxytocin to the posterior pituitary where it enters the blood stream stimulating uterine contraction during labor & supporting milk production for weaning.  Disorders associated with oxytocin abnormalities include ASD. 
that is not as prosocial is prioritizing benefiting others through: Help, Charity, Truth; even if many are acts of restitution to balance out antisocial acts.  Due to the Insula's binding of physical and metaphorical disgust, physically washing your hands can be enough to reduce the need for a prosocial act.  
as often claimed. 

Males have more testosterone and are more aggressive but Sapolsky notes that aggression occurs even when castration has removed testosterone from males.  Sapolsky concludes aggression is typically more about social learning than testosterone.  We reward aggression too often.  Testosterone has effects but they are hugely context dependent and typically amplify preexisting tendencies.  Testosterone boosts impulsivity, risk taking and feeling good from such activities.  It also makes winning a fight feel good.  And it supports the formation of a dominance hierarchy and encourages us to maintain status is a publically accepted, signal that one possesses assets: wealth, beauty, talent, expertise, access & trust of powerful people; to be able to help others. 
.  Only during a challenge does an increases in testosterone make aggression more likely.  When a person's pride depends on being prosocial rising testosterone will increase acts of kindness. 

Oxytocin became interesting to neurobiologists when it was realized the hypothalamic is essential to many operations of the body.  It has many small sub-regions whose main functions are to regulate hunger, thirst, temperature, sexual behavior, and similar body operations.  Its (paraventricular nucleus) is closely connected to the pituitary which secrets hormones into the bloodstream ( => acth -> adrenal cortex => cortisol (+)->  amygdala & (-)-> hippocampus).   neurons, specialized eukaryotic cells include channels which control flows of sodium and potassium ions across the massively extended cell membrane supporting an electro-chemical wave which is then converted into an outgoing chemical signal transmission from synapses which target nearby neuron or muscle cell receptors.  Neurons are supported by glial cells.  Neurons include a:
  • Receptive element - dendrites
  • Transmitting element - axon and synaptic terminals
  • Highly variable DNA schema using transposons. 
that sent axons, a long extension of a neuron which has a membrane constructed to support the uni-directional flow of action potential from the dendritic tree and cell body to the synaptic terminals.   to the posterior pituitary also sent projections throughout the brain, including to the: ventral tegmentum is part of the tegmentum which contains dopamine long system cell bodies (high levels of D(2)).  Their axons branch extensively and reach many areas including: Mesolimbic to the limbic system: amygdala, hippocampus; Tegmentostriatal to the nucleus accumbens, Mesocortical to the forebrain including the prefrontal cortex.  The terminals are fairly evenly distributed through out layers 1 - 6.  So dopamine can modulate input and output excitatory and inhibitory transmissions.  If a rodent wins a fight on his home territory, there are long-lasting increases in levels of testosterone receptors enhancing pleasurable effects.  The lateral ventral tegmental area is now known to be one of two large adrenergic pathways, along with the locus ceruleus.  During child birth the ventral tegmentum deploys more oxytocin receptors increasing its sensitivity to the neuropeptide.  
, accumbens is a region of the basal forebrain rostral to the preoptic area and immediately adjacent to the septum.  The nucleus accumbens was closely associated with the limbic system and plays an important role in reinforcement.  If a rodent wins a fight on his home territory, there are long-lasting increases in levels of testosterone receptors enhancing pleasurable effects.  When prairie voles first mate, epi-genetic state changes are induced in the accumbens to support pair-bonding.  The accumbens projects to brain regions associated with movement.  The major pathways of dopaminergic neurons begin in the substantia nigra and the ventral tegmental area.  The amygdala projects back to the accumbens.  The tegmentostriatal system begins in the ventral tegmental area and projects to the nucleus accumbens.  The nucleus accumbens includes high levels of D1, D2 and D3 dopamine receptors located on the spine & shafts of dendrites of excitatory cells reduce the transfer of excitation from the dendrites to the cell bodies, so only especially strong excitatory inputs get through to the cell body to elicit excitation.  It also has D4 dopamine receptors which are highly variable.  The accumbens responds differently to rewards depending on maturity: In juveniles all reward levels result in the same response activity level, During adolescence the accumbens responds to small rewards negatively, and large rewards hugely, In adults the rewards result in measured scaled positive responses.  Chronic stress depletes dopamine from the nucleus accumbens biasing humans towards depression.  , hippocampus is a part of the brain involved in the temporary storage or coding of long-term episodic memory.  Memory formation in the cells of the hippocampus uses the MAP kinase signalling network which is impacted by sleep deprivation.  The hippocampus dependent memory system is directly affected by cholinergic changes throughout the wake-sleep cycle.  Increased acetylcholine during REM sleep promotes information attained during wakefulness to be stored in the hippocampus by suppressing previous excitatory connections while facilitating encoding without interference from previously stored information.  During slow-wave sleep low levels of acetylcholine cause the release of the suppression and allow for spontaneous recovery of hippocampal neurons resulting in memory consolidation.  It was initially associated with memory formation by McGill University's Dr. Brenda Milner, via studies of 'HM' Henry Molaison, whose medial temporal lobes had been surgically destroyed leaving him unable to create new memories.  The size of neurons' dendritic trees expands and contracts over a female rat's ovulatory cycle, with the peak in size and cognitive skills at the estrogen high point.  Adult neurogenesis occurs in the hippocampus (3% of neurons are replaced each month) where the new neurons integrate into preexisting circuits.  It is enhanced by learning, exercise, estrogen, antidepressants, environmental enrichment, and brain injury and inhibited by various stressors explains Sapolsky.  Prolonged stress makes the hippocampus atrophy.  He notes the new neurons are essential for integrating new information into preexisting schemas -- learning that two things you thought were the same are actually different. 
, amygdala contains > 12 distinct areas: Central, Lateral.  It receives simple signals from the lower parts of the brain: pain from the PAG; and abstract complex information from the highest areas: Disgust from the insula cortex.  It sends signals to almost every other part of the brain, including to the decision-making circuitry of the frontal lobes.  It has high levels of D(1) dopamine receptors.  During extreme fear the amygdala drives the hippocampus into fear learning.  It outputs directly to subcortical reflexive motor pathways when speed is required.  Its central nucleus projects to the BNST.  It signals the locus ceruleus.  The amygdala:
  • Promotes aggression.  Stimulating the amygdala promotes rage.  It converts anger into aggression and when impaired it impacts the ability to detect angry facial expressions.  
  • Participates in disgust
  • Perceives fear promoting stimuli.  In PTSD sufferers the Amygdala overreacts to mildly fearful stimuli and is slow to calm down and the amygdala expands in size over a period of months.  Fear is processed by the lateral nucleus which serves as the input from various senses, and the central nucleus which outputs to the brain stem (central grey - freezing, lateral hypothalamus - blood pressure, activates paraventricular hypothalamus => crf -> hormone adjustments). 
  • Has lots of receptors for and is highly sensitive to glucocorticoids.  Stress inhibits the GABA interneurons in the basolateral amygdala (BLA) allowing the excitatory glutamate releasing neurons to excite more. 
  • Is sensitive to unsettling/uncertain social situations where it promotes anxiety.  It is also interested in uncertain but potentially painful situations.  The amygdala contributes to social and emotional decision making where the BLA supports rejecting an unacceptable offer, as allowed in the Ultimatum Game, by injecting implicit mistrust and vigilance, generating an anger driven rejection that is used as punishment.  The amygdala is very rapidly excited by subliminal signals from the thalamus of outgroup skin color.  The amygdala subsequently tips social emotions against outgroups unless restrained by the frontal lobe or influenced by subliminal priming to prioritize inclusion.  The fast path from the thalamus rapidly but inaccurately signals its identified a weapon. 
  • Promotes male, but not female, sexual motivation when it is an uncertain potential pleasure. 
  • Responds to the longing for uncertain potential pleasures and fear that the reward will not be worth it if it happens.  The amygdala turns off during orgasm. 
  • Uses but is not directly involved in vision. 
& frontal cortex of the cerebral cortex is at the front of the brain.  It includes the: prefrontal cortex, motor cortex.  Sapolsky asserts it makes you do the harder thing when it's the right thing to do.  The frontal cortex supports working memory to sustain focus on a task.  It also coordinates the strategic actions necessary to achieve success.  It provides impulse control, regulation of emotion, and willpower.  The prefrontal cortex maintains focus by deprioritizing currently irrelevant streams of information.  The frontal cortex tracks rules.  Over a lifetime that builds into a costly activity.  Once it tires responses become less prosocial.  But practice shifts operation of tasks to the cerebellum.  The frontal cortex signals the tegmentum and accumbens with the conclusions of its expectancy/discrepancy calculations.  The frontal lobe provides executive function, considering bits of information, assessing patterns and then prioritizing the strategies.  The frontal lobe is the most recent part of the brain to evolve and involves a disproportionate percentage of primate-unique genes in its development and operation.  It does not complete development until the mid-20s.  It includes spindle neurons.  It is easily damaged.  Sapolsky (Nauta) notes that its ventromedial prefrontal cortex is a quasi-member of the limbic system. 
.  Maternal behaviors are induced by oxytocin and its production is boosted during birth by the hypothalamus.  And the ventral tegmentum increases its sensitivity to oxytocin by increasing the volume of receptors, in biological cells these proteins are able to span the cell membrane and present an active site which is tailored to interact with a specific signal.  When the receptor pairs with its signal, its overall shape changes resulting in changes in the part internal to the cell which can be relayed by the cells signalling infrastructure.  In neuron synapses one type of receptor (fast) is associated with an ion channel.  The other (slow) is associated with a signalling enzyme chain and modulates the neuron's response. 
.  In rats olfactory stimulation ensures the new mum learns the smell of her offspring. 
The male analog vasopressin developed by duplication and subsequent mutation of the vasotocin gene, along with oxytocin, during the initial formation of mammals.   It acts as a hormone regulating water retention in the kidneys.  It supports paternal behavior stimulated by a female giving birth.  Sex releases vasopressin in the nucleus accumbens of male prairie voles.  And prairie voles have more receptors in the accumbens than other voles supporting their pair bonding.  This situation is similar in bonobos relative to chimps where it encourages social bonding - but not monogamy.  Vasopressin is made in hypothalamic neurons which project to the posterior pituitary for release as a hormone.  It is also a neuropeptide transmitting from hypothalamic projections to the ventral tegmentum, nucleus accumbens, hippocampus, amygdala, and frontal cortex.  And it is made & secreted in other areas of the brain.  Vasopressin enhances aggression in paternal prairie vole males.  But the aggression is then maintained by social learning. 
supported paternal behavior.  Vasopressin receptor levels increase in the male's brain when the female gives birth. 

Oxytocin was also found to support pair bonding is an increase in the strength of relationship between parents and parents and children in some species: prairie voles, bonobos - not monogamous, and humans.  NIMH's Thomas Insel, Emory's Larry Young & Illinois's Sue Carter's research highlighted prairie voles, where pair-bonding is enabled by a genetic difference from montane voles in the operon controlling generation of the vasopressin receptor.  Oxytocin is associated with pair bonding.  There are: Higher levels of receptors in males (vasopressin) having lots of sex and in females (oxytocin) performing grooming & physical contact, Sex releases oxytocin in the nucleus accumbens of female prairie voles.  Such pair-bonded males are less interested in other females. Insel, Young & Carter engineered: (1) Male mice brains to express the prairie vole version of the vasopressin receptor in their brains resulting in grooming and huddling with familiar females.  (2) Male montane vole brains to add vasopressin receptors to the nucleus accumbens resulting in their being more socially affiliative with individual females.  .  And by inhibiting the central amygdala is a relatively ancient evolved part of the amygdala which processes 'innate' fears.  It projects to the BNST to raise the heart rate and blood pressure in preparation for fight and flight. 
it suppresses fear is an emotion which prepares the body for time sensitive action: Blood is sent to the muscles from the gut and skin, Adrenalin is released stimulating: Fuel to be released from the liver, Blood is encouraged to clot, and Face is wide-eyed and fearful.  The short-term high priority goal, experienced as a sense of urgency, is to flee, fight or deflect the danger.  There are both 'innate' - really high priority learning - which are mediated by the central amydala and learned fears which are mediated by the BLA which learns to fear a stimulus and then signals the central amygdala. 
and anxiety is manifested in the amygdala mediating inhibition of dopamine rewards.  Major anxiety results in elevated glucocorticoids and reduces hippocampal dendrite & spine density.  Some estrogen receptor variants are associated with anxiety in women. 
.  Except when a mother has to defend her babies where the central amygdala's instinctual fear supports oxytocin stimulated enhanced aggression. 

Oxytocin supports prosocial capabilities improving detection of happy faces and socially positive words.  It makes people study eyes for longer improving accuracy of reading emotions.  In a social-recognition-task it increases activity in the TPJ is the temporoparietal juncture: left, right; where the temporal and parietal lobes meet, an area of the cortex with projections from the thalamus, limbic system, visual, auditory and somatosensory networks.  It supports modeling of self and others. 
enhancing ideas about what other people are thinking.  But oxytocin decreases cooperation with strangers.  It helps us identify who is an Us.  Abnormalities in the oxytocin vasopressin network are associated with ASD is autism spectrum disorder.  People who suffer from ASD show a reduced fusiform response to faces.  ASD is linked to gene variants affecting oxytocin and vasopressin, to nongenetic mechanisms that silence the oxytocin receptor gene and to lower levels of the receptor itself. 


Oxytocin has contingent effects: It makes charitable people more charitable, Seeking social support occurred more when the subjects were from individual cultures, with a specific receptor variant & under stress is a multi-faceted condition reflecting high cortisol levels.  Dr. Robert Sapolsky's studies of baboons indicate that stress helps build readiness for fight or flight.  As these actions occur the levels of cortisol return to the baseline rate.  A stressor is anything that disrupts the regular homeostatic balance.  The stress response is the array of neural and endocrine changes that occur to respond effectively to the crisis and reestablish homeostasis. 
  • The short term response to the stressor
    • activates the amygdala which: Stimulates the brain stem resulting in inhibition of the parasympathetic nervous system and activation of the sympathetic nervous system with the hormones epinephrine and norepinephrine deployed around the body, Activates the PVN which generates a cascade resulting in glucocorticoid secretion to: get energy to the muscles with increased blood pressure for a powerful response.  The brain's acuity and cognition are stimulated.  The immune system is stimulated with beta-endorphin and repair activities curtail.  But when the stressor is
  • long term: loneliness, debt; and no action is necessary, or possible, long term damage ensues.  Damage from such stress may only occur in specific situations: Nuclear families coping with parents moving in.  Sustained stress provides an evolved amplifier of a position of dominance and status.  It is a strategy in female aggression used to limit reproductive competition.  Sustained stress:
    • Stops the frontal cortex from ensuring we do the harder thing, instead substituting amplification of the individual's propensity for risk-taking and impairing risk assessment! 
    • Activates the integration between the thalamus and amygdala. 
      • Acts differently on the amygdala in comparison to the frontal cortex and hippocampus: Stress strengthens the integration between the Amygdala and the hippocampus, making the hippocampus fearful. 
      • BLA & BNST respond with increased BDNF levels and expanded dendrites persistently increasing anxiety and fear conditioning. 
    • Makes it easier to learn a fear association and to consolidate it into long-term memory.  Sustained stress makes it harder to unlearn fear by making the prefrontal cortex inhibit the BLA from learning to break the fear association and weakening the prefrontal cortex's hold over the amygdala.  And glucocorticoids decrease activation of the medial prefrontal cortex during processing of emotional faces.  Accuracy of assessing emotions from faces suffers.  A terrified rat generating lots of glucocorticoids will cause dendrites in the hippocampus to atrophy but when it generates the same amount from excitement of running on a wheel the dendrites expand.  The activation of the amygdala seems to determine how the hippocampus responds. 
    • Depletes the nucleus accumbens of dopamine biasing rats toward social subordination and biasing humans toward depression. 
    • Disrupts working memory by amplifying norepinephrine signalling in the prefrontal cortex and amygdala to prefrontal cortex signalling until they become destructive.  It also desynchronizes activation in different frontal lobe regions impacting shifting of attention. 
  • During depression, stress inhibits dopamine signalling. 
  • Strategies for stress reduction include: Mindfulness. 
.  The actions of oxytocin & vasopressin depend on context: who you are, your environment, who they are.  Sapolsky notes that is also true for the regulation of the genes for these peptides

The endocrinology of Aggression in females
Sapolsky explains that this subject confuses him:

Maternal Aggression
Typically females increase aggression during pregnancy.  The increase is most significant in species with significant amounts of infanticide

In:

Bare-Knuckled Female Aggression
While non-maternal female competition is typically assumed to be passive and covert, Sapolsky asserts this is mistaken because:

Perimenstrual aggression and irritability
PMS is premenstrual syndrome associated with: negative mood, irritability, bloating cramps, acne; immediately prior to menstruation.  It occurs in various species.  Its symptoms vary by culture.  There is little evidence of an association with aggression. 
and PMDD is premenstrual dysphoric disorder which affects 2 to 5% of women impairing their normal functioning. 
 studies have not identified an association with aggression. 

Stress and imprudent brain function
Sapolsky notes the dilemma that important, consequential behaviors can be stressed situations and stress is a multi-faceted condition reflecting high cortisol levels.  Dr. Robert Sapolsky's studies of baboons indicate that stress helps build readiness for fight or flight.  As these actions occur the levels of cortisol return to the baseline rate.  A stressor is anything that disrupts the regular homeostatic balance.  The stress response is the array of neural and endocrine changes that occur to respond effectively to the crisis and reestablish homeostasis. 
  • The short term response to the stressor
    • activates the amygdala which: Stimulates the brain stem resulting in inhibition of the parasympathetic nervous system and activation of the sympathetic nervous system with the hormones epinephrine and norepinephrine deployed around the body, Activates the PVN which generates a cascade resulting in glucocorticoid secretion to: get energy to the muscles with increased blood pressure for a powerful response.  The brain's acuity and cognition are stimulated.  The immune system is stimulated with beta-endorphin and repair activities curtail.  But when the stressor is
  • long term: loneliness, debt; and no action is necessary, or possible, long term damage ensues.  Damage from such stress may only occur in specific situations: Nuclear families coping with parents moving in.  Sustained stress provides an evolved amplifier of a position of dominance and status.  It is a strategy in female aggression used to limit reproductive competition.  Sustained stress:
    • Stops the frontal cortex from ensuring we do the harder thing, instead substituting amplification of the individual's propensity for risk-taking and impairing risk assessment! 
    • Activates the integration between the thalamus and amygdala. 
      • Acts differently on the amygdala in comparison to the frontal cortex and hippocampus: Stress strengthens the integration between the Amygdala and the hippocampus, making the hippocampus fearful. 
      • BLA & BNST respond with increased BDNF levels and expanded dendrites persistently increasing anxiety and fear conditioning. 
    • Makes it easier to learn a fear association and to consolidate it into long-term memory.  Sustained stress makes it harder to unlearn fear by making the prefrontal cortex inhibit the BLA from learning to break the fear association and weakening the prefrontal cortex's hold over the amygdala.  And glucocorticoids decrease activation of the medial prefrontal cortex during processing of emotional faces.  Accuracy of assessing emotions from faces suffers.  A terrified rat generating lots of glucocorticoids will cause dendrites in the hippocampus to atrophy but when it generates the same amount from excitement of running on a wheel the dendrites expand.  The activation of the amygdala seems to determine how the hippocampus responds. 
    • Depletes the nucleus accumbens of dopamine biasing rats toward social subordination and biasing humans toward depression. 
    • Disrupts working memory by amplifying norepinephrine signalling in the prefrontal cortex and amygdala to prefrontal cortex signalling until they become destructive.  It also desynchronizes activation in different frontal lobe regions impacting shifting of attention. 
  • During depression, stress inhibits dopamine signalling. 
  • Strategies for stress reduction include: Mindfulness. 
often impacts decision making.  Sight of a large carnivore is detected in the amygdala contains > 12 distinct areas: Central, Lateral.  It receives simple signals from the lower parts of the brain: pain from the PAG; and abstract complex information from the highest areas: Disgust from the insula cortex.  It sends signals to almost every other part of the brain, including to the decision-making circuitry of the frontal lobes.  It has high levels of D(1) dopamine receptors.  During extreme fear the amygdala drives the hippocampus into fear learning.  It outputs directly to subcortical reflexive motor pathways when speed is required.  Its central nucleus projects to the BNST.  It signals the locus ceruleus.  The amygdala:
  • Promotes aggression.  Stimulating the amygdala promotes rage.  It converts anger into aggression and when impaired it impacts the ability to detect angry facial expressions.  
  • Participates in disgust
  • Perceives fear promoting stimuli.  In PTSD sufferers the Amygdala overreacts to mildly fearful stimuli and is slow to calm down and the amygdala expands in size over a period of months.  Fear is processed by the lateral nucleus which serves as the input from various senses, and the central nucleus which outputs to the brain stem (central grey - freezing, lateral hypothalamus - blood pressure, activates paraventricular hypothalamus => crf -> hormone adjustments). 
  • Has lots of receptors for and is highly sensitive to glucocorticoids.  Stress inhibits the GABA interneurons in the basolateral amygdala (BLA) allowing the excitatory glutamate releasing neurons to excite more. 
  • Is sensitive to unsettling/uncertain social situations where it promotes anxiety.  It is also interested in uncertain but potentially painful situations.  The amygdala contributes to social and emotional decision making where the BLA supports rejecting an unacceptable offer, as allowed in the Ultimatum Game, by injecting implicit mistrust and vigilance, generating an anger driven rejection that is used as punishment.  The amygdala is very rapidly excited by subliminal signals from the thalamus of outgroup skin color.  The amygdala subsequently tips social emotions against outgroups unless restrained by the frontal lobe or influenced by subliminal priming to prioritize inclusion.  The fast path from the thalamus rapidly but inaccurately signals its identified a weapon. 
  • Promotes male, but not female, sexual motivation when it is an uncertain potential pleasure. 
  • Responds to the longing for uncertain potential pleasures and fear that the reward will not be worth it if it happens.  The amygdala turns off during orgasm. 
  • Uses but is not directly involved in vision. 
which:
Sapolsky notes how we love stress in benevolent, mild, transient amounts.  In these situations the levels of glucocorticoids secreted stimulates production of dopamine is a synaptic signal supporting generalized goal-directed behavior & anticipation of reward.  Its significance is that the receptors that detect the signal are of the slow acting type and are used to alter (modulate) the response of fast acting dopaminergic neural circuits in which the receptors are deployed (LTP).  The signal detects significant changes including predictions of models and actual results which differ unexpectedly.  The dopamine network architecture is designed to signal the possibility of any type of reward: Norm violation punishment, Winning a lottery, & Misfortune of an envied competitor.  Dopamine signalling:
  • Rescales continuously to accommodate the range of intensity offered by different stimuli.  So dopamine's responses to any reward habituate.  GABA is released by some tegmental neurons to induce habituation. 
  • Reflects the anticipation of reward.  It supports establishment of a relationship between a signal, working for a reward and obtaining the reward, but subsequently dopamine is mainly released encouraging the work, right after the signal supporting anticipation of the reward.  Anticipation requires learning and is reflected in hippocampus activity.  That explains context dependent cravings.  And the learning architecture means reliable cues become rewarding.  The accumbens supports willpower.  And dopamine
  • Promotes goal-oriented behavior needed to obtain & likely to achieve the reward - through the dopamine projections to the prefrontal cortex.  That makes dopamine central to:
    • Motivation.  This binding fails in depression - due to stress and in anxiety - due to signals from the amygdala.  
    • The prefrontal cortex's mesocortically stimulated support for willpower to act to delay rewards.  To sustain work for delayed rewards additional dopamine is released based on the length of the delay and the rewards uncertainty (modelled in the dorsolateral prefrontal cortex - which promotes the long term and the ventromedial prefrontal cortex - which promotes the short term) and the anticipated size of the reward (modelled in the accumbens).  Impulsiveness in ADHD is reflected in abnormal dopamine processing.  Addictive drugs bias the dopamine network towards impulsiveness.  
  • Is lowered by certain gene variants which induce: less dopamine in the synapse, fewer receptors, lower responsiveness of receptors; associated with (as tiny effects in hugely varying social scenarios): sensation seeking, risk taking, attentional problems, extroversion; where:
    • The receptor D4's gene shows high variability.  The D47R form is relatively unresponsive to dopamine.  
    • Dopamine is degraded by COMT.  The COMT gene includes a variant which is highly efficient reducing dopamine signalling but with complicating gene/environment interactions.  
    • Dopamine is removed from the synapse by a reuptake transporter DAT. 
based
Read Montague explores how brains make decisions.  In particular he explains how:
  • Evolution can create indirect abstract models, such as the dopamine system, that allow
  • Life changing real-time decisions to be made, and how
  • Schematic structures provide encodings of computable control structures which operate through and on incomputable, schematically encoded, physically active structures and operationally associated production functions. 
rewards


Some important debunking: Alcohol
Sapolsky notes alcohols impact on judgement.  But he writes it does not cause everyone to become aggressive.  Instead people prone to aggression do become aggressive as do people who believe alcohol makes you aggressive.  That second situation demonstrates social learnings power to shape biology. 

Days to Months Before
Any behavior was produced in prior seconds by the nervous system based on sensory cues minutes to hours before with the brain's sensitivity adjusted by hormones are signalling molecules: ACTH, TRH, Melanocyte stimulating hormone, Testosterone, Oxytocin, Vasopressin, Insulin, Growth hormone, Estrogen, Progesterone, Angiotensin II, Asprosin, EPO, Irisin, Leptin, FGF21 hormone, Prostaglandins, TSH, Thyroxine, Glococorticoids; that are transported by the circulatory system to interact with target organs having appropriate receptors.  The levels of hormones can fluctuate massively, as in pregnancy. 
in the previous hours and days.  But in the prior days to months enormous changes occurred is lasting change to the brain that occurs throughout life.  It is also termed neural plasticity.  The changes include:
  • The strength of dendritic input alters due to genetic, neural and hormonal signals
    • Hebb notes that memories require strengthening of preexisting synapses.  Glutamate responsive neurons' post synaptic dendritic spines have two types of receptor: non-NMDA and NMDA.  NMDA channels are responsible for this strengthening mechanism.  LTP then occurs to prolong the increase in excitability of the synapse. 
    • The LTP operation results in calcium diffusion which triggers new spine formation in adjacent parts of the dendrite.  Eventually that can stimulate dentrite growth enabling more neurons to connect. 
    • Short term stress promotes hippocampal LTP.  
    • Sustained stress promotes:
      • Hippocampal & frontal cortex  LTD & suppresses LTP.  Subsequent reductions in NCAM then reduce dendrite and synapse density. 
      • Amygdala LTP and suppresses LTD boosting fear conditioning.  It increases BDNF levels and expands dendrites in the BLA. 
    • Depression and anxiety reduce hippocampal dendrite and spine number by reducing BDNF. 
  • The axon's conditions for
    • Initiating an action potential. 
      • Progesterone boosts GABA-ergic neurons response to GABA decreasing the excitability of other neurons over a period of hours. 
    • Duration of a neuron's refractory period.  Testosterone shortens the refractory period of amygdala and amygdala target neurons over a period of hours. 
  • Synaptic connections being constantly removed and recreated
  • Synapses being created or destroyed.  Stimulation generates additional dendritic spines which become associated with a nearby axon terminal and within weeks a synapse forms.  The synapse then contributes calcium diffusion through LTP triggering more spine formation.  When dendritic spines recede synapses disappear. 
  • Cortical maps change to reflect alterations in the inputs and outputs from the body. 
  • Birth of brain cells in many areas of adult brains: the hippocampus (where 3% are replaced each month) and olfactory bulb and lesser amounts in the cortex. 
  • Restructuring after brain damage including axonal plasticity.  Distant rerouting of axons is observed but no mechanism has been identified yet. 
  • Vision is plastic in predators, where the eyes are moved during final development.  Dehaene argues for neuronal recycling supporting reading.  
in the brain's structure (neuroplasticity) including:

The realization that the brain is able to change throughout its existence introduces reason for optimism in how all of us can behave. 

The mechanisms of memory in the brain includes functionally different types: Declarative (episodic and semantic), Implicit, Procedural, Spatial, Temporal, Verbal; Hebb noted that glutamate receptive neurons learn by (NMDA channel based) synaptic strengthening.  This strengthening is sustained by subsequent LTP.  The non-realtime learning and planning processes operate through consciousness using the working memory structures, and then via sleep, the salient ones are consolidated while the rest are destroyed and garbage collected.   formation and neural plasticity refers to lasting changes to the brain that occur throughout the life span of the organism.  Many aspects of the brain can be altered into adulthood.  Almost anything in the nervous system can change in response to sustained stimulus.  And in a different environment the changes will often reverse.  The changes include:
  • The strength of dendritic input alters due to genetic, neural and hormonal signals
    • Hebb notes that memories require strengthening of preexisting synapses.  Glutamate responsive neurons' post synaptic dendritic spines have two types of receptor: non-NMDA and NMDA.  NMDA channels are responsible for this strengthening mechanism.  LTP then occurs to prolong the increase in excitability of the synapse. 
    • The LTP operation results in calcium diffusion which triggers new spine formation in adjacent parts of the dendrite.  Eventually that can stimulate dentrite growth enabling more neurons to connect. 
    • Short term stress promotes hippocampal LTP.  
    • Sustained stress promotes:
      • Hippocampal & frontal cortex  LTD & suppresses LTP.  Subsequent reductions in NCAM then reduce dendrite and synapse density. 
      • Amygdala LTP and suppresses LTD boosting fear conditioning.  It increases BDNF levels and expands dendrites in the BLA. 
    • Depression and anxiety reduce hippocampal dendrite and spine number by reducing BDNF. 
  • The axon's conditions for
    • Initiating an action potential. 
      • Progesterone boosts GABA-ergic neurons response to GABA decreasing the excitability of other neurons over a period of hours. 
    • Duration of a neuron's refractory period.  Testosterone shortens the refractory period of amygdala and amygdala target neurons over a period of hours. 
  • Synaptic connections being constantly removed and recreated
  • Synapses being created or destroyed.  Stimulation generates additional dendritic spines which become associated with a nearby axon terminal and within weeks a synapse forms.  The synapse then contributes calcium diffusion through LTP triggering more spine formation.  When dendritic spines recede synapses disappear. 
  • Cortical maps change to reflect alterations in the inputs and outputs from the body. 
  • Birth of brain cells in many areas of adult brains: the hippocampus (where 3% are replaced each month) and olfactory bulb and lesser amounts in the cortex. 
  • Restructuring after brain damage including axonal plasticity.  Distant rerouting of axons is observed but no mechanism has been identified yet. 
  • Vision is plastic in predators, where the eyes are moved during final development.  Dehaene argues for neuronal recycling supporting reading.  
are flexible and reversible in different environments:

Adolescence; Or, Dude, Where's my Frontal Cortex?
Adolescence in humans supports the transition from a juvenile configuration, dependent on parents and structured to learn & logistically transform, to adult optimized to the proximate environment.  And it is staged encouraging the adolescents to escape the hierarchy they grew up in and enter other groups where they may bring in: fresh ideas, risk taking; and alter the existing hierarchy: Steve Jobs & Steve Wozniak, Bill Gates & Paul Allen.  It marks the beginning of Piaget's formal operational stage of cognitive development.  The limbic, autonomic and hormone networks are already deployed and functioning effectively.  The frontal cortex has to be pruned: winning neurons move to their final highly connected positions, and are myelinated over time.  The rest dissolve.  So the frontal lobe does not obtain its adult configuration and networked integration until the mid-twenties when prefrontal cortex control becomes optimal.  The evolutionarily oldest areas of the frontal cortex mature first.  The PFC must be iteratively customized by experience to do the right thing as an adult.  Adolescents:
  • Don't detect irony effectively.  They depend on the DMPFC to do this, unlike adults who leverage the fusiform face area.  
  • Regulate emotions with the ventral striatum while the prefrontal cortex is still being setup.  Dopamine projection density and signalling increase from the ventral tegmentum catalyzing increased interest in dopamine based rewards.  Novelty seeking allows for creative exploration which was necessary to move beyond the familial pack.  Criticisms do not get incorporated into learning models by adolescents leaving their risk assessments very poor.  The target of the dopamine networks, the adolescent accumbens, responds to rewards like a gyrating top - hugely to large rewards, and negatively to small rewards.  Eventually as the frontal regions increase in contribution there are steady improvements in: working memory, flexible rule use, executive organization and task shifting.  And adolescents start to see other people's perspective. 
  • Drive the cellular transformations with post-pubescent high levels of testosterone in males, and high but fluctuating estrogen & progesterone levels in females.  Blood flow to the frontal cortex is also diverted on occasion to the groin.  
  • Peer pressure is exceptionally influential in adolescents.  Admired peer comments reduce vmPFC activity and enhance ventral striatal activity.  Adults modulate the mental impact of socially mean treatment: the initial activation of the PAG, anterior cingulate, amygdala, insula cortex; which generate feelings of pain, anger, and disgust, with the VLPFC but that does not occur in adolescents.  
  • Feel empathy intensely, supported by their rampant emotions, interest in novelty, ego.  But feeling the pain of others can induce self-oriented avoidance of the situations. 
is the later part of the human development is a phase during the operation of a CAS agent.  It allows for schematic strategies to be iteratively blended with environmental signals to solve the logistical issues of migrating newly built and transformed sub-agents.  That is needed to achieve the adult configuration of the agent and optimize it for the proximate environment.  Smiley includes examples of the developmental phase agents required in an emergent CAS.  In situations where parents invest in the growth and memetic learning of their offspring the schematic grab bag can support optimizations to develop models, structures and actions to construct an adept adult.  In humans, adolescence leverages neural plasticity, elder sibling advice and adult coaching to help prepare the deploying neuronal network and body to successfully compete. 
phase where it generates different behaviors from young children & adults.  It reflects the slow maturation of the frontal cortex of the cerebral cortex is at the front of the brain.  It includes the: prefrontal cortex, motor cortex.  Sapolsky asserts it makes you do the harder thing when it's the right thing to do.  The frontal cortex supports working memory to sustain focus on a task.  It also coordinates the strategic actions necessary to achieve success.  It provides impulse control, regulation of emotion, and willpower.  The prefrontal cortex maintains focus by deprioritizing currently irrelevant streams of information.  The frontal cortex tracks rules.  Over a lifetime that builds into a costly activity.  Once it tires responses become less prosocial.  But practice shifts operation of tasks to the cerebellum.  The frontal cortex signals the tegmentum and accumbens with the conclusions of its expectancy/discrepancy calculations.  The frontal lobe provides executive function, considering bits of information, assessing patterns and then prioritizing the strategies.  The frontal lobe is the most recent part of the brain to evolve and involves a disproportionate percentage of primate-unique genes in its development and operation.  It does not complete development until the mid-20s.  It includes spindle neurons.  It is easily damaged.  Sapolsky (Nauta) notes that its ventromedial prefrontal cortex is a quasi-member of the limbic system. 
which does not obtain its adult structure and function until the mid-twenties.  Sapolsky stresses the key implications:
  1. Adolescence shapes the frontal cortex
  2. Adolescence is only understood in the context of the delayed frontal cortex's maturation, making it: frustrating, great, asinine, impulsive, inspiring, destructive, self-destructive, selfless, selfish, impossible and world changing.  Sapolsky adds it's when someone is most likely to kill or be killed.  A time of life of maximal risk taking, novelty seeking, and affiliation with peers.  
The delayed development allows the brain to get all its parts optimized from experience to leverage the social toolset: social memory, emotional perspective taking, impulse control, empathy is the capability to relate to another person from their perspective.  It is implemented by spindle neurons.  It is context dependently mediated by estrogen.  It develops over time: Piaget's preoperational stage includes rudimentary empathy, Theory of mind supports the development; initially feeling someone's pain as one integrated being, then for them and eventually as them.  In adults, when someone else is hurt the anterior cingulate cortex, amygdala & insula activates projecting [scapegoating] to the vmPFC.  If the pain is physical the PAG activates and motor neurons for the area where the other person was injured.  The intertwining of the ACC amygdala & insula in adults results in attribution of fault even when there is none which can make it hard to step in and actually help.  But in seven-year-olds the activation is concrete: PAG and sensory & motor cortexes with minimal coupling to the rudimentary vmPFC.  In older children the vmPFC is coupled to limbic structures.  Ten to twelve year olds abstract empathy to classes of people.  By adolescence the vmPFC is coupled to theory of mind regions and intentional harm induces disgust via the amygdala.  Sapolsky explains adolescent boys are utilitarian and tend to accept inequality more than girls do.  But both sexes accept inequality as the way it is.  Sociopaths do not develop empathy. 
, self-regulation, teamwork; appropriately in this particular proximate environment. 

The nuts and bolts of frontal cortical maturation
During adolescence in humans supports the transition from a juvenile configuration, dependent on parents and structured to learn & logistically transform, to adult optimized to the proximate environment.  And it is staged encouraging the adolescents to escape the hierarchy they grew up in and enter other groups where they may bring in: fresh ideas, risk taking; and alter the existing hierarchy: Steve Jobs & Steve Wozniak, Bill Gates & Paul Allen.  It marks the beginning of Piaget's formal operational stage of cognitive development.  The limbic, autonomic and hormone networks are already deployed and functioning effectively.  The frontal cortex has to be pruned: winning neurons move to their final highly connected positions, and are myelinated over time.  The rest dissolve.  So the frontal lobe does not obtain its adult configuration and networked integration until the mid-twenties when prefrontal cortex control becomes optimal.  The evolutionarily oldest areas of the frontal cortex mature first.  The PFC must be iteratively customized by experience to do the right thing as an adult.  Adolescents:
  • Don't detect irony effectively.  They depend on the DMPFC to do this, unlike adults who leverage the fusiform face area.  
  • Regulate emotions with the ventral striatum while the prefrontal cortex is still being setup.  Dopamine projection density and signalling increase from the ventral tegmentum catalyzing increased interest in dopamine based rewards.  Novelty seeking allows for creative exploration which was necessary to move beyond the familial pack.  Criticisms do not get incorporated into learning models by adolescents leaving their risk assessments very poor.  The target of the dopamine networks, the adolescent accumbens, responds to rewards like a gyrating top - hugely to large rewards, and negatively to small rewards.  Eventually as the frontal regions increase in contribution there are steady improvements in: working memory, flexible rule use, executive organization and task shifting.  And adolescents start to see other people's perspective. 
  • Drive the cellular transformations with post-pubescent high levels of testosterone in males, and high but fluctuating estrogen & progesterone levels in females.  Blood flow to the frontal cortex is also diverted on occasion to the groin.  
  • Peer pressure is exceptionally influential in adolescents.  Admired peer comments reduce vmPFC activity and enhance ventral striatal activity.  Adults modulate the mental impact of socially mean treatment: the initial activation of the PAG, anterior cingulate, amygdala, insula cortex; which generate feelings of pain, anger, and disgust, with the VLPFC but that does not occur in adolescents.  
  • Feel empathy intensely, supported by their rampant emotions, interest in novelty, ego.  But feeling the pain of others can induce self-oriented avoidance of the situations. 
the neurons, specialized eukaryotic cells include channels which control flows of sodium and potassium ions across the massively extended cell membrane supporting an electro-chemical wave which is then converted into an outgoing chemical signal transmission from synapses which target nearby neuron or muscle cell receptors.  Neurons are supported by glial cells.  Neurons include a:
  • Receptive element - dendrites
  • Transmitting element - axon and synaptic terminals
  • Highly variable DNA schema using transposons. 
of the frontal cortex of the cerebral cortex is at the front of the brain.  It includes the: prefrontal cortex, motor cortex.  Sapolsky asserts it makes you do the harder thing when it's the right thing to do.  The frontal cortex supports working memory to sustain focus on a task.  It also coordinates the strategic actions necessary to achieve success.  It provides impulse control, regulation of emotion, and willpower.  The prefrontal cortex maintains focus by deprioritizing currently irrelevant streams of information.  The frontal cortex tracks rules.  Over a lifetime that builds into a costly activity.  Once it tires responses become less prosocial.  But practice shifts operation of tasks to the cerebellum.  The frontal cortex signals the tegmentum and accumbens with the conclusions of its expectancy/discrepancy calculations.  The frontal lobe provides executive function, considering bits of information, assessing patterns and then prioritizing the strategies.  The frontal lobe is the most recent part of the brain to evolve and involves a disproportionate percentage of primate-unique genes in its development and operation.  It does not complete development until the mid-20s.  It includes spindle neurons.  It is easily damaged.  Sapolsky (Nauta) notes that its ventromedial prefrontal cortex is a quasi-member of the limbic system. 
that are not used in the proximate environment prune themselves, programmed cell death is a signal initiated DNA controlled process which results in eukaryotic cells self-destructing.   while the rest migrate to their target locations, leaving an optimized network.  The
This page reviews the implications of selection, variation and heredity in a complex adaptive system (CAS).  The mechanism and its emergence are discussed. 
evolutionarily
oldest areas of the frontal cortex mature first leaving the relatively young dl PFC is dorsolateral prefrontal cortex which is:
  • At the heart of decision making - highly rational, unsentimental 
  • A major agent of working memory
  • The most recently evolved part of the prefrontal cortex.  
  • Mainly interconnected with other parts of the cortex.  
changes waiting until late adolescence.  And as the configuration streamlines myelination is the fatty insulating material deployed by Schwann cells & oligodendrocytes, both types of glial cells, around axons to improve their conduction rate.  In humans it is still occurring 25 years after birth.  It has great impact on long axons, in neurons that project over long distances, where it helps brain inter-region signalling.  The long development time of myelination allows for the later myelinated brain regions to be particularly shaped by the proximate environment. 
increases the axon, a long extension of a neuron which has a membrane constructed to support the uni-directional flow of action potential from the dendritic tree and cell body to the synaptic terminals.  's conduction speeds.  While the frontal cortex takes shape other brain regions provide cover: Ventral striatum is a region within the basal ganglia.   It is a target of the tegmentostriatal dopamine pathway.  It has been captured by brain imaging assigning values to subliminal symbols experimentally associated with winning (highly valued) and losing (low valuation) money.  During adolescence, prior to the deployment of the prefrontal cortex, the ventral striatum helps balance/control emotional decision making. 
helps out with emotional regulation. 

The pubescent generation of gonadal hormones: Testosterone is a hormone secreted by the testes in response to stimulation from the hypothalamic/pituitary/testicular cascade, that makes humans more willing to do what it takes to attain and maintain status, according to Sapolsky.  That means players of the Ultimatum Game, if previously given testosterone can become more generous.  Testosterones effect is highly socially contextual so it may encourage acts of kindness or aggression (when challenged).  The level of testosterone does not predict which individuals will be aggressive in: Birds, Fish, Mammals including primates.  Genes impact the potency of testosterone by altering the enzymes that: Construct it, Convert it to estrogen, code the androgen receptor.   This androgen receptor includes a variable polyglutamine repeat which alters the sensitivity to the testosterone signal.  The more potent form is associated with boys showing more dramatic 'masculinization' of the cortex.  But the detected genetic influences are small.  Testosterone decreases activity in the prefrontal cortex and its functional coupling to the amygdala while increasing the coupling between the amygdala & the thalamus.  Testosterone shortens the refactory period of amygdaloid & amygdaloid target neurons.  This results in impulsive risk taking and more focus on unfamiliar faces and distrust of them.  Testosterone increases activity in the ventral tegmentum projecting dopamine to enhance place preference.  Winners of fights become more willing to fight in part due to testosterone increasing confidence and optimism and reducing fear and anxiety.  And winning at: Chess, Athletics, Stock trades; induces the BNST to add testosterone receptors increasing its sensitivity to the hormone.  People become overconfident and overly optimistic. 
, Estrogen is a generic term for a number of related steroid hormones each of which works differently.  Estrogen:
  • Contributes to maternal aggression but it can reduce aggression and enhance empathy, depending on brain state.  There are two different estrogen receptor types which mediate these conflicting effects.  The levels of each type of receptor is independently regulated.  Different receptor variants are associated with:
    • Higher rates of anxiety among women
    • Higher rates of antisocial behavior and conduct disorder in men
, Progesterone is a steroid hormone.  It:
  • Rarely directly effects areas of the brain.  Instead it is converted into other sterioids which have different actions in different brain areas. 
  • Increases maternal aggression in concert with estrogen by increasing oxytocin release in certain brain regions.  However, on its own progresterone decreases aggression and anxiety.  It decreases anxiety by entering neurons where it is converted to allopregnanolone which binds to GABA receptors increasing their sensitivity to GABA. 
; is dramatic and transformational.  Adult female endocrine function is cyclic but at puberty the activity is more intermittent.  Adolescent male brains also receive washes of gonadal hormones and often suffer hypoxia from blood flow diverted to the crotch. 

Frontal Cortical Changes in Cognition in Adolescence
Sapolsky reviews how frontal cortex of the cerebral cortex is at the front of the brain.  It includes the: prefrontal cortex, motor cortex.  Sapolsky asserts it makes you do the harder thing when it's the right thing to do.  The frontal cortex supports working memory to sustain focus on a task.  It also coordinates the strategic actions necessary to achieve success.  It provides impulse control, regulation of emotion, and willpower.  The prefrontal cortex maintains focus by deprioritizing currently irrelevant streams of information.  The frontal cortex tracks rules.  Over a lifetime that builds into a costly activity.  Once it tires responses become less prosocial.  But practice shifts operation of tasks to the cerebellum.  The frontal cortex signals the tegmentum and accumbens with the conclusions of its expectancy/discrepancy calculations.  The frontal lobe provides executive function, considering bits of information, assessing patterns and then prioritizing the strategies.  The frontal lobe is the most recent part of the brain to evolve and involves a disproportionate percentage of primate-unique genes in its development and operation.  It does not complete development until the mid-20s.  It includes spindle neurons.  It is easily damaged.  Sapolsky (Nauta) notes that its ventromedial prefrontal cortex is a quasi-member of the limbic system. 
maturation is reflected in cognition is the ability to orchestrate thought and action in accordance with internal goals according to Princeton's Jonathan Cohen. 
.  During adolescence in humans supports the transition from a juvenile configuration, dependent on parents and structured to learn & logistically transform, to adult optimized to the proximate environment.  And it is staged encouraging the adolescents to escape the hierarchy they grew up in and enter other groups where they may bring in: fresh ideas, risk taking; and alter the existing hierarchy: Steve Jobs & Steve Wozniak, Bill Gates & Paul Allen.  It marks the beginning of Piaget's formal operational stage of cognitive development.  The limbic, autonomic and hormone networks are already deployed and functioning effectively.  The frontal cortex has to be pruned: winning neurons move to their final highly connected positions, and are myelinated over time.  The rest dissolve.  So the frontal lobe does not obtain its adult configuration and networked integration until the mid-twenties when prefrontal cortex control becomes optimal.  The evolutionarily oldest areas of the frontal cortex mature first.  The PFC must be iteratively customized by experience to do the right thing as an adult.  Adolescents:
  • Don't detect irony effectively.  They depend on the DMPFC to do this, unlike adults who leverage the fusiform face area.  
  • Regulate emotions with the ventral striatum while the prefrontal cortex is still being setup.  Dopamine projection density and signalling increase from the ventral tegmentum catalyzing increased interest in dopamine based rewards.  Novelty seeking allows for creative exploration which was necessary to move beyond the familial pack.  Criticisms do not get incorporated into learning models by adolescents leaving their risk assessments very poor.  The target of the dopamine networks, the adolescent accumbens, responds to rewards like a gyrating top - hugely to large rewards, and negatively to small rewards.  Eventually as the frontal regions increase in contribution there are steady improvements in: working memory, flexible rule use, executive organization and task shifting.  And adolescents start to see other people's perspective. 
  • Drive the cellular transformations with post-pubescent high levels of testosterone in males, and high but fluctuating estrogen & progesterone levels in females.  Blood flow to the frontal cortex is also diverted on occasion to the groin.  
  • Peer pressure is exceptionally influential in adolescents.  Admired peer comments reduce vmPFC activity and enhance ventral striatal activity.  Adults modulate the mental impact of socially mean treatment: the initial activation of the PAG, anterior cingulate, amygdala, insula cortex; which generate feelings of pain, anger, and disgust, with the VLPFC but that does not occur in adolescents.  
  • Feel empathy intensely, supported by their rampant emotions, interest in novelty, ego.  But feeling the pain of others can induce self-oriented avoidance of the situations. 
improvements are seen in:

Frontal Cortical Changes in Emotional Regulation
Adolescents in humans supports the transition from a juvenile configuration, dependent on parents and structured to learn & logistically transform, to adult optimized to the proximate environment.  And it is staged encouraging the adolescents to escape the hierarchy they grew up in and enter other groups where they may bring in: fresh ideas, risk taking; and alter the existing hierarchy: Steve Jobs & Steve Wozniak, Bill Gates & Paul Allen.  It marks the beginning of Piaget's formal operational stage of cognitive development.  The limbic, autonomic and hormone networks are already deployed and functioning effectively.  The frontal cortex has to be pruned: winning neurons move to their final highly connected positions, and are myelinated over time.  The rest dissolve.  So the frontal lobe does not obtain its adult configuration and networked integration until the mid-twenties when prefrontal cortex control becomes optimal.  The evolutionarily oldest areas of the frontal cortex mature first.  The PFC must be iteratively customized by experience to do the right thing as an adult.  Adolescents:
  • Don't detect irony effectively.  They depend on the DMPFC to do this, unlike adults who leverage the fusiform face area.  
  • Regulate emotions with the ventral striatum while the prefrontal cortex is still being setup.  Dopamine projection density and signalling increase from the ventral tegmentum catalyzing increased interest in dopamine based rewards.  Novelty seeking allows for creative exploration which was necessary to move beyond the familial pack.  Criticisms do not get incorporated into learning models by adolescents leaving their risk assessments very poor.  The target of the dopamine networks, the adolescent accumbens, responds to rewards like a gyrating top - hugely to large rewards, and negatively to small rewards.  Eventually as the frontal regions increase in contribution there are steady improvements in: working memory, flexible rule use, executive organization and task shifting.  And adolescents start to see other people's perspective. 
  • Drive the cellular transformations with post-pubescent high levels of testosterone in males, and high but fluctuating estrogen & progesterone levels in females.  Blood flow to the frontal cortex is also diverted on occasion to the groin.  
  • Peer pressure is exceptionally influential in adolescents.  Admired peer comments reduce vmPFC activity and enhance ventral striatal activity.  Adults modulate the mental impact of socially mean treatment: the initial activation of the PAG, anterior cingulate, amygdala, insula cortex; which generate feelings of pain, anger, and disgust, with the VLPFC but that does not occur in adolescents.  
  • Feel empathy intensely, supported by their rampant emotions, interest in novelty, ego.  But feeling the pain of others can induce self-oriented avoidance of the situations. 
experience emotions are low level agents distributed across the brain and body which associate, via the amygdala and rich club hubs, important environmental signals with encoded high speed sensors, and distributed programs of action to model: predict, prioritize guidance signals, select and respond effectively, coherently and rapidly to the initial signal.  The majority of emotion centered brain regions interface to the midbrain through the hypothalamus.  The most accessible signs of emotions are the hard to control and universal facial expressions.  Emotions provide prioritization for conscious access given that an animal has only one body, but possibly many cells, with which to achieve its highest level goals.  Because of this emotions clash with group goals and are disparaged by the powerful.  Evolutionary psychology argues evolution shaped human emotions during the long period of hunter-gatherer existence in the African savanna.  Human emotions are universal and include: Anger, Appreciation of natural beauty, Disgust, Fear, Gratitude, Grief, Guilt, Happiness, Honor, Jealousy, Liking, Love, Rage, Romantic love, Lust for revenge, Passion, Sadness, Self-control, Shame, Sympathy, Surprise; and the sham emotions and distrust induced by reciprocal altruism.   more intensely than adults or young children.  They respond more to affective visual displays is a face expressing strong emotions as interpreted by an observer's brain.  In:
  • Adult observers this activates that amygdala and subsequently the vmPFC as they become habituated to the emotional content. 
  • Adolescents the vmPFC responds less allowing the amygdala's activation to keep growing. 
.  Coping using reappraisal gets more effective during adolescence as the ventral striatum is a region within the basal ganglia.   It is a target of the tegmentostriatal dopamine pathway.  It has been captured by brain imaging assigning values to subliminal symbols experimentally associated with winning (highly valued) and losing (low valuation) money.  During adolescence, prior to the deployment of the prefrontal cortex, the ventral striatum helps balance/control emotional decision making. 
hands over the dampening activity to the frontal cortex of the cerebral cortex is at the front of the brain.  It includes the: prefrontal cortex, motor cortex.  Sapolsky asserts it makes you do the harder thing when it's the right thing to do.  The frontal cortex supports working memory to sustain focus on a task.  It also coordinates the strategic actions necessary to achieve success.  It provides impulse control, regulation of emotion, and willpower.  The prefrontal cortex maintains focus by deprioritizing currently irrelevant streams of information.  The frontal cortex tracks rules.  Over a lifetime that builds into a costly activity.  Once it tires responses become less prosocial.  But practice shifts operation of tasks to the cerebellum.  The frontal cortex signals the tegmentum and accumbens with the conclusions of its expectancy/discrepancy calculations.  The frontal lobe provides executive function, considering bits of information, assessing patterns and then prioritizing the strategies.  The frontal lobe is the most recent part of the brain to evolve and involves a disproportionate percentage of primate-unique genes in its development and operation.  It does not complete development until the mid-20s.  It includes spindle neurons.  It is easily damaged.  Sapolsky (Nauta) notes that its ventromedial prefrontal cortex is a quasi-member of the limbic system. 
.  While the striatum is involved dopamine is a synaptic signal supporting generalized goal-directed behavior & anticipation of reward.  Its significance is that the receptors that detect the signal are of the slow acting type and are used to alter (modulate) the response of fast acting dopaminergic neural circuits in which the receptors are deployed (LTP).  The signal detects significant changes including predictions of models and actual results which differ unexpectedly.  The dopamine network architecture is designed to signal the possibility of any type of reward: Norm violation punishment, Winning a lottery, & Misfortune of an envied competitor.  Dopamine signalling:
  • Rescales continuously to accommodate the range of intensity offered by different stimuli.  So dopamine's responses to any reward habituate.  GABA is released by some tegmental neurons to induce habituation. 
  • Reflects the anticipation of reward.  It supports establishment of a relationship between a signal, working for a reward and obtaining the reward, but subsequently dopamine is mainly released encouraging the work, right after the signal supporting anticipation of the reward.  Anticipation requires learning and is reflected in hippocampus activity.  That explains context dependent cravings.  And the learning architecture means reliable cues become rewarding.  The accumbens supports willpower.  And dopamine
  • Promotes goal-oriented behavior needed to obtain & likely to achieve the reward - through the dopamine projections to the prefrontal cortex.  That makes dopamine central to:
    • Motivation.  This binding fails in depression - due to stress and in anxiety - due to signals from the amygdala.  
    • The prefrontal cortex's mesocortically stimulated support for willpower to act to delay rewards.  To sustain work for delayed rewards additional dopamine is released based on the length of the delay and the rewards uncertainty (modelled in the dorsolateral prefrontal cortex - which promotes the long term and the ventromedial prefrontal cortex - which promotes the short term) and the anticipated size of the reward (modelled in the accumbens).  Impulsiveness in ADHD is reflected in abnormal dopamine processing.  Addictive drugs bias the dopamine network towards impulsiveness.  
  • Is lowered by certain gene variants which induce: less dopamine in the synapse, fewer receptors, lower responsiveness of receptors; associated with (as tiny effects in hugely varying social scenarios): sensation seeking, risk taking, attentional problems, extroversion; where:
    • The receptor D4's gene shows high variability.  The D47R form is relatively unresponsive to dopamine.  
    • Dopamine is degraded by COMT.  The COMT gene includes a variant which is highly efficient reducing dopamine signalling but with complicating gene/environment interactions.  
    • Dopamine is removed from the synapse by a reuptake transporter DAT. 
Read Montague explores how brains make decisions.  In particular he explains how:
  • Evolution can create indirect abstract models, such as the dopamine system, that allow
  • Life changing real-time decisions to be made, and how
  • Schematic structures provide encodings of computable control structures which operate through and on incomputable, schematically encoded, physically active structures and operationally associated production functions. 
based rewards
are very significant encouraging risk taking

Adolescent Risk Taking
Adolescents in humans supports the transition from a juvenile configuration, dependent on parents and structured to learn & logistically transform, to adult optimized to the proximate environment.  And it is staged encouraging the adolescents to escape the hierarchy they grew up in and enter other groups where they may bring in: fresh ideas, risk taking; and alter the existing hierarchy: Steve Jobs & Steve Wozniak, Bill Gates & Paul Allen.  It marks the beginning of Piaget's formal operational stage of cognitive development.  The limbic, autonomic and hormone networks are already deployed and functioning effectively.  The frontal cortex has to be pruned: winning neurons move to their final highly connected positions, and are myelinated over time.  The rest dissolve.  So the frontal lobe does not obtain its adult configuration and networked integration until the mid-twenties when prefrontal cortex control becomes optimal.  The evolutionarily oldest areas of the frontal cortex mature first.  The PFC must be iteratively customized by experience to do the right thing as an adult.  Adolescents:
  • Don't detect irony effectively.  They depend on the DMPFC to do this, unlike adults who leverage the fusiform face area.  
  • Regulate emotions with the ventral striatum while the prefrontal cortex is still being setup.  Dopamine projection density and signalling increase from the ventral tegmentum catalyzing increased interest in dopamine based rewards.  Novelty seeking allows for creative exploration which was necessary to move beyond the familial pack.  Criticisms do not get incorporated into learning models by adolescents leaving their risk assessments very poor.  The target of the dopamine networks, the adolescent accumbens, responds to rewards like a gyrating top - hugely to large rewards, and negatively to small rewards.  Eventually as the frontal regions increase in contribution there are steady improvements in: working memory, flexible rule use, executive organization and task shifting.  And adolescents start to see other people's perspective. 
  • Drive the cellular transformations with post-pubescent high levels of testosterone in males, and high but fluctuating estrogen & progesterone levels in females.  Blood flow to the frontal cortex is also diverted on occasion to the groin.  
  • Peer pressure is exceptionally influential in adolescents.  Admired peer comments reduce vmPFC activity and enhance ventral striatal activity.  Adults modulate the mental impact of socially mean treatment: the initial activation of the PAG, anterior cingulate, amygdala, insula cortex; which generate feelings of pain, anger, and disgust, with the VLPFC but that does not occur in adolescents.  
  • Feel empathy intensely, supported by their rampant emotions, interest in novelty, ego.  But feeling the pain of others can induce self-oriented avoidance of the situations. 
activate the prefrontal cortex (PFC) is
  • The front part of the frontal lobe of the cerebral cortex.  It evolved most recently.  During adolescence when the PFC is still deploying, older brain agents provide equivalent strategies: ventral striatum.  The PFC has been implicated in planning, working memory: dorsolateral; decision making: Orbitofrontal cortex; and social behavior.  Different PFC circuits track internal reward driven strategies and externally signalled advice.  The PFC chooses between conflicting options, letting go or restraint, especially between cognition and emotions.  It imposes an overarching strategy for managing working memory.  It is essential for thinking about multiple items with different labels.  It includes neurons that are interested in particular sub-categories: Dog, Cat.  Once it has made a decision it signals the rest of the frontal lobe just behind it.  Glucocorticoids decrease excitability of the PFC.  
less often than adults when making risky, is an assessment of the likelihood of an independent problem occurring.  It can be assigned an accurate probability since it is independent of other variables in the system.  As such it is different from uncertainty. 
decisions.  They do not incorporate feedback based on their poor assessments about bad news!  So they are willing to take more risks.  They also seek more novelty, developing new and subsequently stable tastes in music, food, and fashion.  In most primates adolescents of one sex make their way into other groups driven by this desire for novelty.  Sapolsky suggests the novelty seeking reflects the increasing development of the mesolimbic projection includes projections between the tegmentum and the accumbens, amygdala, hippocampus;
to the nucleus accumbens is a region of the basal forebrain rostral to the preoptic area and immediately adjacent to the septum.  The nucleus accumbens was closely associated with the limbic system and plays an important role in reinforcement.  If a rodent wins a fight on his home territory, there are long-lasting increases in levels of testosterone receptors enhancing pleasurable effects.  When prairie voles first mate, epi-genetic state changes are induced in the accumbens to support pair-bonding.  The accumbens projects to brain regions associated with movement.  The major pathways of dopaminergic neurons begin in the substantia nigra and the ventral tegmental area.  The amygdala projects back to the accumbens.  The tegmentostriatal system begins in the ventral tegmental area and projects to the nucleus accumbens.  The nucleus accumbens includes high levels of D1, D2 and D3 dopamine receptors located on the spine & shafts of dendrites of excitatory cells reduce the transfer of excitation from the dendrites to the cell bodies, so only especially strong excitatory inputs get through to the cell body to elicit excitation.  It also has D4 dopamine receptors which are highly variable.  The accumbens responds differently to rewards depending on maturity: In juveniles all reward levels result in the same response activity level, During adolescence the accumbens responds to small rewards negatively, and large rewards hugely, In adults the rewards result in measured scaled positive responses.  Chronic stress depletes dopamine from the nucleus accumbens biasing humans towards depression.   and the mesocortical projection includes projections between the tegmentum and the prefrontal cortex;
to the frontal cortex of the cerebral cortex is at the front of the brain.  It includes the: prefrontal cortex, motor cortex.  Sapolsky asserts it makes you do the harder thing when it's the right thing to do.  The frontal cortex supports working memory to sustain focus on a task.  It also coordinates the strategic actions necessary to achieve success.  It provides impulse control, regulation of emotion, and willpower.  The prefrontal cortex maintains focus by deprioritizing currently irrelevant streams of information.  The frontal cortex tracks rules.  Over a lifetime that builds into a costly activity.  Once it tires responses become less prosocial.  But practice shifts operation of tasks to the cerebellum.  The frontal cortex signals the tegmentum and accumbens with the conclusions of its expectancy/discrepancy calculations.  The frontal lobe provides executive function, considering bits of information, assessing patterns and then prioritizing the strategies.  The frontal lobe is the most recent part of the brain to evolve and involves a disproportionate percentage of primate-unique genes in its development and operation.  It does not complete development until the mid-20s.  It includes spindle neurons.  It is easily damaged.  Sapolsky (Nauta) notes that its ventromedial prefrontal cortex is a quasi-member of the limbic system. 
.  Novelty seeking peaks as the
This page discusses the mechanisms and effects of emergence underpinning any complex adaptive system (CAS).  Key research is reviewed. 
emerging
frontal regulation is prefrontal cortex which is:
  • The front part of the frontal lobe of the cerebral cortex.  It evolved most recently.  During adolescence when the PFC is still deploying, older brain agents provide equivalent strategies: ventral striatum.  The PFC has been implicated in planning, working memory: dorsolateral; decision making: Orbitofrontal cortex; and social behavior.  Different PFC circuits track internal reward driven strategies and externally signalled advice.  The PFC chooses between conflicting options, letting go or restraint, especially between cognition and emotions.  It imposes an overarching strategy for managing working memory.  It is essential for thinking about multiple items with different labels.  It includes neurons that are interested in particular sub-categories: Dog, Cat.  Once it has made a decision it signals the rest of the frontal lobe just behind it.  Glucocorticoids decrease excitability of the PFC.  
starts to inhibit activity. 

Peers, social acceptance, and social exclusion
Adolescents in humans supports the transition from a juvenile configuration, dependent on parents and structured to learn & logistically transform, to adult optimized to the proximate environment.  And it is staged encouraging the adolescents to escape the hierarchy they grew up in and enter other groups where they may bring in: fresh ideas, risk taking; and alter the existing hierarchy: Steve Jobs & Steve Wozniak, Bill Gates & Paul Allen.  It marks the beginning of Piaget's formal operational stage of cognitive development.  The limbic, autonomic and hormone networks are already deployed and functioning effectively.  The frontal cortex has to be pruned: winning neurons move to their final highly connected positions, and are myelinated over time.  The rest dissolve.  So the frontal lobe does not obtain its adult configuration and networked integration until the mid-twenties when prefrontal cortex control becomes optimal.  The evolutionarily oldest areas of the frontal cortex mature first.  The PFC must be iteratively customized by experience to do the right thing as an adult.  Adolescents:
  • Don't detect irony effectively.  They depend on the DMPFC to do this, unlike adults who leverage the fusiform face area.  
  • Regulate emotions with the ventral striatum while the prefrontal cortex is still being setup.  Dopamine projection density and signalling increase from the ventral tegmentum catalyzing increased interest in dopamine based rewards.  Novelty seeking allows for creative exploration which was necessary to move beyond the familial pack.  Criticisms do not get incorporated into learning models by adolescents leaving their risk assessments very poor.  The target of the dopamine networks, the adolescent accumbens, responds to rewards like a gyrating top - hugely to large rewards, and negatively to small rewards.  Eventually as the frontal regions increase in contribution there are steady improvements in: working memory, flexible rule use, executive organization and task shifting.  And adolescents start to see other people's perspective. 
  • Drive the cellular transformations with post-pubescent high levels of testosterone in males, and high but fluctuating estrogen & progesterone levels in females.  Blood flow to the frontal cortex is also diverted on occasion to the groin.  
  • Peer pressure is exceptionally influential in adolescents.  Admired peer comments reduce vmPFC activity and enhance ventral striatal activity.  Adults modulate the mental impact of socially mean treatment: the initial activation of the PAG, anterior cingulate, amygdala, insula cortex; which generate feelings of pain, anger, and disgust, with the VLPFC but that does not occur in adolescents.  
  • Feel empathy intensely, supported by their rampant emotions, interest in novelty, ego.  But feeling the pain of others can induce self-oriented avoidance of the situations. 
are particularly vulnerable to peer pressure from friends.  They frantically need to belong.  Peers' signals, is an emergent capability which is used by cooperating agents to support coordination & rival agents to support control and dominance.  In eukaryotic cells signalling is used extensively.  A signal interacts with the exposed region of a receptor molecule inducing it to change shape to an activated form.  Chains of enzymes interact with the activated receptor relaying, amplifying and responding to the signal to change the state of the cell.  Many of the signalling pathways pass through the nuclear membrane and interact with the DNA to change its state.  Enzymes sensitive to the changes induced in the DNA then start to operate generating actions including sending further signals.  Cell signalling is reviewed by Helmreich.  Signalling is a fundamental aspect of CAS theory and is discussed from the abstract CAS perspective in signals and sensors.  In AWF the eukaryotic signalling architecture has been abstracted in a codelet based implementation.  To be credible signals must be hard to fake.  To be effective they must be easily detected by the target recipient.  To be efficient they are low cost to produce and destroy. 
lessen vmPFC is ventromedial prefrontal cortex which is:
  • Focused on the impact of emotion on decision making
  • A participant in limbic system operations 
  • Many human behaviors involve interactions between the vmPFC, the limbic system & the dorsolateral prefrontal cortex.  Part of decision making is for the limbic system to internally simulate (often with the help of the sympathetic nervous system) what alternative outcomes of a decision will feel like with the results of these somatic marker analyses being reported to the vmPFC.  
  • Damage to the vmPFC results in bad decision making: Poor judgement in choosing friends & partners, Failure to respond to negative feedback; because they can't feel the issues. 
activity & enhance ventral striatal is a region within the basal ganglia.   It is a target of the tegmentostriatal dopamine pathway.  It has been captured by brain imaging assigning values to subliminal symbols experimentally associated with winning (highly valued) and losing (low valuation) money.  During adolescence, prior to the deployment of the prefrontal cortex, the ventral striatum helps balance/control emotional decision making. 
activity in adolescents but not adults.  Rejection hurts them more: Periaqueductal gray, ACC is either the
  • Anterior cingulate cortex which:
    • Is a central focus of empathy supporting people relating to other's pain.  This is dependent on oxytocin. 
    • In non-human mammals it processes interoceptive signals.  The ACC focuses the internal signals into high level 'gut intuitions.'  Pain catches the ACC's attention. 
    • Performs discrepancy detection from the outcome that was predicted - at a high level.  The ACC cares about the meaning of what is predicted. 
      • If the ACC has been convinced that a pain killer placebo has inhibited pain signals, the ACC will stay silent about actual pain that is signalled from interoceptive networks.  
      • The ACC will signal: physical pain, emotional pain, metaphorical pain, anxiety, disgust, embarrassment, social exclusion especially in adolescence; as one and the same.  The ACC's abnormalities being associated with major depression. 
    • Has a bridging role between the empathetic and self-interested pain monitor.  Sapolsky notes the ACC is essential for learning fear and conditioned avoidance by observation alone through an intermediate step of shared representation of self.  He concludes "At its core the ACC is about self-interest, with caring about the other person in pain as an add-on." 
  • American College of Cardiology
, Amygdala contains > 12 distinct areas: Central, Lateral.  It receives simple signals from the lower parts of the brain: pain from the PAG; and abstract complex information from the highest areas: Disgust from the insula cortex.  It sends signals to almost every other part of the brain, including to the decision-making circuitry of the frontal lobes.  It has high levels of D(1) dopamine receptors.  During extreme fear the amygdala drives the hippocampus into fear learning.  It outputs directly to subcortical reflexive motor pathways when speed is required.  Its central nucleus projects to the BNST.  It signals the locus ceruleus.  The amygdala:
  • Promotes aggression.  Stimulating the amygdala promotes rage.  It converts anger into aggression and when impaired it impacts the ability to detect angry facial expressions.  
  • Participates in disgust
  • Perceives fear promoting stimuli.  In PTSD sufferers the Amygdala overreacts to mildly fearful stimuli and is slow to calm down and the amygdala expands in size over a period of months.  Fear is processed by the lateral nucleus which serves as the input from various senses, and the central nucleus which outputs to the brain stem (central grey - freezing, lateral hypothalamus - blood pressure, activates paraventricular hypothalamus => crf -> hormone adjustments). 
  • Has lots of receptors for and is highly sensitive to glucocorticoids.  Stress inhibits the GABA interneurons in the basolateral amygdala (BLA) allowing the excitatory glutamate releasing neurons to excite more. 
  • Is sensitive to unsettling/uncertain social situations where it promotes anxiety.  It is also interested in uncertain but potentially painful situations.  The amygdala contributes to social and emotional decision making where the BLA supports rejecting an unacceptable offer, as allowed in the Ultimatum Game, by injecting implicit mistrust and vigilance, generating an anger driven rejection that is used as punishment.  The amygdala is very rapidly excited by subliminal signals from the thalamus of outgroup skin color.  The amygdala subsequently tips social emotions against outgroups unless restrained by the frontal lobe or influenced by subliminal priming to prioritize inclusion.  The fast path from the thalamus rapidly but inaccurately signals its identified a weapon. 
  • Promotes male, but not female, sexual motivation when it is an uncertain potential pleasure. 
  • Responds to the longing for uncertain potential pleasures and fear that the reward will not be worth it if it happens.  The amygdala turns off during orgasm. 
  • Uses but is not directly involved in vision. 
, Insula is part of the cerebral cortex folded deep within the lateral sulcus.  It includes: anterior, posterior insula; and is overlaid by the operculum.  It is assumed to participate in consciousness where it has been linked to emotion, salience & body homeostasis functions:
  • Perception,
  • Motor control: Hand-&-eye motor movement, Swallowing, Gastric motility, Speech articulation;
  • Self-awareness,
  • Inter-personal experiences: Disgust at smells, contamination & mutilation which generate visceral responses, that are projected to the amygdala; binding physical and moral aspects of purity (Macbeth effect)
  • Homeostatic regulation of the sympathetic network, parasympathetic network, and immune system.  
; activation with no calming signals from the vlPFC is ventrolateral prefrontal cortex


Empathy, sympathy, and moral reasoning
Adolescents in humans supports the transition from a juvenile configuration, dependent on parents and structured to learn & logistically transform, to adult optimized to the proximate environment.  And it is staged encouraging the adolescents to escape the hierarchy they grew up in and enter other groups where they may bring in: fresh ideas, risk taking; and alter the existing hierarchy: Steve Jobs & Steve Wozniak, Bill Gates & Paul Allen.  It marks the beginning of Piaget's formal operational stage of cognitive development.  The limbic, autonomic and hormone networks are already deployed and functioning effectively.  The frontal cortex has to be pruned: winning neurons move to their final highly connected positions, and are myelinated over time.  The rest dissolve.  So the frontal lobe does not obtain its adult configuration and networked integration until the mid-twenties when prefrontal cortex control becomes optimal.  The evolutionarily oldest areas of the frontal cortex mature first.  The PFC must be iteratively customized by experience to do the right thing as an adult.  Adolescents:
  • Don't detect irony effectively.  They depend on the DMPFC to do this, unlike adults who leverage the fusiform face area.  
  • Regulate emotions with the ventral striatum while the prefrontal cortex is still being setup.  Dopamine projection density and signalling increase from the ventral tegmentum catalyzing increased interest in dopamine based rewards.  Novelty seeking allows for creative exploration which was necessary to move beyond the familial pack.  Criticisms do not get incorporated into learning models by adolescents leaving their risk assessments very poor.  The target of the dopamine networks, the adolescent accumbens, responds to rewards like a gyrating top - hugely to large rewards, and negatively to small rewards.  Eventually as the frontal regions increase in contribution there are steady improvements in: working memory, flexible rule use, executive organization and task shifting.  And adolescents start to see other people's perspective. 
  • Drive the cellular transformations with post-pubescent high levels of testosterone in males, and high but fluctuating estrogen & progesterone levels in females.  Blood flow to the frontal cortex is also diverted on occasion to the groin.  
  • Peer pressure is exceptionally influential in adolescents.  Admired peer comments reduce vmPFC activity and enhance ventral striatal activity.  Adults modulate the mental impact of socially mean treatment: the initial activation of the PAG, anterior cingulate, amygdala, insula cortex; which generate feelings of pain, anger, and disgust, with the VLPFC but that does not occur in adolescents.  
  • Feel empathy intensely, supported by their rampant emotions, interest in novelty, ego.  But feeling the pain of others can induce self-oriented avoidance of the situations. 
are better at seeing how they would feel in someone else's situation than seeing how the other person might feel.  Similarly their moral judgements provides rules for identifying right from wrong.  It develops in stages with children using play to work out rules of appropriate behavior.  Kohlberg's 1950s experiments using children led him to conclude moral judgement is a cognitive process that develops in three stages.  Sapolsky raises issues with the framework: Its a model, It does not apply to other cultures, Intuition & emotion are as significant as cognition, Moral reasoning doesn't predict moral actions; and notes the capacity of the frontal cortex to regulate emotions and behavior is far more predictive.  The marshmallow test, performed on three to six year olds, actually predicted their subsequent SAT scores at high school, social success and lack of aggression, and forty years on more PFC activation during a frontal task and a lower BMI!  Jonathan Haidt argues people's moral decisions are rationalizations rather than using reasoning. 
are not as sophisticated as adults since they fail to account for systemic effects. 

As they get older intentional harm to others becomes of interest to the dlPFC is dorsolateral prefrontal cortex which is:
  • At the heart of decision making - highly rational, unsentimental 
  • A major agent of working memory
  • The most recently evolved part of the prefrontal cortex.  
  • Mainly interconnected with other parts of the cortex.  
& vmPFC is ventromedial prefrontal cortex which is:
  • Focused on the impact of emotion on decision making
  • A participant in limbic system operations 
  • Many human behaviors involve interactions between the vmPFC, the limbic system & the dorsolateral prefrontal cortex.  Part of decision making is for the limbic system to internally simulate (often with the help of the sympathetic nervous system) what alternative outcomes of a decision will feel like with the results of these somatic marker analyses being reported to the vmPFC.  
  • Damage to the vmPFC results in bad decision making: Poor judgement in choosing friends & partners, Failure to respond to negative feedback; because they can't feel the issues. 
rather than just the amygdala contains > 12 distinct areas: Central, Lateral.  It receives simple signals from the lower parts of the brain: pain from the PAG; and abstract complex information from the highest areas: Disgust from the insula cortex.  It sends signals to almost every other part of the brain, including to the decision-making circuitry of the frontal lobes.  It has high levels of D(1) dopamine receptors.  During extreme fear the amygdala drives the hippocampus into fear learning.  It outputs directly to subcortical reflexive motor pathways when speed is required.  Its central nucleus projects to the BNST.  It signals the locus ceruleus.  The amygdala:
  • Promotes aggression.  Stimulating the amygdala promotes rage.  It converts anger into aggression and when impaired it impacts the ability to detect angry facial expressions.  
  • Participates in disgust
  • Perceives fear promoting stimuli.  In PTSD sufferers the Amygdala overreacts to mildly fearful stimuli and is slow to calm down and the amygdala expands in size over a period of months.  Fear is processed by the lateral nucleus which serves as the input from various senses, and the central nucleus which outputs to the brain stem (central grey - freezing, lateral hypothalamus - blood pressure, activates paraventricular hypothalamus => crf -> hormone adjustments). 
  • Has lots of receptors for and is highly sensitive to glucocorticoids.  Stress inhibits the GABA interneurons in the basolateral amygdala (BLA) allowing the excitatory glutamate releasing neurons to excite more. 
  • Is sensitive to unsettling/uncertain social situations where it promotes anxiety.  It is also interested in uncertain but potentially painful situations.  The amygdala contributes to social and emotional decision making where the BLA supports rejecting an unacceptable offer, as allowed in the Ultimatum Game, by injecting implicit mistrust and vigilance, generating an anger driven rejection that is used as punishment.  The amygdala is very rapidly excited by subliminal signals from the thalamus of outgroup skin color.  The amygdala subsequently tips social emotions against outgroups unless restrained by the frontal lobe or influenced by subliminal priming to prioritize inclusion.  The fast path from the thalamus rapidly but inaccurately signals its identified a weapon. 
  • Promotes male, but not female, sexual motivation when it is an uncertain potential pleasure. 
  • Responds to the longing for uncertain potential pleasures and fear that the reward will not be worth it if it happens.  The amygdala turns off during orgasm. 
  • Uses but is not directly involved in vision. 
and insula is part of the cerebral cortex folded deep within the lateral sulcus.  It includes: anterior, posterior insula; and is overlaid by the operculum.  It is assumed to participate in consciousness where it has been linked to emotion, salience & body homeostasis functions:
  • Perception,
  • Motor control: Hand-&-eye motor movement, Swallowing, Gastric motility, Speech articulation;
  • Self-awareness,
  • Inter-personal experiences: Disgust at smells, contamination & mutilation which generate visceral responses, that are projected to the amygdala; binding physical and moral aspects of purity (Macbeth effect)
  • Homeostatic regulation of the sympathetic network, parasympathetic network, and immune system.  


Sapolsky's core interest in the best & worst behaviors is rewarded by adolescents ability to feel other's pain empathetically is the capability to relate to another person from their perspective.  It is implemented by spindle neurons.  It is context dependently mediated by estrogen.  It develops over time: Piaget's preoperational stage includes rudimentary empathy, Theory of mind supports the development; initially feeling someone's pain as one integrated being, then for them and eventually as them.  In adults, when someone else is hurt the anterior cingulate cortex, amygdala & insula activates projecting [scapegoating] to the vmPFC.  If the pain is physical the PAG activates and motor neurons for the area where the other person was injured.  The intertwining of the ACC amygdala & insula in adults results in attribution of fault even when there is none which can make it hard to step in and actually help.  But in seven-year-olds the activation is concrete: PAG and sensory & motor cortexes with minimal coupling to the rudimentary vmPFC.  In older children the vmPFC is coupled to limbic structures.  Ten to twelve year olds abstract empathy to classes of people.  By adolescence the vmPFC is coupled to theory of mind regions and intentional harm induces disgust via the amygdala.  Sapolsky explains adolescent boys are utilitarian and tend to accept inequality more than girls do.  But both sexes accept inequality as the way it is.  Sociopaths do not develop empathy. 
and try with boundless energy and confidence to make 'things' right. 

Except that doing the right thing in an intense empathetic or moral situation is about prior practice:
  • Allowing the brain's signals to act unconsciously, thus
  • Avoiding overwhelming emotional pain becoming one's total conscious inhibiting focus. 

Adolescent violence
Sapolsky notes violence peaks in late adolescence in humans supports the transition from a juvenile configuration, dependent on parents and structured to learn & logistically transform, to adult optimized to the proximate environment.  And it is staged encouraging the adolescents to escape the hierarchy they grew up in and enter other groups where they may bring in: fresh ideas, risk taking; and alter the existing hierarchy: Steve Jobs & Steve Wozniak, Bill Gates & Paul Allen.  It marks the beginning of Piaget's formal operational stage of cognitive development.  The limbic, autonomic and hormone networks are already deployed and functioning effectively.  The frontal cortex has to be pruned: winning neurons move to their final highly connected positions, and are myelinated over time.  The rest dissolve.  So the frontal lobe does not obtain its adult configuration and networked integration until the mid-twenties when prefrontal cortex control becomes optimal.  The evolutionarily oldest areas of the frontal cortex mature first.  The PFC must be iteratively customized by experience to do the right thing as an adult.  Adolescents:
  • Don't detect irony effectively.  They depend on the DMPFC to do this, unlike adults who leverage the fusiform face area.  
  • Regulate emotions with the ventral striatum while the prefrontal cortex is still being setup.  Dopamine projection density and signalling increase from the ventral tegmentum catalyzing increased interest in dopamine based rewards.  Novelty seeking allows for creative exploration which was necessary to move beyond the familial pack.  Criticisms do not get incorporated into learning models by adolescents leaving their risk assessments very poor.  The target of the dopamine networks, the adolescent accumbens, responds to rewards like a gyrating top - hugely to large rewards, and negatively to small rewards.  Eventually as the frontal regions increase in contribution there are steady improvements in: working memory, flexible rule use, executive organization and task shifting.  And adolescents start to see other people's perspective. 
  • Drive the cellular transformations with post-pubescent high levels of testosterone in males, and high but fluctuating estrogen & progesterone levels in females.  Blood flow to the frontal cortex is also diverted on occasion to the groin.  
  • Peer pressure is exceptionally influential in adolescents.  Admired peer comments reduce vmPFC activity and enhance ventral striatal activity.  Adults modulate the mental impact of socially mean treatment: the initial activation of the PAG, anterior cingulate, amygdala, insula cortex; which generate feelings of pain, anger, and disgust, with the VLPFC but that does not occur in adolescents.  
  • Feel empathy intensely, supported by their rampant emotions, interest in novelty, ego.  But feeling the pain of others can induce self-oriented avoidance of the situations. 
- early adulthood, and then rates plummet.  Adolescent violence is driven by:

The
Matt Taibbi describes the phenotypic alignment of the American justice system.  The result he explains relentlessly grinds the poor and undocumented into resources to be constrained, consumed and ejected.  Even as it supports and aligns the financial infrastructure into a potent weapon capable of targeting any company or nation to extract profits and leave the victim deflated. 

Taibbi uses five scenarios to provide a broad picture of the: activities, crimes, policing, prosecutions, court processes, prisons and deportation network.  The scenarios are: Undocumented people's neighborhoods, Poor neighborhoods, Welfare recipients, Credit card debtors and Financial institutions.

Following our summary of his arguments, RSS comments on them framed by complex adaptive system (CAS) theory.  The alignment of the justice system reflects a set of long term strategies and responses to a powerful global arms race that the US leadership intends to win. 

criminal justice system
has to decide if adolescent criminals are to be considered fully in control of their judgements.  Sapolsky concludes that Behave's scientific foundation demands the transformation of the criminal justice system. 

Back to the Crib, Back to the Womb
Childhood is Sapolsky's next significant influence on our current good or bad behavior

Complexification
Childhood's progressively increasing
This page introduces the complex adaptive system (CAS) theory frame.  The theory is positioned relative to the natural sciences.  It catalogs the laws and strategies which underpin the operation of systems that are based on the interaction of emergent agents. 
John Holland's framework for representing complexity is outlined.  Links to other key aspects of CAS theory discussed at the site are presented. 
complexity
occurs in stages.  Neuron, specialized eukaryotic cells include channels which control flows of sodium and potassium ions across the massively extended cell membrane supporting an electro-chemical wave which is then converted into an outgoing chemical signal transmission from synapses which target nearby neuron or muscle cell receptors.  Neurons are supported by glial cells.  Neurons include a:
  • Receptive element - dendrites
  • Transmitting element - axon and synaptic terminals
  • Highly variable DNA schema using transposons. 
formation, migration and synaptogenesis are mostly prenatal in humans.  But myelination is the fatty insulating material deployed by Schwann cells & oligodendrocytes, both types of glial cells, around axons to improve their conduction rate.  In humans it is still occurring 25 years after birth.  It has great impact on long axons, in neurons that project over long distances, where it helps brain inter-region signalling.  The long development time of myelination allows for the later myelinated brain regions to be particularly shaped by the proximate environment. 
occurs when a region has setup its mature structure and is optimizing.  Sapolsky reiterates the later a brain region matures the more it is influenced by the environment: internal & external.  And he concludes that it is useful to remember that the significant aspects of childhood that will be highlighted all alter genes in the brain.  That is the only way it can work and the linkage reveals how bad outcomes might be reversed and good outcomes reinforced. 

Stages
For Jean Piaget the brain's developmental stages defines a series including the: Sensorimotor stage, Preoperational stage, Concrete operational stage, Formal operational stage; of child development.  It is an experimentally derived early model of behavioral development allowing an assessment of the: role of evolved stages, impact of experience, implications for the resulting adult; of the complex process.  Neural formation, migration and synaptogenesis are prenatal.  Myelination proceeds in stages for 25 years.  Sapolsky notes that Piaget's cognitive model ignores emotions and social factors which can't be right.  And while the network of mental agents cooperate to develop coherent responses, our developmental models should represent the separate development of the agents that process objects, faces etc., that allow a child to miss transitivity between objects but detect it in faces. 
were cognitive is the ability to orchestrate thought and action in accordance with internal goals according to Princeton's Jonathan Cohen. 
.  Sapolsky adds an early cognitive stage occurs when toddlers form ego boundaries are a cognitive developmental stage that starts when a child realizes there is a 'me,' separate from everyone else.  Prior to the ego boundary a toddler that sees his mother has cut her finger claims his finger hurts. 
.  By nine months infants understand that others have information they don't.  Older children eventually reach a stage where they achieve Theory of Mind of mind is the capability of adults, and even young children, to see that others think and perceive the world differently to them.  It typically develops around age three to four.  It supports the child's development of empathy.  It is associated with the DMPFC, precuneus, superior temporal sulcus & temporoparietal junction.  Subsequently more capabilities appear including: Understanding a second person's theory of mind about a third person, Perspectives and Irony.  .  But Sapolsky reminds us that Behave's premise is that cognitive development is a phase during the operation of a CAS agent.  It allows for schematic strategies to be iteratively blended with environmental signals to solve the logistical issues of migrating newly built and transformed sub-agents.  That is needed to achieve the adult configuration of the agent and optimize it for the proximate environment.  Smiley includes examples of the developmental phase agents required in an emergent CAS.  In situations where parents invest in the growth and memetic learning of their offspring the schematic grab bag can support optimizations to develop models, structures and actions to construct an adept adult.  In humans, adolescence leverages neural plasticity, elder sibling advice and adult coaching to help prepare the deploying neuronal network and body to successfully compete. 
can't be viewed in isolation from emotion and social effects. 

Feeling someone else's pain
Children next realize that others can have different feelings.  Sapolsky stresses the path to full empathy is the capability to relate to another person from their perspective.  It is implemented by spindle neurons.  It is context dependently mediated by estrogen.  It develops over time: Piaget's preoperational stage includes rudimentary empathy, Theory of mind supports the development; initially feeling someone's pain as one integrated being, then for them and eventually as them.  In adults, when someone else is hurt the anterior cingulate cortex, amygdala & insula activates projecting [scapegoating] to the vmPFC.  If the pain is physical the PAG activates and motor neurons for the area where the other person was injured.  The intertwining of the ACC amygdala & insula in adults results in attribution of fault even when there is none which can make it hard to step in and actually help.  But in seven-year-olds the activation is concrete: PAG and sensory & motor cortexes with minimal coupling to the rudimentary vmPFC.  In older children the vmPFC is coupled to limbic structures.  Ten to twelve year olds abstract empathy to classes of people.  By adolescence the vmPFC is coupled to theory of mind regions and intentional harm induces disgust via the amygdala.  Sapolsky explains adolescent boys are utilitarian and tend to accept inequality more than girls do.  But both sexes accept inequality as the way it is.  Sociopaths do not develop empathy. 
progresses in stages.  Young children's empathy makes no distinction between:
By age seven children express their own empathy.  By 10 to 12 empathy is abstracted and generalized.  And Sapolsky notes children first negatively stereotype categories of people. 

Similar stages are seen with responses to injustice is when someone has been treated unfairly.  Young children, aged four to six in all cultures, are aware of being treated unjustly.  By age eight to ten kids in some cultures respond negatively to others being treated unjustly.  At that point they attempt to rectify previous inequity.  But by pre-adolescence they are accepting of inequality, justified by merit, effort, or team needs.  And by adolescence boys accept injustice on utilitarian grounds and both sexes accept it can't be fixed; it's the way it is.  Mitigated free will argues that a person should be held responsible for their actions except for mitigating circumstances of a defect of reason produced by a psychosis. 


Moral Development
Kohlberg leveraged Piaget's ideas defines a series including the: Sensorimotor stage, Preoperational stage, Concrete operational stage, Formal operational stage; of child development.  It is an experimentally derived early model of behavioral development allowing an assessment of the: role of evolved stages, impact of experience, implications for the resulting adult; of the complex process.  Neural formation, migration and synaptogenesis are prenatal.  Myelination proceeds in stages for 25 years.  Sapolsky notes that Piaget's cognitive model ignores emotions and social factors which can't be right.  And while the network of mental agents cooperate to develop coherent responses, our developmental models should represent the separate development of the agents that process objects, faces etc., that allow a child to miss transitivity between objects but detect it in faces. 
including: Children working out rules of appropriate behavior; and experimentally demonstrated stages of moral development defines a three stage moral process:
  1. Preconventional reasoning (until age 8 - 10) initially allows aggression.  But around two to four years old adult and peer punishment inhibits the use of aggression.  And subsequently rewards become significant in the decision process.  When these judgements about aggression don't change at age 8 - 10 it predicts increasing risk of adult sociopathy.  For the others post conflict reconciliation decreases glucocorticoid levels and anxiety. 
  2. Conventional reasoning (adolescents and adults) reviews the consequences to others and if they are friends, as well as what others would likely do.  Being well regarded is also significant.  The law becomes significant in the later stage of this phase. 
  3. Post conventional reasoning (Inspiring heroes & insufferable pedants) first increases the value of self-judgement when judging rules.  And beyond this are moral acts valued because they represent a position more important than a law. 
.  He concluded such judgements are cognitive is the ability to orchestrate thought and action in accordance with internal goals according to Princeton's Jonathan Cohen. 
.  Sapolsky notes studies suggesting conservatives and liberals reason at different stages of Kohlberg's
The agents in complex adaptive systems (CAS) must model their environment to respond effectively to it.  Samuel modeling is described as an approach. 
model
.  But the studies focused on Americans, missing the cross-cultural moral differences and undervaluing intuition and emotion are low level agents distributed across the brain and body which associate, via the amygdala and rich club hubs, important environmental signals with encoded high speed sensors, and distributed programs of action to model: predict, prioritize guidance signals, select and respond effectively, coherently and rapidly to the initial signal.  The majority of emotion centered brain regions interface to the midbrain through the hypothalamus.  The most accessible signs of emotions are the hard to control and universal facial expressions.  Emotions provide prioritization for conscious access given that an animal has only one body, but possibly many cells, with which to achieve its highest level goals.  Because of this emotions clash with group goals and are disparaged by the powerful.  Evolutionary psychology argues evolution shaped human emotions during the long period of hunter-gatherer existence in the African savanna.  Human emotions are universal and include: Anger, Appreciation of natural beauty, Disgust, Fear, Gratitude, Grief, Guilt, Happiness, Honor, Jealousy, Liking, Love, Rage, Romantic love, Lust for revenge, Passion, Sadness, Self-control, Shame, Sympathy, Surprise; and the sham emotions and distrust induced by reciprocal altruism.  .  And they fail to predict who will do the harder-but-right thing.  Instead that is powerfully predicted by children's scores in Walter Mischel's experiments

Marshmallows
In the 1960s Mischel was studying gratification postponement and developed the marshmallow test is Stanford psychologist Walter Mischel's experiment studying gratification postponement.  A child is presented with a marshmallow and told that the experimenter is leaving the room and if the child doesn't eat the marshmallow before the experimenter returns he will be given another.  The child is then observed through a two way mirror.  Only a third of the children lasted the fifteen minutes.  To achieve the task the children had to: Trust in the system, avoid hot ideation of the marshmallow, or displace it with cold ideation of the marshmallow or hot ideation of some other object.   It was performed on three to six year olds, but subsequently predicted their SAT scores at high school, social success and lack of aggression, and forty years on more PFC activation during a frontal task and a lower BMI!  .  Significantly Mischel tracked his subject children's subsequent adult lives.  Forty years later, the children who were able to delay gratification had: Averaged higher SATs, More social success/resilience and less aggressive and oppositional behavior, Lower average BMI is body mass index.  s and excelled at frontal cortical of the cerebral cortex is at the front of the brain.  It includes the: prefrontal cortex, motor cortex.  Sapolsky asserts it makes you do the harder thing when it's the right thing to do.  The frontal cortex supports working memory to sustain focus on a task.  It also coordinates the strategic actions necessary to achieve success.  It provides impulse control, regulation of emotion, and willpower.  The prefrontal cortex maintains focus by deprioritizing currently irrelevant streams of information.  The frontal cortex tracks rules.  Over a lifetime that builds into a costly activity.  Once it tires responses become less prosocial.  But practice shifts operation of tasks to the cerebellum.  The frontal cortex signals the tegmentum and accumbens with the conclusions of its expectancy/discrepancy calculations.  The frontal lobe provides executive function, considering bits of information, assessing patterns and then prioritizing the strategies.  The frontal lobe is the most recent part of the brain to evolve and involves a disproportionate percentage of primate-unique genes in its development and operation.  It does not complete development until the mid-20s.  It includes spindle neurons.  It is easily damaged.  Sapolsky (Nauta) notes that its ventromedial prefrontal cortex is a quasi-member of the limbic system. 
function. 

Consequences
Sapolsky reviews what childhood events significantly contribute through biology to their adult best or worst behaviors: Mothers, Childhood adversity, Observing violence, Bullying; and subsequently explores: Why most children are still resilient enough to avoid the worst impacts, What happens when everything is wrong, The process by which parents mediate cultural memes into their children.   

Let's start at the very beginning: The importance of mothers
John Bowlby's attachment theory is John Bowlby's model of mother infant bonding.  He argued that infants need: love, warmth, affection, responsiveness, stimulation, consistency, reliability; or they become anxious, depressed, and/or poorly attached adults.  Evolutionarily, sociopaths may be highly successful as managers and leaders but they are probably anxious.  Sapolsky notes the powerful association between murder rates and stopping pregnant girls from terminating unwanted pregnancies.  Typical mothers also provide training on social conventions and their children's position in the group hierarchy.  Children raised without a mother's support fail to understand social constraints and when to use social behaviors.  And in the presence of unsupportive mothers newborns attach to negative stimuli.  This response is explained by the SHRP.  Abused children subsequently seek out abusive relationships as adults.  And a percentage of infants abused by their mothers become abusive mothers. 
launched the modern view of the importance of the mother-infant bond which was subsequently validated experimentally by Wisconsin's Harry Harlow in the infant monkey placed the infants with constructed mothers.  The mothers were made of wire mesh with a feeding bottle attached to the chest.  Some of the 'mothers' differed in being covered with terry cloth.  If the infants were given a choice they chose the terry-cloth mother, even if they were forced to feed from the wire cage mother.  And the infant monkeys that only experienced the wire cage mother and could not 'cling' failed to attach.  


And, the eventual explanation for a massive drop in crime rates observed starting in the 1990s, provides further evidence of mother-infant bond's significance in humans: Stanford's Donohue & Chicago's Levitt identified that the drop in violence correlated with state abortion legalization passed 20 years prior.  The problem was being born to a mother the child knows would prefer you don't exist.  Again Harlow's results confirmed Donohue & Levitt's proposals.  And the infant monkeys Harlow wrecked had the full set of behaviors, but didn't know when to use them effectively.  Without mothers to teach them, they had no social context. 

Any kind of mother in a storm
Sapolsky explains the SHRP is stress hypo-responsive period, an evolved strategy to protect the developing brain of a newborn from stress generated glucocorticoid based damage.  The many adverse effects on brain development of glucocorticoids induce an advantage from temporarily shrinking the adrenals.  SHRP makes sense since a newborn's mother will be better equipped to handle the stress.  During the SHRP any strong stimulus is assumed positive and the newborn attaches to it.  Without access to a mother the SHRP strategy is abandoned and the adrenals expand and secrete glucocorticoids. 
evolved to protect the very young brain from damage from glucocorticoids are corticosteroids which bind the glucocorticoid receptor.  They decrease excitability of prefrontal cortical neurons.  They have adverse effects in fetal/infant development having organizational effects on fetal brain construction and decreasing levels of: growth factors, neurons, synapses; resulting in an adult brain that is more sensitive to environmental triggers of depression and anxiety.  Glucocorticoids affect gene control structures and induce epi-genetic changes.  They have been found associated with high sodium chloride consumption (May 2017). 
.  But when mom is the source of the aversive stimuli the SHRP ensures abused children will seek similarly abusive relationships as adults, and the girls are more likely to become abusive mothers. 

Different routes to the same place
Most other significant events have similar impacts - Sapolsky groups them as forms of childhood adversity includes different types of stressor: No mother, Unsupportive mother, paternal deprivation, Childhood poverty, [Observing ]violence, Natural disasters, Bullying; which impact development and produce adult problems. 
  • The adversities are stressful and alter stress physiology producing children and adults with elevated: Glucocorticoids, CRH and ACTH, Sympathetic nervous system activity.  Early stress permanently impacts the brains ability to control glucocorticoid secretion.  The more stressors experienced and the less protective factors, the less likely it is that the child will cope and become a resilient adult.  The stressors expand the size and activity of the amygdala helping it ignore prefrontal cortex constraints.  And they degrade the dopamine network through impacts to the development of the mesolimbic system and elevated adult glucocorticoids depleting dopamine. 
  • The problems include attachment issues and adults with: depression (dopamine depletion and lowered thresholds making adult stressors more influential), anxiety, substance abuse (dopamine depletion, excessive adult exposure to glucocorticoids increasing drug craving & poorly developed frontal cortex), impaired cognitive abilities especially frontocortical with impaired hippocampal-dependent learning, impaired impulse control (amygdala), impaired emotional control, antisocial behavior and violence, relationships that replicate the childhood adversities.  Abused children who develop PTSD show decreased hippocampal volume.  Glucocorticoids decrease hippocampal production of BDNF.  Childhood poverty impacts development of the corpus callosum & ensures by kindergarten, poor marshmallow test performance.  Childhood poverty increases impacts from environmental stressors: Toxins, Liquor stores instead of fresh food markets, No transport infrastructure, Limited jobs in the immediate vicinity, Little access to low cost capital, Low positions in all social hierarchies.  
resulting in the same biological profile:
  • The adversities are stressful and alter stress physiology producing children and adults with elevated: Glucocorticoids, CRH and ACTH, Sympathetic nervous system activity.  Early stress permanently impacts the brains ability to control glucocorticoid secretion.  The more stressors experienced and the less protective factors, the less likely it is that the child will cope and become a resilient adult.  The stressors expand the size and activity of the amygdala helping it ignore prefrontal cortex constraints.  And they degrade the dopamine network through impacts to the development of the mesolimbic system and elevated adult glucocorticoids depleting dopamine. 
  • The problems include attachment issues and adults with: depression (dopamine depletion and lowered thresholds making adult stressors more influential), anxiety, substance abuse (dopamine depletion, excessive adult exposure to glucocorticoids increasing drug craving & poorly developed frontal cortex), impaired cognitive abilities especially frontocortical with impaired hippocampal-dependent learning, impaired impulse control (amygdala), impaired emotional control, antisocial behavior and violence, relationships that replicate the childhood adversities.  Abused children who develop PTSD show decreased hippocampal volume.  Glucocorticoids decrease hippocampal production of BDNF.  Childhood poverty impacts development of the corpus callosum & ensures by kindergarten, poor marshmallow test performance.  Childhood poverty increases impacts from environmental stressors: Toxins, Liquor stores instead of fresh food markets, No transport infrastructure, Limited jobs in the immediate vicinity, Little access to low cost capital, Low positions in all social hierarchies.  


Observing Violence
Exposure to scenes of violence such as: domestic abuse, warfare, murders, massacres; as children result in impaired concentration and impulse control Sapolsky explains.  Observed gun violence increases the child's adopting violence as a coping strategy.  Later as adults there is increased risk of depression, anxiety and aggression.  Even observing realistic violence indirectly on media causes the responses. 
desensitizes and normalizes the aggression of those already prone to violence. 

Bullying
While bullying is a complex childhood adversity.  Sapolsky explains that targets of bullying aren't selected at random.  And bullies often come from families with single mothers or younger parents with low educational and economic prospects.  Sapolsky notes that someone who bullies and is also bullied is likely to be a real mess as an adult. 
is one of the many forms of childhood adversity includes different types of stressor: No mother, Unsupportive mother, paternal deprivation, Childhood poverty, [Observing ]violence, Natural disasters, Bullying; which impact development and produce adult problems. 
  • The adversities are stressful and alter stress physiology producing children and adults with elevated: Glucocorticoids, CRH and ACTH, Sympathetic nervous system activity.  Early stress permanently impacts the brains ability to control glucocorticoid secretion.  The more stressors experienced and the less protective factors, the less likely it is that the child will cope and become a resilient adult.  The stressors expand the size and activity of the amygdala helping it ignore prefrontal cortex constraints.  And they degrade the dopamine network through impacts to the development of the mesolimbic system and elevated adult glucocorticoids depleting dopamine. 
  • The problems include attachment issues and adults with: depression (dopamine depletion and lowered thresholds making adult stressors more influential), anxiety, substance abuse (dopamine depletion, excessive adult exposure to glucocorticoids increasing drug craving & poorly developed frontal cortex), impaired cognitive abilities especially frontocortical with impaired hippocampal-dependent learning, impaired impulse control (amygdala), impaired emotional control, antisocial behavior and violence, relationships that replicate the childhood adversities.  Abused children who develop PTSD show decreased hippocampal volume.  Glucocorticoids decrease hippocampal production of BDNF.  Childhood poverty impacts development of the corpus callosum & ensures by kindergarten, poor marshmallow test performance.  Childhood poverty increases impacts from environmental stressors: Toxins, Liquor stores instead of fresh food markets, No transport infrastructure, Limited jobs in the immediate vicinity, Little access to low cost capital, Low positions in all social hierarchies.  
, Sapolsky singles it out because of the complexity: the targets aren't selected at random.  More often they have personal or family psychiatric issues and poor social and emotional intelligence.  and the bullies often come from families with single mothers or younger parents with low educational and economic prospects.  They may be bullied as they bully which results in very poorly adjusted adults who see bullying as ok and are likely to suffer from depression, anxiety and suicidality.  And otherwise bullies are either:
  • Anxious, isolated children with poor social skills aiming to achieve acceptance and reduce their own frustration. 
  • Future sociopaths. 
are the product of additional adversity and the targets aren't selected at random. 

A key question
Sapolsky considers why are many children resilient to childhood adversity includes different types of stressor: No mother, Unsupportive mother, paternal deprivation, Childhood poverty, [Observing ]violence, Natural disasters, Bullying; which impact development and produce adult problems. 
  • The adversities are stressful and alter stress physiology producing children and adults with elevated: Glucocorticoids, CRH and ACTH, Sympathetic nervous system activity.  Early stress permanently impacts the brains ability to control glucocorticoid secretion.  The more stressors experienced and the less protective factors, the less likely it is that the child will cope and become a resilient adult.  The stressors expand the size and activity of the amygdala helping it ignore prefrontal cortex constraints.  And they degrade the dopamine network through impacts to the development of the mesolimbic system and elevated adult glucocorticoids depleting dopamine. 
  • The problems include attachment issues and adults with: depression (dopamine depletion and lowered thresholds making adult stressors more influential), anxiety, substance abuse (dopamine depletion, excessive adult exposure to glucocorticoids increasing drug craving & poorly developed frontal cortex), impaired cognitive abilities especially frontocortical with impaired hippocampal-dependent learning, impaired impulse control (amygdala), impaired emotional control, antisocial behavior and violence, relationships that replicate the childhood adversities.  Abused children who develop PTSD show decreased hippocampal volume.  Glucocorticoids decrease hippocampal production of BDNF.  Childhood poverty impacts development of the corpus callosum & ensures by kindergarten, poor marshmallow test performance.  Childhood poverty increases impacts from environmental stressors: Toxins, Liquor stores instead of fresh food markets, No transport infrastructure, Limited jobs in the immediate vicinity, Little access to low cost capital, Low positions in all social hierarchies.  
?  He argues that:

A sledgehammer
Sadly 1980s Romanian infant institutions provided a human equivalent to Harlow's experimental setup in the infant monkey placed the infants with constructed mothers.  The mothers were made of wire mesh with a feeding bottle attached to the chest.  Some of the 'mothers' differed in being covered with terry cloth.  If the infants were given a choice they chose the terry-cloth mother, even if they were forced to feed from the wire cage mother.  And the infant monkeys that only experienced the wire cage mother and could not 'cling' failed to attach.  
.  Ceausescu's banning of abortion and requirement for families to have at least five children resulted in impoverished families abandoning newborns.  Overwhelmed institutions generated deprivation and neglect.  The children's lives were followed by Harvard's Charles Nelson who characterized them as adults:

Culture, with both a big and a little c
Prior to the general review of culture's effects on our best and worst behaviors, Sapolsky reviews how: culture is how we do and think about things, transmitted by non-genetic means as defined by Frans de Waal.  CAS theory views cultures as operating via memetic schemata evolved by memetic operators to support a cultural superorganism.  Evolutionary psychology asserts that human culture reflects adaptations generated while hunting and gathering.  Dehaene views culture as essentially human, shaped by exaptations and reading, transmitted with support of the neuronal workspace and stabilized by neuronal recycling.  Sapolsky argues that parents must show children how to transform their genetically derived capabilities into a culturally effective toolset.  He is interested in the broad differences across cultures of: Life expectancy, GDP, Death in childbirth, Violence, Chronic bullying, Gender equality, Happiness, Response to cheating, Individualist or collectivist, Enforcing honor, Approach to hierarchy; illustrating how different a person's life will be depending on the culture where they are raised.  Culture:
  • Is deployed during pregnancy & childhood, with parental mediation.  Nutrients, immune messages and hormones all affect the prenatal brain.  Hormones: Testosterone with anti-Mullerian hormone masculinizes the brain by entering target cells and after conversion to estrogen binding to intracellular estrogen receptors; have organizational effects producing lifelong changes.  Parenting style typically produces adults who adopt the same approach.  And mothering style can alter gene regulation in the fetus in ways that transfer epigenetically to future generations!  PMS symptoms vary by culture. 
  • Is also significantly transmitted to children by their peers during play.  So parents try to control their children's peer group.  
  • Is transmitted to children by their neighborhoods, tribes, nations etc. 
  • Influences the parenting style that is considered appropriate. 
  • Can transform dominance into honor.  There are ecological correlates of adopting honor cultures.  Parents in honor cultures are typically authoritarian. 
  • Is strongly adapted across a meta-ethnic frontier according to Turchin.  
  • Across Europe was shaped by the Carolingian empire. 
  • Can provide varying levels of support for innovation.  
  • Produces consciousness according to Dennet. 
influences childhood and parents mediate the process.  Sapolsky explains this begins with parenting style is a psychological classification including: authoritative, authoritarian, permissive, neglectful; ways of parenting that provide children with their first experience of proximate culture.  Each style typically produces adults that use that same approach.  Different cultures value different styles. 
.  He stresses that each style usually produces adults with that same approach, with different cultures valuing different styles. 
He adds transmission by peers from a young age as identified by Judith Harris is a book by Judith Rich Harris which argues children's personality is most significantly influenced by peers.  Her reasons include:
  • Parental influence is mediated by peers
  • Peers impact linguistic development
  • Other primates are socialized by peers, not mothers. 
in The Nurture Assumption.  Sapolsky explains that children play is a powerful tool for learning social competence.  It appears universal among socially complex species.  It has an evolved high priority displacing foraging and using up energy stores.  There are a number of behavioral strategies practiced in play that train those participating.  Play:
  • Supports safe practice of using social tools.  Allows for trying out roles and honing motor skills. 
  • Allows a peer group to provide context for these developments: Approach to hierarchy; etc.  Parents will aim to select the peer group. 
  • It integrates cultural details from local neighborhoods
  • It supports practice of aggression. 
  • Demonstrates that transient stress is enjoyable. 
  • It allows the developing neuron network to identify which synapses to prune. 
with peers, which builds social competence.  And neighborhoods transmit their cultural characteristics.  As do tribes, nations and states

Parents in cultures of honor tend to be authoritarian.  The children are aggressive and endorse violent responses to honor is a doomsday machine emotional signal, which Pinker explains as an advertisement of the desire to publically avenge even minor trespasses and insults.   violations. 

Collectivist versus individualist cultures
Mothers in individualist cultures, speak louder, play music louder, are more animated and see their role as teacher.  They encourage individually competitive games and hobbies where children do things.  The children are taught to be assertive, autonomous & influential.  They acquire theory of mind of mind is the capability of adults, and even young children, to see that others think and perceive the world differently to them.  It typically develops around age three to four.  It supports the child's development of empathy.  It is associated with the DMPFC, precuneus, superior temporal sulcus & temporoparietal junction.  Subsequently more capabilities appear including: Understanding a second person's theory of mind about a third person, Perspectives and Irony.   later than collectivist children. 

Mothers in collectivist cultures, spend more time soothing their children, maintaining contact, and facilitating contact with other adults.  They pick cooperative games.  They stress getting along, thinking of others, accepting and adapting.  These children see social competence as taking someone else's perspective. 

Class differences
St. Michael's College anthropologist Adrie Kusserow observed parent child interactions in three US is the United States of America.   neighborhoods:

Nine long months
The demonstration that fetuses were affected by what they tasted & heard outside the womb encouraged interest among the public.  For our best and worst behaviors Sapolsky judges other neonatal environmental influences are even more significant: Glands are active in the fetus and their action is transformative, Hostile external environment. 

Boy and girl brains, whatever that might mean
Sapolsky stresses that hormones are signalling molecules: ACTH, TRH, Melanocyte stimulating hormone, Testosterone, Oxytocin, Vasopressin, Insulin, Growth hormone, Estrogen, Progesterone, Angiotensin II, Asprosin, EPO, Irisin, Leptin, FGF21 hormone, Prostaglandins, TSH, Thyroxine, Glococorticoids; that are transported by the circulatory system to interact with target organs having appropriate receptors.  The levels of hormones can fluctuate massively, as in pregnancy. 
have lifelong organizational effects in the fetus, rather than the activation effects seen in adults.  Eight weeks post-conception: Male brains are masculinized by fetal gonads secreting testosterone is a hormone secreted by the testes in response to stimulation from the hypothalamic/pituitary/testicular cascade, that makes humans more willing to do what it takes to attain and maintain status, according to Sapolsky.  That means players of the Ultimatum Game, if previously given testosterone can become more generous.  Testosterones effect is highly socially contextual so it may encourage acts of kindness or aggression (when challenged).  The level of testosterone does not predict which individuals will be aggressive in: Birds, Fish, Mammals including primates.  Genes impact the potency of testosterone by altering the enzymes that: Construct it, Convert it to estrogen, code the androgen receptor.   This androgen receptor includes a variable polyglutamine repeat which alters the sensitivity to the testosterone signal.  The more potent form is associated with boys showing more dramatic 'masculinization' of the cortex.  But the detected genetic influences are small.  Testosterone decreases activity in the prefrontal cortex and its functional coupling to the amygdala while increasing the coupling between the amygdala & the thalamus.  Testosterone shortens the refactory period of amygdaloid & amygdaloid target neurons.  This results in impulsive risk taking and more focus on unfamiliar faces and distrust of them.  Testosterone increases activity in the ventral tegmentum projecting dopamine to enhance place preference.  Winners of fights become more willing to fight in part due to testosterone increasing confidence and optimism and reducing fear and anxiety.  And winning at: Chess, Athletics, Stock trades; induces the BNST to add testosterone receptors increasing its sensitivity to the hormone.  People become overconfident and overly optimistic. 
& "anti-Mullerian hormone," Female gonad's secret estrogen is a generic term for a number of related steroid hormones each of which works differently.  Estrogen:
  • Contributes to maternal aggression but it can reduce aggression and enhance empathy, depending on brain state.  There are two different estrogen receptor types which mediate these conflicting effects.  The levels of each type of receptor is independently regulated.  Different receptor variants are associated with:
    • Higher rates of anxiety among women
    • Higher rates of antisocial behavior and conduct disorder in men
and progesterone is a steroid hormone.  It:
  • Rarely directly effects areas of the brain.  Instead it is converted into other sterioids which have different actions in different brain areas. 
  • Increases maternal aggression in concert with estrogen by increasing oxytocin release in certain brain regions.  However, on its own progresterone decreases aggression and anxiety.  It decreases anxiety by entering neurons where it is converted to allopregnanolone which binds to GABA receptors increasing their sensitivity to GABA. 
.  But Sapolsky warns of messy complications:

Expanding the scope of "environment"
Many external factors have an impact:

Back to when you were just a fertilized egg
Sapolsky reviews the contribution of the
Plans emerge in complex adaptive systems (CAS) to provide the instructions that agents use to perform actions.  The component architecture and structure of the plans is reviewed. 
genome
to our best and worst behavior.  It is a contentious subject with
Russ Abbott explores the impact on science of epiphenomena and the emergence of agents. 
reductionists
arguing genes imply essentialism at one extreme and the other where people resist linking genes and behavior.  He notes that gene alleles, one of multiple alternative forms of a schematic sequence with the same address on a schematic string. 
affect:

Part 1: Genes from the bottom up
But Sapolsky concludes that while genes are important, they are not
Plans are interpreted and implemented by agents.  This page discusses the properties of agents in a complex adaptive system (CAS). 
It then presents examples of agents in different CAS.  The examples include a computer program where modeling and actions are performed by software agents.  These software agents are aggregates. 
The participation of agents in flows is introduced and some implications of this are outlined. 
agents
and they are constrained in what they can do by the 'environment':

Transcription factors in neurons, specialized eukaryotic cells include channels which control flows of sodium and potassium ions across the massively extended cell membrane supporting an electro-chemical wave which is then converted into an outgoing chemical signal transmission from synapses which target nearby neuron or muscle cell receptors.  Neurons are supported by glial cells.  Neurons include a:
  • Receptive element - dendrites
  • Transmitting element - axon and synaptic terminals
  • Highly variable DNA schema using transposons. 
are regulated by the
This page discusses the potential of the vast state space which supports the emergence of complex adaptive systems (CAS).  Kauffman describes the mechanism by which the system expands across the space. 
environment
: Intracellular where the immediate chemical state can activate a transcription factor, Adjacent neuron which signals, is an emergent capability which is used by cooperating agents to support coordination & rival agents to support control and dominance.  In eukaryotic cells signalling is used extensively.  A signal interacts with the exposed region of a receptor molecule inducing it to change shape to an activated form.  Chains of enzymes interact with the activated receptor relaying, amplifying and responding to the signal to change the state of the cell.  Many of the signalling pathways pass through the nuclear membrane and interact with the DNA to change its state.  Enzymes sensitive to the changes induced in the DNA then start to operate generating actions including sending further signals.  Cell signalling is reviewed by Helmreich.  Signalling is a fundamental aspect of CAS theory and is discussed from the abstract CAS perspective in signals and sensors.  In AWF the eukaryotic signalling architecture has been abstracted in a codelet based implementation.  To be credible signals must be hard to fake.  To be effective they must be easily detected by the target recipient.  To be efficient they are low cost to produce and destroy. 
this neuron across a synapse, a neuron structure which provides a junction with other neurons.  It generates signal molecules, either excitatory or inhibitory, which are kept in vesicles until the synapse is stimulated when the signal molecules are released across the synaptic cleft from the neuron.  The provisioning of synapses is under genetic control and is part of long term memory formation as identified by Eric Kandel.  Modulation signals (from slow receptors) initiate the synaptic strengthening which occurs in memory. 
, Distant organ that generates a hormone transported in the blood to a receptor on the neural circuit, a network of interconnected neurons which perform signalling, modeling and control functions.  In Cajal's basic neural circuits the signalling is unidirectional.  He identified three classes of neurons in the circuits:
  • Sensory, Interneurons, Motor; which are biochemically distinct and suffer different disease states. 
, Outside world of smells and sights.  The transcription factors and the genes they activate operate in a CAS where:
  • Genes encode the 1,500 different transcription factors
  • The human genome contains four million transcription factor binding sites. 
  • The presence of a particular set of transcription factors control the transcription of a network of genes, including those coding for other transcription factors.  
  • An average cell uses 200,000 transcription factor binding sites to generate its distinctive gene-expression profile. 
Sapolsky explains that epi-genetic represent state surfaces within cells and eggs which can be operationally modified so as to provide a heritable structure.  DNA, histones and other stable structures provide surfaces where these states may be setup.  Egg carriers are in a particularly powerful position to induce epi-genetic changes.  Sapolsky notes [childhood] events which persistently alter brain structure and behavior via epi-genetic mechanisms including: pair-bonding in prairie voles, as they first mate, is supported by changes in oxytocin & vasopressin receptor gene regulation in the nucleus accumbens. 
changes can reflect chemical & structural shifts which alter the ability of transcription factors to bind to the target DNA.  Lamarckian inheritance of environmental effects can occur over generations! 

And Sapolsky notes that proteins, a relatively long chain (polymer) of peptides.  Shorter chains of peptides are termed polypeptides.   are often constructed uses the product of the transcription of DNA into messenger m-RNA and is mostly the translation of the m-RNA into a folded protein by Ribosomes and t-RNA. 
from m-RNA (RNA), a polymer composed of a chain of ribose sugars.  It does not naturally form into a paired double helix and so is far less stable than DNA.  Chains of DNA are converted by transcription into equivalently sequenced messenger m-RNA.  RNA also provides the associations that encode the genetic code.  Transfer t-RNAs have a site that maps to the codon and match the associated amino-acid.  Stuart Kauffman argues that RNA polymers may be the precursor to our current DNA based genome and protein based enzymes.  In the adaptive web framework's (AWF) Smiley we use a similar paradigm with no proteins. 
sequences that contain non-coding introns.  Enzymes, a protein with a structure which allows it to operate as a chemical catalyst and a control switch. 
then remove the introns.  The mechanism enables multiple protein variants to be generated from one DNA sequence.  90% of human genes with exons are alternately spliced. 

Sapolsky considers the impact of transposons are regions of DNA which are copied and inserted into other areas of the DNA schemata.  Their effect and existence was first noted by Barbara McClintock.  Plants, when under stress, utilize transposons to rapidly evolve their genomes.  Animal's immune systems and neuronal stem cells when forming memories, also leverage transposons.  , which he notes operate within neuronal stem cells is a biological cell which is partly or wholly undifferentiated.  A totipotent cell can generate a complete embryo and placenta.  Embryos include pluripotent cells which can generate any tissue in the body.  Adult humans' cells have turned off this ability but still include multipotent stem cells that differentiate into multiple cell types.   Typically a cell's local environment will have the signals required for it to obtain context and differentiate appropriately.  This will include both the external environment and the internal state of the cell which has replicated from a parent and obtained its epi-genetic state.   So introduction of undifferentiated stem cells into an injured area is not likely to have either aspect of the environment suitable.  Consequently development is aiming to encourage differentiation to progenitor cells for the damaged region.  This requires delivering the cells to the appropriate part of the body.  To avoid rejection by the immune system techniques aim to use cell lines developed from the patient's cells.  The techniques to generate these cell lines include: SCNT, iPS.  Possible mechanisms of stem cell therapy are: Generation of new differentiated cells, Stimulation of growth of new blood vessels to repopulate damaged regions, Secretion of growth factors, Treatment of diabetes (1 and 2) with addition of pancreatic cells, Assistance of other mechanisms; in the brain expanding the genome of neurons forming memories. 

Part 2: Genes from the top down--Behavior Genetics
Sapolsky argues that 'behavioral genetics' had struggled to quantify the magnitude of genetic effects on: IQ, Sexual orientation; forcing additional approaches: 
Part 3: So what do genes actually have to do with behaviors we're interested in?
Sapolsky notes that behavioral genetics has beneficially incorporated molecular approaches.  But many interesting findings are complicated by gene/environment interactions:


Centuries to millennia before
Sapolsky notes that the cultures is how we do and think about things, transmitted by non-genetic means as defined by Frans de Waal.  CAS theory views cultures as operating via memetic schemata evolved by memetic operators to support a cultural superorganism.  Evolutionary psychology asserts that human culture reflects adaptations generated while hunting and gathering.  Dehaene views culture as essentially human, shaped by exaptations and reading, transmitted with support of the neuronal workspace and stabilized by neuronal recycling.  Sapolsky argues that parents must show children how to transform their genetically derived capabilities into a culturally effective toolset.  He is interested in the broad differences across cultures of: Life expectancy, GDP, Death in childbirth, Violence, Chronic bullying, Gender equality, Happiness, Response to cheating, Individualist or collectivist, Enforcing honor, Approach to hierarchy; illustrating how different a person's life will be depending on the culture where they are raised.  Culture:
  • Is deployed during pregnancy & childhood, with parental mediation.  Nutrients, immune messages and hormones all affect the prenatal brain.  Hormones: Testosterone with anti-Mullerian hormone masculinizes the brain by entering target cells and after conversion to estrogen binding to intracellular estrogen receptors; have organizational effects producing lifelong changes.  Parenting style typically produces adults who adopt the same approach.  And mothering style can alter gene regulation in the fetus in ways that transfer epigenetically to future generations!  PMS symptoms vary by culture. 
  • Is also significantly transmitted to children by their peers during play.  So parents try to control their children's peer group.  
  • Is transmitted to children by their neighborhoods, tribes, nations etc. 
  • Influences the parenting style that is considered appropriate. 
  • Can transform dominance into honor.  There are ecological correlates of adopting honor cultures.  Parents in honor cultures are typically authoritarian. 
  • Is strongly adapted across a meta-ethnic frontier according to Turchin.  
  • Across Europe was shaped by the Carolingian empire. 
  • Can provide varying levels of support for innovation.  
  • Produces consciousness according to Dennet. 
that surround us have huge effects on how individuals behave.  These behaviors reflect the culture the individuals were raised in as well as where they live.  He illustrates his case with:
  • Gender differences in mathematics where the more gender equality the less difference there was in math scores with Iceland's girls scoring better than boys, while most other countries have boys scoring better especially in unequal cultures like Turkey's. 
    • Earlier adoption of the hoe over the plow predicts gender equality in today's culture. 
  • Long term impacts - population density in 1500 significantly predicts how authoritarian a government was in 2000. 
He explores where we are now and how we probably got here: Patterns of cultural variation effect behaviors, Culture & biology coevolve, and Ecology effects culture. 

Life is unrecognizably different, depending on which culture you were born into:

Hobbes or Rousseau- how we got there
Sapolsky explains that contemporary evaluations of the applicability of these philosophies leverage data:
  • Archaeological
  • Contemporary humans living in pre-state tribal societies
But there are definitional disagreements.  University of Illinois's Lawrence Keeley and Harvard's Steven Pinker argue archaeological evidence of war is broad, ancient, & barbaric and violence is declining.  They argue that archaeologists are ignoring the evidence to pacify the past.  Rutgers R. Brian Ferguson strikes back arguing: Their evidence does not show what they claim and they cherry-picked their data. 

Keeley, Pinker & Santa Fe Institute's Samuel Bowles conclude from looking at contemporary hunter-gatherers: New Guinea: Gebusi, Mae Enga; & Borneo head hunters, Maasai & Zulu warriors from Africa, Amazonians: Waorani, Jivaro; Brazilian and Venezuelan Yanomamos; that warfare is nearly universal.  Anthropologists again argue there was cherry-picking with Global surveys showing less violence.  There are issues with the statistical analyses.  And other studies do not find the levels of violence of the initial reports. 

Sapolsky argues 95 to 99% of hominin history was spent in small nomadic bands that foraged for edible plants and hunted cooperatively.  Few records exist: cave paintings from forty thousand years ago which show hunting but not war.  So most people infer from the current day hunter-gatherer tribes the: Hazda, Mbuti, Batwa, Gunwinggi, Andaman Islanders, Batak, Semang and Inuit cultures.  Mostly the men discuss how fantastic their last hunt was while generations of females provide the calories from foraging.  Most hunter-gatherers:
Sapolsky writes the evidence suggests a mixture of Hobbes & Rousseau.  He does not consider hunter-gatherers as angels but sees war as rare until the nomadic lifestyle was abandoned. 

So Sapolsky suggests agriculture/herding was one of the all-time human blunders because it:

He concludes by asserting the most consequential aspect of the coevolution of brains and culture is how we do and think about things, transmitted by non-genetic means as defined by Frans de Waal.  CAS theory views cultures as operating via memetic schemata evolved by memetic operators to support a cultural superorganism.  Evolutionary psychology asserts that human culture reflects adaptations generated while hunting and gathering.  Dehaene views culture as essentially human, shaped by exaptations and reading, transmitted with support of the neuronal workspace and stabilized by neuronal recycling.  Sapolsky argues that parents must show children how to transform their genetically derived capabilities into a culturally effective toolset.  He is interested in the broad differences across cultures of: Life expectancy, GDP, Death in childbirth, Violence, Chronic bullying, Gender equality, Happiness, Response to cheating, Individualist or collectivist, Enforcing honor, Approach to hierarchy; illustrating how different a person's life will be depending on the culture where they are raised.  Culture:
  • Is deployed during pregnancy & childhood, with parental mediation.  Nutrients, immune messages and hormones all affect the prenatal brain.  Hormones: Testosterone with anti-Mullerian hormone masculinizes the brain by entering target cells and after conversion to estrogen binding to intracellular estrogen receptors; have organizational effects producing lifelong changes.  Parenting style typically produces adults who adopt the same approach.  And mothering style can alter gene regulation in the fetus in ways that transfer epigenetically to future generations!  PMS symptoms vary by culture. 
  • Is also significantly transmitted to children by their peers during play.  So parents try to control their children's peer group.  
  • Is transmitted to children by their neighborhoods, tribes, nations etc. 
  • Influences the parenting style that is considered appropriate. 
  • Can transform dominance into honor.  There are ecological correlates of adopting honor cultures.  Parents in honor cultures are typically authoritarian. 
  • Is strongly adapted across a meta-ethnic frontier according to Turchin.  
  • Across Europe was shaped by the Carolingian empire. 
  • Can provide varying levels of support for innovation.  
  • Produces consciousness according to Dennet. 
is childhood, when cultures inculcate individuals.  The delayed maturation in humans supports the transition from a juvenile configuration, dependent on parents and structured to learn & logistically transform, to adult optimized to the proximate environment.  And it is staged encouraging the adolescents to escape the hierarchy they grew up in and enter other groups where they may bring in: fresh ideas, risk taking; and alter the existing hierarchy: Steve Jobs & Steve Wozniak, Bill Gates & Paul Allen.  It marks the beginning of Piaget's formal operational stage of cognitive development.  The limbic, autonomic and hormone networks are already deployed and functioning effectively.  The frontal cortex has to be pruned: winning neurons move to their final highly connected positions, and are myelinated over time.  The rest dissolve.  So the frontal lobe does not obtain its adult configuration and networked integration until the mid-twenties when prefrontal cortex control becomes optimal.  The evolutionarily oldest areas of the frontal cortex mature first.  The PFC must be iteratively customized by experience to do the right thing as an adult.  Adolescents:
  • Don't detect irony effectively.  They depend on the DMPFC to do this, unlike adults who leverage the fusiform face area.  
  • Regulate emotions with the ventral striatum while the prefrontal cortex is still being setup.  Dopamine projection density and signalling increase from the ventral tegmentum catalyzing increased interest in dopamine based rewards.  Novelty seeking allows for creative exploration which was necessary to move beyond the familial pack.  Criticisms do not get incorporated into learning models by adolescents leaving their risk assessments very poor.  The target of the dopamine networks, the adolescent accumbens, responds to rewards like a gyrating top - hugely to large rewards, and negatively to small rewards.  Eventually as the frontal regions increase in contribution there are steady improvements in: working memory, flexible rule use, executive organization and task shifting.  And adolescents start to see other people's perspective. 
  • Drive the cellular transformations with post-pubescent high levels of testosterone in males, and high but fluctuating estrogen & progesterone levels in females.  Blood flow to the frontal cortex is also diverted on occasion to the groin.  
  • Peer pressure is exceptionally influential in adolescents.  Admired peer comments reduce vmPFC activity and enhance ventral striatal activity.  Adults modulate the mental impact of socially mean treatment: the initial activation of the PAG, anterior cingulate, amygdala, insula cortex; which generate feelings of pain, anger, and disgust, with the VLPFC but that does not occur in adolescents.  
  • Feel empathy intensely, supported by their rampant emotions, interest in novelty, ego.  But feeling the pain of others can induce self-oriented avoidance of the situations. 
of the frontal cortex of the cerebral cortex is at the front of the brain.  It includes the: prefrontal cortex, motor cortex.  Sapolsky asserts it makes you do the harder thing when it's the right thing to do.  The frontal cortex supports working memory to sustain focus on a task.  It also coordinates the strategic actions necessary to achieve success.  It provides impulse control, regulation of emotion, and willpower.  The prefrontal cortex maintains focus by deprioritizing currently irrelevant streams of information.  The frontal cortex tracks rules.  Over a lifetime that builds into a costly activity.  Once it tires responses become less prosocial.  But practice shifts operation of tasks to the cerebellum.  The frontal cortex signals the tegmentum and accumbens with the conclusions of its expectancy/discrepancy calculations.  The frontal lobe provides executive function, considering bits of information, assessing patterns and then prioritizing the strategies.  The frontal lobe is the most recent part of the brain to evolve and involves a disproportionate percentage of primate-unique genes in its development and operation.  It does not complete development until the mid-20s.  It includes spindle neurons.  It is easily damaged.  Sapolsky (Nauta) notes that its ventromedial prefrontal cortex is a quasi-member of the limbic system. 
allows for sculpting by the environment including cultural norms. 


The Evolution of Behavior
Sapolsky asserts that only
This page reviews the implications of selection, variation and heredity in a complex adaptive system (CAS).  The mechanism and its emergence are discussed. 
evolution
can explain behavior.  He argues like anatomy and physiology, behaviors evolve. 

Kin selection is a strategy of selfish genes, which aims to maximize gene survival & replication across all the bodies where a copy of the gene probably exists: relatives.  Altruism is beneficial to gene replication in this situation.  Love supports the agent's prioritization of appropriate altruistic strategies.  Sapolsky describes an array of strategies used to identify kin:
  • Genetically shaped pheromonal signatures.  Rodents leverage the immune systems MHC super variable gene regions to develop unique signals.  The more similar the signals are the closer is the relative.  Pregnancy triggers adult neurogenesis in the olfactory system of rat mothers to allow them to learn the smell of their newborn.
  • Imprinting on the female whose birdsong a chick heard while still in the egg
  • Degree of paternalism depending on likelihood of being the father in primates
  • Humans use cognition
allows relatives to group together to help their genes persist into the next generation.  Sapolsky notes that across species mating third-cousins is an evolved behavior.  Women prefer the smell of moderately related over unrelated men.  Identifying kin works differently across species.  Human kin recognition is mainly cognitive is the ability to orchestrate thought and action in accordance with internal goals according to Princeton's Jonathan Cohen. 
.  Sapolsky notes it is far from accurate which is significant. 

Non-kin can also evolve to cooperate through reciprocal altruism benefits another organism at a cost to the behaver.  It is differentiated from kin altruism, by Williams and Trivers, since it can apply between unrelated individuals.  It can be induced by natural selection when there is mutual survival benefit in group activities and cheating can be detected and discouraged.  Humans, leveraging the cognitive niche, can particularly easily, build an evolved amplifier, through sharing information at little cost and significant benefit.  But African savanna hunters similarly gain from sharing large game meat with other un-related altruistic group members since the meat would otherwise spoil before it could be eaten. 
.  Sapolsky notes the strategies to maximize cooperation and limit cheating have been investigated using game theory investigates the optimal strategies for cooperating.  The iterative Prisoner's Dilemma scenario was shown by Axelrod & Hamilton to reward a Tit for Tat strategy with perfect signals.  An effective way to limit the risks of initiating Tit for Tat in a potential sea of Always Defectors is a genetically encoded signal of cooperation such as the green-beard gene.  Other evolved strategies include genetic algorithms. 
including
Plans change in complex adaptive systems (CAS) due to the action of genetic operations such as mutation, splitting and recombination.  The nature of the operations is described. 
genetic algorithms


Sapolsky views the three foundations of behavioral evolution: individual
This page reviews the implications of selection, variation and heredity in a complex adaptive system (CAS).  The mechanism and its emergence are discussed. 
natural
&
This page describes the consequences of the asymmetries caused by genotypic traits creating a phenotypic signal in males and selection activity in the female - sexual selection.   
The impact of this asymmetry is to create a powerful alternative to natural selection with sexual selection's leverage of positive returns.  The mechanisms are described. 
sexual
selection, kin selection, reciprocal altruism; as a lens to understand:
But he notes that neo-group selection is David Sloan Wilson and subsequently Harvard's E.O. Wilson's, multi-level evolutionary selection mechanism.  It is seen to operate in humans and is explained by Tit for Tat style cooperation and prosociality. 
is also a significant force in evolving humans' behaviors. 

Still Sapolsky stresses that we are not descended from any of the other living species.  He reviews a variety of measures that all suggest we are profoundly confused.  We exhibit behaviors: Violence driven by male-male competition for reproductive access to females, Violence against females for coercive sex or punishing rejection; that make sense to an elephant seal or a baboon have asymmetric sizes, and musculature.  Males may be much bigger and more muscular than females and have conspicuous facial markings.  Sapolsky's tournament primates include: baboons, mandrills, rhesus monkeys, vervets, and chimps.  And he additionally lists gazelles, lions, sheep, peacocks, & elephant seals.  Selection has supported evolved capture of fighting skills and display.  Such males:
  • Use aggressive conflict to obtain high dominance rankings.  
  • A small percentage of high rank males do all the mating.  They will mate with any female anytime.  So they have evolved to invest in larger testes and higher sperm counts. 
  • Males do no investment in parenting of their children.  
  • Females look for signals of good genes.  This encourages sexual selection of signals of male health, status and dominance.  
  • Females don't compete with each other since they will all breed with the high rank males. 
.  But other behaviors don't fit that mold: People forgoing reproduction, Individuals sacrificing themselves for strangers. 

Sapolsky sees broad evidence of human kin selection.  But there are complications:
  • Clan fighting occurs - but many wars show evidence of families fighting on both sides. 
  • Intra-family individual violence occurs
  • Patricide occurs, typically as revenge for abuse and fratricide. 
  • Parents killing children because of mental illness, abuse that unintentionally turned fatal
  • Money is bequeathed to non-kin.  Sapolsky judges that our cognitive based kin assessment helps explain these situations.  If you 'feel' like a relative to me you are a relative.  And we can be manipulated into feeling more or less related to someone than we really are. 
Sapolsky asserts reciprocal altruism and neo-group selection drive: rules, laws, treaties, penalties, social conscience, inner voice, morals, ethics, etc.  He notes that a study of present day hunter-gatherers showed, only 40% of people within the bands are blood relatives.  So we potentially evolved from hunter-gatherers that were driven by reciprocal altruism.  And neo-group selection is at the core of cooperation and competition among human groups and cultures is how we do and think about things, transmitted by non-genetic means as defined by Frans de Waal.  CAS theory views cultures as operating via memetic schemata evolved by memetic operators to support a cultural superorganism.  Evolutionary psychology asserts that human culture reflects adaptations generated while hunting and gathering.  Dehaene views culture as essentially human, shaped by exaptations and reading, transmitted with support of the neuronal workspace and stabilized by neuronal recycling.  Sapolsky argues that parents must show children how to transform their genetically derived capabilities into a culturally effective toolset.  He is interested in the broad differences across cultures of: Life expectancy, GDP, Death in childbirth, Violence, Chronic bullying, Gender equality, Happiness, Response to cheating, Individualist or collectivist, Enforcing honor, Approach to hierarchy; illustrating how different a person's life will be depending on the culture where they are raised.  Culture:
  • Is deployed during pregnancy & childhood, with parental mediation.  Nutrients, immune messages and hormones all affect the prenatal brain.  Hormones: Testosterone with anti-Mullerian hormone masculinizes the brain by entering target cells and after conversion to estrogen binding to intracellular estrogen receptors; have organizational effects producing lifelong changes.  Parenting style typically produces adults who adopt the same approach.  And mothering style can alter gene regulation in the fetus in ways that transfer epigenetically to future generations!  PMS symptoms vary by culture. 
  • Is also significantly transmitted to children by their peers during play.  So parents try to control their children's peer group.  
  • Is transmitted to children by their neighborhoods, tribes, nations etc. 
  • Influences the parenting style that is considered appropriate. 
  • Can transform dominance into honor.  There are ecological correlates of adopting honor cultures.  Parents in honor cultures are typically authoritarian. 
  • Is strongly adapted across a meta-ethnic frontier according to Turchin.  
  • Across Europe was shaped by the Carolingian empire. 
  • Can provide varying levels of support for innovation.  
  • Produces consciousness according to Dennet. 


Sapolsky notes that these gene driven behavioral arguments were intended to be threatened by Punctuated Equilibrium is Steven Jay Gould and Niles Eldridge's proposal about how evolution occurs.  They suggested that most of the time 'nothing happens' and then there are intermittent rapid lurches.  The idea is analogous to Schumpeter's waves of creative destruction.  Complex adaptive systems will operate this way driven by the underlying slow mutations of the germ-line schemata and the action of infrastructure and evolved amplifiers. 
.  While Dawkins dismissed macro mutations and species selection, there is evidence of rapid gene spreading and our understanding of transcription-factors are enzymes which associate with a transcription complex to bind to the DNA and control its transcription and hence translation into proteins.  The regulation of DNA transcription and protein synthesis are reviewed by Tsonis.  Transcription factors allow environmental state to become reflected in the control of DNA transcription.  Transcription factors can regulate multiple genes, allowing network effects & multiple transcription factors can regulate a gene allowing sophisticated control processes.  In AWF the transcription, translation and deployment infrastructure of the eukaryotic cell has been abstracted in a codelet based implementation. 
, transposons are regions of DNA which are copied and inserted into other areas of the DNA schemata.  Their effect and existence was first noted by Barbara McClintock.  Plants, when under stress, utilize transposons to rapidly evolve their genomes.  Animal's immune systems and neuronal stem cells when forming memories, also leverage transposons.   and other
This page reviews the strategy of setting up an arms race.  At its core this strategy depends on being able to alter, or take advantage of an alteration in, the genome or equivalent.  The situation is illustrated with examples from biology, high tech and politics. 
amplifiers
enables us to accept macro-mutational events.  Still Sapolsky concludes that depending on what genes are being affected, gradualism and punctuated change occur in evolution without the edifice collapsing. 

Steven Jay Gould's tinkering exaptation, initially termed preadaptation refers to the coopting of some function for a new use.   allows evolution to produce good enough solutions.  They suggest other traits were purely spandrels are Steven Jay Gould and Richard Lewontin's evolved traits that appear as side effects of exaptations and have no selection value themselves.  A spandrel is an architectural term for the space between two arches.  Gould and Lenontin's point was that adaptationists would suggest spandrels were created to provide a medium for the artwork that was often placed on them because they were there. 
.  Sapolsky suggests there are less adaptations than socio-biologists once asserted and less spandrels than Gould and other critics argued, with reality anchored in the complexity of evolution.  


Us Versus Them
Sapolsky explores our tendency to form 'Us' versus 'Them' dichotomies and to favor the former, as detected by the IAT is the implicit association test:
  • Pictures of likely in-group and out-group members and positive and negative words are displayed on a computer screen
  • Sometimes the viewer is asked "If you see an in-group member or positive word press the red button; if you see an out-group or negative word press the blue button." 
  • Alternatively the viewer may be asked "If you see an in-group member or negative term press the red button; if you see an out-group or positive word press the blue button." 
  • The discordance of the second rule causes a pause of a few milliseconds before the viewer presses a button.  Over a series of views a pattern of delay emerges indicating the viewer's bias. 
.  He notes how this grouping process is supported by the brain's architecture and happens rapidly:
Bristol's Tajfel showed 'Us' versus 'Them' formed rapidly in minimal group paradigms, demonstrating prosociality is prioritizing benefiting others through: Help, Charity, Truth; even if many are acts of restitution to balance out antisocial acts.  Due to the Insula's binding of physical and metaphorical disgust, physically washing your hands can be enough to reduce the need for a prosocial act.  
requires only meaningless traits for group identification.  It emerges before age three.  Sapolsky argues this augments genetic green-beard genes code for expressing an evolved signal, such as a green-beard, and cooperating with other "green-beard displaying" people.  Humans can adjust how we view these signals and so how we define us and them. 
effects which depend on both kin selection is a strategy of selfish genes, which aims to maximize gene survival & replication across all the bodies where a copy of the gene probably exists: relatives.  Altruism is beneficial to gene replication in this situation.  Love supports the agent's prioritization of appropriate altruistic strategies.  Sapolsky describes an array of strategies used to identify kin:
  • Genetically shaped pheromonal signatures.  Rodents leverage the immune systems MHC super variable gene regions to develop unique signals.  The more similar the signals are the closer is the relative.  Pregnancy triggers adult neurogenesis in the olfactory system of rat mothers to allow them to learn the smell of their newborn.
  • Imprinting on the female whose birdsong a chick heard while still in the egg
  • Degree of paternalism depending on likelihood of being the father in primates
  • Humans use cognition
and reciprocal altruism benefits another organism at a cost to the behaver.  It is differentiated from kin altruism, by Williams and Trivers, since it can apply between unrelated individuals.  It can be induced by natural selection when there is mutual survival benefit in group activities and cheating can be detected and discouraged.  Humans, leveraging the cognitive niche, can particularly easily, build an evolved amplifier, through sharing information at little cost and significant benefit.  But African savanna hunters similarly gain from sharing large game meat with other un-related altruistic group members since the meat would otherwise spoil before it could be eaten. 
; with psychological equivalents. 

Sapolsky suggests what helps define a culture is how we do and think about things, transmitted by non-genetic means as defined by Frans de Waal.  CAS theory views cultures as operating via memetic schemata evolved by memetic operators to support a cultural superorganism.  Evolutionary psychology asserts that human culture reflects adaptations generated while hunting and gathering.  Dehaene views culture as essentially human, shaped by exaptations and reading, transmitted with support of the neuronal workspace and stabilized by neuronal recycling.  Sapolsky argues that parents must show children how to transform their genetically derived capabilities into a culturally effective toolset.  He is interested in the broad differences across cultures of: Life expectancy, GDP, Death in childbirth, Violence, Chronic bullying, Gender equality, Happiness, Response to cheating, Individualist or collectivist, Enforcing honor, Approach to hierarchy; illustrating how different a person's life will be depending on the culture where they are raised.  Culture:
  • Is deployed during pregnancy & childhood, with parental mediation.  Nutrients, immune messages and hormones all affect the prenatal brain.  Hormones: Testosterone with anti-Mullerian hormone masculinizes the brain by entering target cells and after conversion to estrogen binding to intracellular estrogen receptors; have organizational effects producing lifelong changes.  Parenting style typically produces adults who adopt the same approach.  And mothering style can alter gene regulation in the fetus in ways that transfer epigenetically to future generations!  PMS symptoms vary by culture. 
  • Is also significantly transmitted to children by their peers during play.  So parents try to control their children's peer group.  
  • Is transmitted to children by their neighborhoods, tribes, nations etc. 
  • Influences the parenting style that is considered appropriate. 
  • Can transform dominance into honor.  There are ecological correlates of adopting honor cultures.  Parents in honor cultures are typically authoritarian. 
  • Is strongly adapted across a meta-ethnic frontier according to Turchin.  
  • Across Europe was shaped by the Carolingian empire. 
  • Can provide varying levels of support for innovation.  
  • Produces consciousness according to Dennet. 
is a standardized process binding arbitrary markers: dress, ornamentation, accent; to normally invisible meaningful traits: Values, beliefs, attributions, ideologies; allowing a flag to become a symbol to die for

Sapolsky notes that the 'Us' process inflates the merits of in-group core values: correct, wise, moral, and worthy; and arbitrary markers.  Feelings about 'Us' focus on shared obligations, willingness and mutuality from a high frequency of iterative positive interactions and the power to punish defections.  But the prosociality for the in-group is bound quite abstractly.  Sapolsky also reports that we act to improve things for 'Us' AND to maximize the gap with 'Them'.  Maximizing the gap can result in focusing on loyalty rather than equality.  An in-group member, who airs the group's dirty laundry generating a negative stereotype and in-group shame is an emotional reaction to being discovered cheating on a friend. 
, will provoke high levels of punishment as a signal, is an emergent capability which is used by cooperating agents to support coordination & rival agents to support control and dominance.  In eukaryotic cells signalling is used extensively.  A signal interacts with the exposed region of a receptor molecule inducing it to change shape to an activated form.  Chains of enzymes interact with the activated receptor relaying, amplifying and responding to the signal to change the state of the cell.  Many of the signalling pathways pass through the nuclear membrane and interact with the DNA to change its state.  Enzymes sensitive to the changes induced in the DNA then start to operate generating actions including sending further signals.  Cell signalling is reviewed by Helmreich.  Signalling is a fundamental aspect of CAS theory and is discussed from the abstract CAS perspective in signals and sensors.  In AWF the eukaryotic signalling architecture has been abstracted in a codelet based implementation.  To be credible signals must be hard to fake.  To be effective they must be easily detected by the target recipient.  To be efficient they are low cost to produce and destroy. 
to outsiders.  When someone leaves 'Us' to join 'Them' it may invoke stiff punishment if the commitment to 'Us' is based on sacred values rather than a contract. 

Similarly Sapolsky sees patterns in how we view 'Them':
Sapolsky notes that humans display unique aspects of 'Us' versus 'Them'.  They can maintain:

Hierarchy, Obedience, and Resistance

Sapolsky argues that hierarchies are about our automatic tendency to favor people close in rank to us over those who are distant.  And he notes that as with 'Us' versus 'Them' these tendencies appear:
have intertwined cognitive is the ability to orchestrate thought and action in accordance with internal goals according to Princeton's Jonathan Cohen. 
& affective are low level agents distributed across the brain and body which associate, via the amygdala and rich club hubs, important environmental signals with encoded high speed sensors, and distributed programs of action to model: predict, prioritize guidance signals, select and respond effectively, coherently and rapidly to the initial signal.  The majority of emotion centered brain regions interface to the midbrain through the hypothalamus.  The most accessible signs of emotions are the hard to control and universal facial expressions.  Emotions provide prioritization for conscious access given that an animal has only one body, but possibly many cells, with which to achieve its highest level goals.  Because of this emotions clash with group goals and are disparaged by the powerful.  Evolutionary psychology argues evolution shaped human emotions during the long period of hunter-gatherer existence in the African savanna.  Human emotions are universal and include: Anger, Appreciation of natural beauty, Disgust, Fear, Gratitude, Grief, Guilt, Happiness, Honor, Jealousy, Liking, Love, Rage, Romantic love, Lust for revenge, Passion, Sadness, Self-control, Shame, Sympathy, Surprise; and the sham emotions and distrust induced by reciprocal altruism.   underpinnings and the categorizations and status is a publically accepted, signal that one possesses assets: wealth, beauty, talent, expertise, access & trust of powerful people; to be able to help others. 
assessments interact.  But Sapolsky notes hierarchy leads in a different & uniquely human direction:

Nature and varieties of hierarchies
Across species hierarchies:

Rank and hierarchy in humans
Human hierarchies resemble those of other species:
Human hierarchies have unique aspects:
  • Membership in multiple hierarchies - which can overlap and conflict. 
  • The specialization of some ranking systems - while high ranking chimps generally excel at related areas, humans specialize deeply and build hierarchies around these subcultures.  It is not obvious these skills generalize. 
  • Internal standards that don't depend on the real world

The view from the top, the view from the bottom
Rank is so important to humans that we dedicate brain activity to judging it: Sapolsky reviews neurobiological research on hierarchy:
An SES (socioeconomic status) is:
  • The most permeating form of status among primates.  It is reflected in the SES health gradient where
  • Lower life expectancy and higher incidence and morbidity of numerous diseases for poor people.  Stress related diseases are most significant: cardiovascular, gastrointestinal, psychiatric;
  • Health gets worse for each point down the SES health gradient scale and is mostly associated with psychological impacts of poverty: feeling poor, obvious income inequality. 
  • Poverty instigates poor health with low-SES wombs associated with poorer health as the fetus becomes adult.  
  • The SES health gradient is seen in countries with socialized medicine and universal health care and expresses in diseases that don't require health care access. 
  • Environmental risk factors associated with being poor explain less than a third of the SES health gradient effect. 
health gradient is cross-culturally ubiquitous.  Material inequality powerfully subjugates the low ranking. 

A really odd thing that we do now and then
We have and choose leaders aims to develop plans and strategies which ensure effective coordination to improve the common good of the in-group.  John Adair developed a leadership methodology based on the three-circles model. 
.  Sapolsky notes that is different to a primate power hierarchy.  Sapolsky suggests that in choosing leaders we:

Oh, Why not take this one on? Politics and political orientations

Obedience and conformity, disobedience and nonconformity
Sapolsky notes that obedience and conformity are closely linked.  Both reflect value-free terms.  Agreement supports the wisdom of crowds. 


Morality and Doing the Right Thing, Once You've Figured Out What That Is
Sapolsky reviews whether morality provides rules for identifying right from wrong.  It develops in stages with children using play to work out rules of appropriate behavior.  Kohlberg's 1950s experiments using children led him to conclude moral judgement is a cognitive process that develops in three stages.  Sapolsky raises issues with the framework: Its a model, It does not apply to other cultures, Intuition & emotion are as significant as cognition, Moral reasoning doesn't predict moral actions; and notes the capacity of the frontal cortex to regulate emotions and behavior is far more predictive.  The marshmallow test, performed on three to six year olds, actually predicted their subsequent SAT scores at high school, social success and lack of aggression, and forty years on more PFC activation during a frontal task and a lower BMI!  Jonathan Haidt argues people's moral decisions are rationalizations rather than using reasoning. 
is based on emotions are low level agents distributed across the brain and body which associate, via the amygdala and rich club hubs, important environmental signals with encoded high speed sensors, and distributed programs of action to model: predict, prioritize guidance signals, select and respond effectively, coherently and rapidly to the initial signal.  The majority of emotion centered brain regions interface to the midbrain through the hypothalamus.  The most accessible signs of emotions are the hard to control and universal facial expressions.  Emotions provide prioritization for conscious access given that an animal has only one body, but possibly many cells, with which to achieve its highest level goals.  Because of this emotions clash with group goals and are disparaged by the powerful.  Evolutionary psychology argues evolution shaped human emotions during the long period of hunter-gatherer existence in the African savanna.  Human emotions are universal and include: Anger, Appreciation of natural beauty, Disgust, Fear, Gratitude, Grief, Guilt, Happiness, Honor, Jealousy, Liking, Love, Rage, Romantic love, Lust for revenge, Passion, Sadness, Self-control, Shame, Sympathy, Surprise; and the sham emotions and distrust induced by reciprocal altruism.   or cognition is the ability to orchestrate thought and action in accordance with internal goals according to Princeton's Jonathan Cohen. 
according to scientific investigation: 
Context
Sapolsky argues that both reasoning and intuition contribute to moral decisions.  He reviews what situations increase one aspect or the other with the help of Josh Greene's neuroscience experiments uses fMRI to follow brain activity of subjects during Josh Greene's experiments with the trolley problem.  He used two scenarios, (1) The five people are doomed, would you pull the lever so the trolley will hit and kill someone: (2) Would you push someone onto the tracks to stop the trolley?  Consistently 60-70% of people say yes to scenario (1) but only 30% say yes to scenario (2).  The fMRI showed the more the vmPFC and/or amygdala activate the more likely the person is to refuse to act in either case.  Greene concludes that intuitions about intentionality are what is changing.  Pushing that kills someone feels morally wrong.  Greene developed a third scenario where the subject throws the switch to save the five but in doing so pushes a person out of the way and in falling to the ground they die.  This seems more acceptable.  Then Greene adjusts scenario (1) making the side loop switch back to the main track so that if the trolley is not stopped by the body of a person the five will still die - the same result as scenario (2).  Described this way 60-70% find pulling the lever acceptable.  Greene concludes our intuitions are very local and the additional indirection in this last scenario stops us feeling disturbed. 
with the trolley problem is British philosopher Philippa Foot's thought experiment in ethics, which asks if you have the opportunity to switch the tracks ahead of a runaway trolley car so that it kills one person instead of the five on the main track should you: Do nothing or Pull the lever and kill the one.  Foot asks what would be the most ethical choice.  .  Sapolsky concludes moral decision making is highly context dependent, shifting the locality/language impact on intuitionist morals.  Sapolsky highlights Alain Cohn's report on banking's impact on morals was studied by University of Zurich's Alain Cohn.  He used a coin-toss game with financial rewards for guessing outcomes correctly and possibilities to cheat, and for the researchers to detect the cheating.  Cohn then primed the test subjects to implicitly think about their profession.  For bankers ONLY, thinking about their profession, or the word banking, increased rates of cheating by 20%.  The bankers carried two sets of ethical rules concerning cheating in their heads.  Context is important in making moral decisions. 
- bank employees cheating increased 20% when they thought about banking, other industries employees did not show the effect!
Fools rush in: applying the findings of the science of morality
Sapolsky wonders how these insights can help us be at our best. 

Feeling Someone's Pain, Understanding Someone's Pain, Alleviating Someone's Pain
Sapolsky notes that empathy is the capability to relate to another person from their perspective.  It is implemented by spindle neurons.  It is context dependently mediated by estrogen.  It develops over time: Piaget's preoperational stage includes rudimentary empathy, Theory of mind supports the development; initially feeling someone's pain as one integrated being, then for them and eventually as them.  In adults, when someone else is hurt the anterior cingulate cortex, amygdala & insula activates projecting [scapegoating] to the vmPFC.  If the pain is physical the PAG activates and motor neurons for the area where the other person was injured.  The intertwining of the ACC amygdala & insula in adults results in attribution of fault even when there is none which can make it hard to step in and actually help.  But in seven-year-olds the activation is concrete: PAG and sensory & motor cortexes with minimal coupling to the rudimentary vmPFC.  In older children the vmPFC is coupled to limbic structures.  Ten to twelve year olds abstract empathy to classes of people.  By adolescence the vmPFC is coupled to theory of mind regions and intentional harm induces disgust via the amygdala.  Sapolsky explains adolescent boys are utilitarian and tend to accept inequality more than girls do.  But both sexes accept inequality as the way it is.  Sociopaths do not develop empathy. 
's underlying biology includes sensorimotor, emotional are low level agents distributed across the brain and body which associate, via the amygdala and rich club hubs, important environmental signals with encoded high speed sensors, and distributed programs of action to model: predict, prioritize guidance signals, select and respond effectively, coherently and rapidly to the initial signal.  The majority of emotion centered brain regions interface to the midbrain through the hypothalamus.  The most accessible signs of emotions are the hard to control and universal facial expressions.  Emotions provide prioritization for conscious access given that an animal has only one body, but possibly many cells, with which to achieve its highest level goals.  Because of this emotions clash with group goals and are disparaged by the powerful.  Evolutionary psychology argues evolution shaped human emotions during the long period of hunter-gatherer existence in the African savanna.  Human emotions are universal and include: Anger, Appreciation of natural beauty, Disgust, Fear, Gratitude, Grief, Guilt, Happiness, Honor, Jealousy, Liking, Love, Rage, Romantic love, Lust for revenge, Passion, Sadness, Self-control, Shame, Sympathy, Surprise; and the sham emotions and distrust induced by reciprocal altruism.   and cognitive is the ability to orchestrate thought and action in accordance with internal goals according to Princeton's Jonathan Cohen. 
building blocks that ensure varied forms which can be sharpened or dulled by quite logical influences.  He investigates when does empathy encourage us to do something helpful? When we do act, whose benefit is it for?  He concludes our
This page reviews the implications of selection, variation and heredity in a complex adaptive system (CAS).  The mechanism and its emergence are discussed. 
evolution
made us focus on local acts and bound together altruism, is the property that since kin share genes natural selection will improve the replicator's selfish goals by supporting the survival of such relatives.  Improving the chances of survival of non-kin is hard to explain with a gene preservation theory.  Why help a competitive gene?  Trivers explanation of reciprocal altruism shows the special conditions under which it can occur. 
and reciprocity benefits another organism at a cost to the behaver.  It is differentiated from kin altruism, by Williams and Trivers, since it can apply between unrelated individuals.  It can be induced by natural selection when there is mutual survival benefit in group activities and cheating can be detected and discouraged.  Humans, leveraging the cognitive niche, can particularly easily, build an evolved amplifier, through sharing information at little cost and significant benefit.  But African savanna hunters similarly gain from sharing large game meat with other un-related altruistic group members since the meat would otherwise spoil before it could be eaten. 
.  And that it helps to have generated learned habits for automatically acting compassionately. 

"For" versus "as if" and other distinctions
Sapolsky notes that empathy is the capability to relate to another person from their perspective.  It is implemented by spindle neurons.  It is context dependently mediated by estrogen.  It develops over time: Piaget's preoperational stage includes rudimentary empathy, Theory of mind supports the development; initially feeling someone's pain as one integrated being, then for them and eventually as them.  In adults, when someone else is hurt the anterior cingulate cortex, amygdala & insula activates projecting [scapegoating] to the vmPFC.  If the pain is physical the PAG activates and motor neurons for the area where the other person was injured.  The intertwining of the ACC amygdala & insula in adults results in attribution of fault even when there is none which can make it hard to step in and actually help.  But in seven-year-olds the activation is concrete: PAG and sensory & motor cortexes with minimal coupling to the rudimentary vmPFC.  In older children the vmPFC is coupled to limbic structures.  Ten to twelve year olds abstract empathy to classes of people.  By adolescence the vmPFC is coupled to theory of mind regions and intentional harm induces disgust via the amygdala.  Sapolsky explains adolescent boys are utilitarian and tend to accept inequality more than girls do.  But both sexes accept inequality as the way it is.  Sociopaths do not develop empathy. 
, sympathy is an emotion, the desire to help those in need.  Steven Pinker suggests it may develop into a sham emotion to earn gratitide.  Sapolsky adds that it can describe someone with the power to help, but who choses not to.  Alternately it can indicate feeling sorry for someone elses pain while not understanding it, in contrast with empathy.  Or it can mean the emotionally distanced sense of feeling for someone.  Or the state of feeling their pain as if it were happening to you where it may cause such distress as to focus you onto alleviating you own distress. 
, and compassion indicates an emotional state where resonance with someone else's distress leads one to help them. 
are suitcase words have multiple attached meanings which encourage us to think in different ways about the word.  Suitcase words are reviewed by Marvin Minsky. 
that are about internally motivated states.  And insights about these states can be found by observing other animals. 

Emotionally contagious, compassionate animals
Sapolsky asserts many animals:
But Sapolsky admits it is not possible to tell if the animals are being self-serving or compassionate indicates an emotional state where resonance with someone else's distress leads one to help them. 


Emotionally contagious, compassionate children
Sapolsky reminds us of the prior discussion about children & adolescents:
Again Sapolsky notes it is difficult to tell if children are hoping to end a sufferer's distress or their own. 

Affect and/or cognition?
The ACC is either the
  • Anterior cingulate cortex which:
    • Is a central focus of empathy supporting people relating to other's pain.  This is dependent on oxytocin. 
    • In non-human mammals it processes interoceptive signals.  The ACC focuses the internal signals into high level 'gut intuitions.'  Pain catches the ACC's attention. 
    • Performs discrepancy detection from the outcome that was predicted - at a high level.  The ACC cares about the meaning of what is predicted. 
      • If the ACC has been convinced that a pain killer placebo has inhibited pain signals, the ACC will stay silent about actual pain that is signalled from interoceptive networks.  
      • The ACC will signal: physical pain, emotional pain, metaphorical pain, anxiety, disgust, embarrassment, social exclusion especially in adolescence; as one and the same.  The ACC's abnormalities being associated with major depression. 
    • Has a bridging role between the empathetic and self-interested pain monitor.  Sapolsky notes the ACC is essential for learning fear and conditioned avoidance by observation alone through an intermediate step of shared representation of self.  He concludes "At its core the ACC is about self-interest, with caring about the other person in pain as an add-on." 
  • American College of Cardiology
is the center of empathy is the capability to relate to another person from their perspective.  It is implemented by spindle neurons.  It is context dependently mediated by estrogen.  It develops over time: Piaget's preoperational stage includes rudimentary empathy, Theory of mind supports the development; initially feeling someone's pain as one integrated being, then for them and eventually as them.  In adults, when someone else is hurt the anterior cingulate cortex, amygdala & insula activates projecting [scapegoating] to the vmPFC.  If the pain is physical the PAG activates and motor neurons for the area where the other person was injured.  The intertwining of the ACC amygdala & insula in adults results in attribution of fault even when there is none which can make it hard to step in and actually help.  But in seven-year-olds the activation is concrete: PAG and sensory & motor cortexes with minimal coupling to the rudimentary vmPFC.  In older children the vmPFC is coupled to limbic structures.  Ten to twelve year olds abstract empathy to classes of people.  By adolescence the vmPFC is coupled to theory of mind regions and intentional harm induces disgust via the amygdala.  Sapolsky explains adolescent boys are utilitarian and tend to accept inequality more than girls do.  But both sexes accept inequality as the way it is.  Sociopaths do not develop empathy. 
in humans, but in other species it processes interoceptive signals indicate the body's internal state: Pain, Fatigue; seconds to minutes before.  The interoceptive 'networks' project to brain regions that implement social emotions. 
and monitors disconnects from what it predicted.  Sapolsky notes these later two roles intersect in unexpected pain amplifies the aggression response of people by interoceptive signalling of brain regions providing social emotions including the PAG projecting to the amygdala; making aggressive people more so and less aggressive people less so.  Pain is the main reason people visit the ED in the US.   and its meaning, including for social and emotional pain suggesting the ACC is self-oriented and worried about ones well-being rather than empathy for others.  So is an empathic state about one's self?  Sapolsky notes the ACC also allows us to 'empathetically feel' someone's pain, under the influence of oxytocin is a peptide hormone which makes humans more prosocial to and socially competent in their in-group and more antisocial to everyone else.  The effects are contingent; changing during stress and in the presence of a threatening outgroup.  Oxytocin makes people look at eyes longer, encouraging improved accuracy at perceiving emotions.  It enhances activity in the TPJ supporting modeling of other people's thinking.  Dogs and their owners secrete oxytocin increasing the amount of eye contact between them.  It is associated with pair bonding.  It is central to female mammals wanting to nurse, nursing, and remembering their child.  Its effects are context dependent and so is the regulation of the genes that control oxytocin.  Variants of a gene CD38 which facilitates oxytocin secretion from neurons are associated with differing levels of activation of the fusiform face area when looking at faces.  Sapolsky describes an oxytocin receptor gene variant that is associated with children showing: Extreme aggression, A callous unemotional style; foreshadowing adult psychopathy.  And another receptor gene variant which is associated with childhood social disconnection and unstable adult relationships.  Gene/environment interactions complicate the interpretation of the presence of particular gene variants.  Hypothalamic neurons send projections to: ventral tegmentum which also becomes more receptive during child birth, nucleus accumbens, hippocampus, amygdala where it inhibits the central amygdala suppressing fear & anxiety consistently in men while still allowing women to respond to threats to their infants, frontal cortex, olfactory network where it helps new rat mums to learn the smell of their offspring; where oxytocin prepares the brain for in-group bonding, out-grouping, birth and maternal behavior.  Outside the brain hypothalamic neurons in females send oxytocin to the posterior pituitary where it enters the blood stream stimulating uterine contraction during labor & supporting milk production for weaning.  Disorders associated with oxytocin abnormalities include ASD. 
, which helps us learn what someone else is going through and bond with them -- an add-on to self-interest. 

The insula is part of the cerebral cortex folded deep within the lateral sulcus.  It includes: anterior, posterior insula; and is overlaid by the operculum.  It is assumed to participate in consciousness where it has been linked to emotion, salience & body homeostasis functions:
  • Perception,
  • Motor control: Hand-&-eye motor movement, Swallowing, Gastric motility, Speech articulation;
  • Self-awareness,
  • Inter-personal experiences: Disgust at smells, contamination & mutilation which generate visceral responses, that are projected to the amygdala; binding physical and moral aspects of purity (Macbeth effect)
  • Homeostatic regulation of the sympathetic network, parasympathetic network, and immune system.  
and amygdala contains > 12 distinct areas: Central, Lateral.  It receives simple signals from the lower parts of the brain: pain from the PAG; and abstract complex information from the highest areas: Disgust from the insula cortex.  It sends signals to almost every other part of the brain, including to the decision-making circuitry of the frontal lobes.  It has high levels of D(1) dopamine receptors.  During extreme fear the amygdala drives the hippocampus into fear learning.  It outputs directly to subcortical reflexive motor pathways when speed is required.  Its central nucleus projects to the BNST.  It signals the locus ceruleus.  The amygdala:
  • Promotes aggression.  Stimulating the amygdala promotes rage.  It converts anger into aggression and when impaired it impacts the ability to detect angry facial expressions.  
  • Participates in disgust
  • Perceives fear promoting stimuli.  In PTSD sufferers the Amygdala overreacts to mildly fearful stimuli and is slow to calm down and the amygdala expands in size over a period of months.  Fear is processed by the lateral nucleus which serves as the input from various senses, and the central nucleus which outputs to the brain stem (central grey - freezing, lateral hypothalamus - blood pressure, activates paraventricular hypothalamus => crf -> hormone adjustments). 
  • Has lots of receptors for and is highly sensitive to glucocorticoids.  Stress inhibits the GABA interneurons in the basolateral amygdala (BLA) allowing the excitatory glutamate releasing neurons to excite more. 
  • Is sensitive to unsettling/uncertain social situations where it promotes anxiety.  It is also interested in uncertain but potentially painful situations.  The amygdala contributes to social and emotional decision making where the BLA supports rejecting an unacceptable offer, as allowed in the Ultimatum Game, by injecting implicit mistrust and vigilance, generating an anger driven rejection that is used as punishment.  The amygdala is very rapidly excited by subliminal signals from the thalamus of outgroup skin color.  The amygdala subsequently tips social emotions against outgroups unless restrained by the frontal lobe or influenced by subliminal priming to prioritize inclusion.  The fast path from the thalamus rapidly but inaccurately signals its identified a weapon. 
  • Promotes male, but not female, sexual motivation when it is an uncertain potential pleasure. 
  • Responds to the longing for uncertain potential pleasures and fear that the reward will not be worth it if it happens.  The amygdala turns off during orgasm. 
  • Uses but is not directly involved in vision. 
are also involved in developing full empathy. 

The cognitive is the ability to orchestrate thought and action in accordance with internal goals according to Princeton's Jonathan Cohen. 
dlPFC is dorsolateral prefrontal cortex which is:
  • At the heart of decision making - highly rational, unsentimental 
  • A major agent of working memory
  • The most recently evolved part of the prefrontal cortex.  
  • Mainly interconnected with other parts of the cortex.  
and theory of mind of mind is the capability of adults, and even young children, to see that others think and perceive the world differently to them.  It typically develops around age three to four.  It supports the child's development of empathy.  It is associated with the DMPFC, precuneus, superior temporal sulcus & temporoparietal junction.  Subsequently more capabilities appear including: Understanding a second person's theory of mind about a third person, Perspectives and Irony.  
This page discusses the effect of the network on the agents participating in a complex adaptive system (CAS).  Small world and scale free networks are considered. 
networks
: TPJ is the temporoparietal juncture: left, right; where the temporal and parietal lobes meet, an area of the cortex with projections from the thalamus, limbic system, visual, auditory and somatosensory networks.  It supports modeling of self and others. 
& superior central sulcus; help figure out what's going on.  This is especially true for issues of causation and intentionality.  These take shape when children distinguish between self- and other-inflicted pain.  Sapolsky concludes cognitive processes act as a gatekeeper, deciding whether a particular misfortune is worthy of empathy.  And cognition helps when the pain is more abstract or seen indirectly.  Human empathy, like in rodents, is contingent on 'Us' rather than 'Them', unless cognition intervenesWith 'Us' empathy is the capability to relate to another person from their perspective.  It is implemented by spindle neurons.  It is context dependently mediated by estrogen.  It develops over time: Piaget's preoperational stage includes rudimentary empathy, Theory of mind supports the development; initially feeling someone's pain as one integrated being, then for them and eventually as them.  In adults, when someone else is hurt the anterior cingulate cortex, amygdala & insula activates projecting [scapegoating] to the vmPFC.  If the pain is physical the PAG activates and motor neurons for the area where the other person was injured.  The intertwining of the ACC amygdala & insula in adults results in attribution of fault even when there is none which can make it hard to step in and actually help.  But in seven-year-olds the activation is concrete: PAG and sensory & motor cortexes with minimal coupling to the rudimentary vmPFC.  In older children the vmPFC is coupled to limbic structures.  Ten to twelve year olds abstract empathy to classes of people.  By adolescence the vmPFC is coupled to theory of mind regions and intentional harm induces disgust via the amygdala.  Sapolsky explains adolescent boys are utilitarian and tend to accept inequality more than girls do.  But both sexes accept inequality as the way it is.  Sociopaths do not develop empathy. 
activates the ACC is either the
  • Anterior cingulate cortex which:
    • Is a central focus of empathy supporting people relating to other's pain.  This is dependent on oxytocin. 
    • In non-human mammals it processes interoceptive signals.  The ACC focuses the internal signals into high level 'gut intuitions.'  Pain catches the ACC's attention. 
    • Performs discrepancy detection from the outcome that was predicted - at a high level.  The ACC cares about the meaning of what is predicted. 
      • If the ACC has been convinced that a pain killer placebo has inhibited pain signals, the ACC will stay silent about actual pain that is signalled from interoceptive networks.  
      • The ACC will signal: physical pain, emotional pain, metaphorical pain, anxiety, disgust, embarrassment, social exclusion especially in adolescence; as one and the same.  The ACC's abnormalities being associated with major depression. 
    • Has a bridging role between the empathetic and self-interested pain monitor.  Sapolsky notes the ACC is essential for learning fear and conditioned avoidance by observation alone through an intermediate step of shared representation of self.  He concludes "At its core the ACC is about self-interest, with caring about the other person in pain as an add-on." 
  • American College of Cardiology
but with 'Them' the TPJ is the temporoparietal juncture: left, right; where the temporal and parietal lobes meet, an area of the cortex with projections from the thalamus, limbic system, visual, auditory and somatosensory networks.  It supports modeling of self and others. 


Across the socioeconomic spectrum, the wealthier is schematically useful information and its equivalent, schematically useful energy, to paraphrase Beinhocker.  It is useful because an agent has schematic strategies that can utilize the information or energy to extend or leverage control of the cognitive niche.  
people are (or just feel): The less they empathize with people in distress, The poorer they are at recognizing other's emotions are low level agents distributed across the brain and body which associate, via the amygdala and rich club hubs, important environmental signals with encoded high speed sensors, and distributed programs of action to model: predict, prioritize guidance signals, select and respond effectively, coherently and rapidly to the initial signal.  The majority of emotion centered brain regions interface to the midbrain through the hypothalamus.  The most accessible signs of emotions are the hard to control and universal facial expressions.  Emotions provide prioritization for conscious access given that an animal has only one body, but possibly many cells, with which to achieve its highest level goals.  Because of this emotions clash with group goals and are disparaged by the powerful.  Evolutionary psychology argues evolution shaped human emotions during the long period of hunter-gatherer existence in the African savanna.  Human emotions are universal and include: Anger, Appreciation of natural beauty, Disgust, Fear, Gratitude, Grief, Guilt, Happiness, Honor, Jealousy, Liking, Love, Rage, Romantic love, Lust for revenge, Passion, Sadness, Self-control, Shame, Sympathy, Surprise; and the sham emotions and distrust induced by reciprocal altruism.  , The greedier they are and the more likely they are to cheat or steal; which is due to seeing these things as beneath 'Us.' 

Sapolsky concludes for empathetic states, emotion and cognition both apply but the balance between them shifts with more cognitive work needed as the other person becomes more different. 

A mythic leap forward 
Sapolsky explains that University of Parma's Rizzolatti and Gallese identified mirror neurons are part of the premotor cortex, responding to signals from the PFC and sending on signals to the motor cortex to drive muscles.  This subset of Pre-motor cortex neurons, also respond to observation of other animals performing the same act requested by the PFC.  The mirroring can be abstract: See and hear activate same mirror neurons, Activate only when the action has the same intentionality; and none has been shown to be causally related yet. 
.  Sapolsky notes that some people, including UC San Diego's Ramachandran, have suggested they can help support empathy is the capability to relate to another person from their perspective.  It is implemented by spindle neurons.  It is context dependently mediated by estrogen.  It develops over time: Piaget's preoperational stage includes rudimentary empathy, Theory of mind supports the development; initially feeling someone's pain as one integrated being, then for them and eventually as them.  In adults, when someone else is hurt the anterior cingulate cortex, amygdala & insula activates projecting [scapegoating] to the vmPFC.  If the pain is physical the PAG activates and motor neurons for the area where the other person was injured.  The intertwining of the ACC amygdala & insula in adults results in attribution of fault even when there is none which can make it hard to step in and actually help.  But in seven-year-olds the activation is concrete: PAG and sensory & motor cortexes with minimal coupling to the rudimentary vmPFC.  In older children the vmPFC is coupled to limbic structures.  Ten to twelve year olds abstract empathy to classes of people.  By adolescence the vmPFC is coupled to theory of mind regions and intentional harm induces disgust via the amygdala.  Sapolsky explains adolescent boys are utilitarian and tend to accept inequality more than girls do.  But both sexes accept inequality as the way it is.  Sociopaths do not develop empathy. 
.  But there is no evidence of a causal link. 

The core issue: Actually Doing Something
Sapolsky asks what actually predicts that someone will do something compassionate indicates an emotional state where resonance with someone else's distress leads one to help them. 
to lessen another's pain.  Sapolsky argues:

Metaphors We Kill By
Sapolsky argues language separates a signal, is an emergent capability which is used by cooperating agents to support coordination & rival agents to support control and dominance.  In eukaryotic cells signalling is used extensively.  A signal interacts with the exposed region of a receptor molecule inducing it to change shape to an activated form.  Chains of enzymes interact with the activated receptor relaying, amplifying and responding to the signal to change the state of the cell.  Many of the signalling pathways pass through the nuclear membrane and interact with the DNA to change its state.  Enzymes sensitive to the changes induced in the DNA then start to operate generating actions including sending further signals.  Cell signalling is reviewed by Helmreich.  Signalling is a fundamental aspect of CAS theory and is discussed from the abstract CAS perspective in signals and sensors.  In AWF the eukaryotic signalling architecture has been abstracted in a codelet based implementation.  To be credible signals must be hard to fake.  To be effective they must be easily detected by the target recipient.  To be efficient they are low cost to produce and destroy. 
's message from its meaning and this
Rather than oppose the direct thrust of some environmental flow agents can improve their effectiveness with indirect responses.  This page explains how agents are architected to do this and discusses some examples of how it can be done. 
indirection
had significant benefits: Able to represent past and future emotions are low level agents distributed across the brain and body which associate, via the amygdala and rich club hubs, important environmental signals with encoded high speed sensors, and distributed programs of action to model: predict, prioritize guidance signals, select and respond effectively, coherently and rapidly to the initial signal.  The majority of emotion centered brain regions interface to the midbrain through the hypothalamus.  The most accessible signs of emotions are the hard to control and universal facial expressions.  Emotions provide prioritization for conscious access given that an animal has only one body, but possibly many cells, with which to achieve its highest level goals.  Because of this emotions clash with group goals and are disparaged by the powerful.  Evolutionary psychology argues evolution shaped human emotions during the long period of hunter-gatherer existence in the African savanna.  Human emotions are universal and include: Anger, Appreciation of natural beauty, Disgust, Fear, Gratitude, Grief, Guilt, Happiness, Honor, Jealousy, Liking, Love, Rage, Romantic love, Lust for revenge, Passion, Sadness, Self-control, Shame, Sympathy, Surprise; and the sham emotions and distrust induced by reciprocal altruism.  , Messages unrelated to emotion, Lying enabling the prisoner's dilemma is a game theoretical scenario explored by mathematicians and evolutionary biologists.  Two prisoners are to be interrogated separately.  They will benefit if they cooperate with one another - serving one year each.  But if just one defects they will go free while the other will lose serving three years.  If they both defect they will both lose serving two years each.  If the game is played once it is best to defect.  But if the game is iterative, cooperation can be beneficial using a Tit for Tat strategy.  Bootstrapping cooperation can be induced:
  • When a small founder population first becomes isolated and kin-selection increases and then the group connect to the main population.  The group can then use cooperation to out compete the other members.  
  • Through green-beard signals, driving cooperation and forcing non green-beards to shift to cooperation to compete. 
.  He suggests metaphor is the height of the symbolic features of language.  Metaphors are deployed liberally in everyday language.  He now explores the neurobiology of symbolic and metaphorical thinking.  He notes that the very recent evolution of these capabilities is reflected in our brains improvising to cope with metaphor.  And our brain is struggling to distinguish between the metaphorical and literal.  Sapolsky writes that this has enormous consequences for our best and worst behaviors

Feeling someone else's pain
Sapolsky reviews the ACC is either the
  • Anterior cingulate cortex which:
    • Is a central focus of empathy supporting people relating to other's pain.  This is dependent on oxytocin. 
    • In non-human mammals it processes interoceptive signals.  The ACC focuses the internal signals into high level 'gut intuitions.'  Pain catches the ACC's attention. 
    • Performs discrepancy detection from the outcome that was predicted - at a high level.  The ACC cares about the meaning of what is predicted. 
      • If the ACC has been convinced that a pain killer placebo has inhibited pain signals, the ACC will stay silent about actual pain that is signalled from interoceptive networks.  
      • The ACC will signal: physical pain, emotional pain, metaphorical pain, anxiety, disgust, embarrassment, social exclusion especially in adolescence; as one and the same.  The ACC's abnormalities being associated with major depression. 
    • Has a bridging role between the empathetic and self-interested pain monitor.  Sapolsky notes the ACC is essential for learning fear and conditioned avoidance by observation alone through an intermediate step of shared representation of self.  He concludes "At its core the ACC is about self-interest, with caring about the other person in pain as an add-on." 
  • American College of Cardiology
's significant role in interpreting the meaning of pain: Physical, Discrepancy identification, Social anxiety, Feeling as someone you love is in pain; with the brain mixing literal and psychic pain -- elevating Substance P is deployed by cells under stress.  It is a powerful vasodilator and initiates deployment of cytokines.  Substance P targets the NK1-receptor.  It is associated with inflammatory processes and pain. 
during clinical depression is a debilitating state which is facilitated by genetic predisposition - for example genes coding for relatively low serotonin levels; and an accumulation of traumatic events.  There is evidence of shifts in the sleep/wake cycle in affected individuals (Dec 2015).  The affected person will experience a pathological sense of loss of control, prolonged sadness, irritability, sleep disturbances, loss of appetite, and inability to experience pleasure.  It affects 12% of men and 20% of women.  It appears to be associated with androgen deprivation therapy treatment for prostate cancer (Apr 2016).  Chronic stress depletes the nucleus accumbens of dopamine, biasing humans towards depression.  Depression easily leads to following unhealthy pathways: drinking, overeating; which increase the risk of heart disease.   It has been associated with an aging related B12 deficiency (Sep 2016).  During depression, stress mediates inhibition of dopamine signalling.  There is an association between depression and particular brain regions: Hippocampal dendrite and spine number reductions, Dorsal raphe nucleus linked to loneliness, Abnormalities of the ACC.  Childhood adversity can increase depression risk by linking recollections of uncontrollable situations to overgeneralizations that life will always be terrible and uncontrollable.  Treatments include: CBT, UMHS depression management.  As of 2010 drug treatments take weeks to facilitate a response & many patients do not respond to the first drug applied, often prolonging the agony.   Genomic predictions of which treatment will be effective have not been possible because: Not all clinical depressions are the same, a standard definition of drug response is difficult; and dread activating cortical components of pain networks.  The ACC activates if a hated rival succeeds while if he fails the dopaminergic reward pathways activate!

Disgust and Purity
The insula is part of the cerebral cortex folded deep within the lateral sulcus.  It includes: anterior, posterior insula; and is overlaid by the operculum.  It is assumed to participate in consciousness where it has been linked to emotion, salience & body homeostasis functions:
  • Perception,
  • Motor control: Hand-&-eye motor movement, Swallowing, Gastric motility, Speech articulation;
  • Self-awareness,
  • Inter-personal experiences: Disgust at smells, contamination & mutilation which generate visceral responses, that are projected to the amygdala; binding physical and moral aspects of purity (Macbeth effect)
  • Homeostatic regulation of the sympathetic network, parasympathetic network, and immune system.  
supports our various feelings of disgust is a universal human emotion.  Pinker notes it has its own facial expression and is codified in food taboos.  The mind must be associated with the proximate environment and parents minimize the risk for their omnivorous children by teaching them what foods to eat and what to avoid.  The children's minds are initially receptive to trying all foods but their brains subsequently lock in on the foods they have experienced.  These parental choices are affected by schematic influence on what has been beneficial.  Adolescent's brain developments undermine these constraints enabling intergroup transfers.  Disgust is modulated by the insula cortex which projects signals to the amygdala. 
: Protecting us from rancid food, Detecting faces showing disgust, Detecting unattractive faces, and thinking about details of appalling events.  Sapolsky stresses the bidirectional link between visceral is our bodily responses to rancid food, evolved to protect against ingesting toxins and infectious organisms.  These responses: wrinkling the nose, raising upper lip, narrowing eyes, slowing of heart, gaging or vomiting out food reflexively; are bound by the architecture of the brain: insula; to other interests of the insula such as faces indicating disgust, unattractive faces, norm violations, moral disgust, and risks to purity.  There are political effects:
  • Social conservatives have a lower threshold for visceral disgust than progressives.
  • A sense of threat is bound into social situations: Sanctity of marriage, Family values; with disgust serving as an ethnic or out-group marker. 
  • Being clean is associated with being good & close to god. 
& moral disgust.  Both send a signal, is an emergent capability which is used by cooperating agents to support coordination & rival agents to support control and dominance.  In eukaryotic cells signalling is used extensively.  A signal interacts with the exposed region of a receptor molecule inducing it to change shape to an activated form.  Chains of enzymes interact with the activated receptor relaying, amplifying and responding to the signal to change the state of the cell.  Many of the signalling pathways pass through the nuclear membrane and interact with the DNA to change its state.  Enzymes sensitive to the changes induced in the DNA then start to operate generating actions including sending further signals.  Cell signalling is reviewed by Helmreich.  Signalling is a fundamental aspect of CAS theory and is discussed from the abstract CAS perspective in signals and sensors.  In AWF the eukaryotic signalling architecture has been abstracted in a codelet based implementation.  To be credible signals must be hard to fake.  To be effective they must be easily detected by the target recipient.  To be efficient they are low cost to produce and destroy. 
to our emotional networks to respond aversively, binding our physical responses to metaphorical ideas.  The body part: mouth, hand; that communicates a lie becomes associated with it and needs to be cleaned (Macbeth effect is the need to clean part of the body used to perform an immoral act.  University of Toronto's Chen-Bo Zhong & Northwestern University's Katie Liljenquist asked subjects to recount a moral or immoral act in their past.  Subsequently the volunteers were offered a gift of a pencil or pack of antiseptic wipes.  Those recounting immoral acts selected the wipes more often.  Physical and moral purity and actions are bound together by the insula.  Another study asked subjects to speak or write a lie.  This time the complementary cleaning products in a study using western subjects were associated with the mouth if they spoke the lie or hands if they wrote it.  If the subjects were East Asian those that wrote the lie selected face cleansers -- Sapolsky comments "If you are going to save face, it should be a clean one.  ).  The feeling of being dirty can induce a balancing prosocial is prioritizing benefiting others through: Help, Charity, Truth; even if many are acts of restitution to balance out antisocial acts.  Due to the Insula's binding of physical and metaphorical disgust, physically washing your hands can be enough to reduce the need for a prosocial act.  
behavior but studies show washing your hands reduces this drive.  And Sapolsky raises concerns about the political implications and problems for Commonsense Morality of the brain's intermixing visceral & moral disgust. 

Real versus Metaphorical Sensation
Yale's John Bargh showed experimentally that evaluation of the seriousness of a person from reading their resume could be influenced by the weight of the clipboard holding it.  Hard or soft chairs influenced the judgement of if someone was hard-hearted or a softie.  Cups of hot or cold drinks influenced the evaluation of someone's personality. 

Sapolsky notes that our brains also confuse metaphorical and literal interoceptive information indicate the body's internal state: Pain, Fatigue; seconds to minutes before.  The interoceptive 'networks' project to brain regions that implement social emotions. 
: judges refused parole far more when they were hungry.  The brain uses circuits that deal with physical properties of objects to assess metaphorical language. 

Duct Tape
Symbols are interpreted as signals, is an emergent capability which is used by cooperating agents to support coordination & rival agents to support control and dominance.  In eukaryotic cells signalling is used extensively.  A signal interacts with the exposed region of a receptor molecule inducing it to change shape to an activated form.  Chains of enzymes interact with the activated receptor relaying, amplifying and responding to the signal to change the state of the cell.  Many of the signalling pathways pass through the nuclear membrane and interact with the DNA to change its state.  Enzymes sensitive to the changes induced in the DNA then start to operate generating actions including sending further signals.  Cell signalling is reviewed by Helmreich.  Signalling is a fundamental aspect of CAS theory and is discussed from the abstract CAS perspective in signals and sensors.  In AWF the eukaryotic signalling architecture has been abstracted in a codelet based implementation.  To be credible signals must be hard to fake.  To be effective they must be easily detected by the target recipient.  To be efficient they are low cost to produce and destroy. 
by various species.  Even in the setting of experiments, for the subjects: rats; the symbol can become rewarding.   Metaphorical symbols have this power and we struggle to remember these metaphors aren't literal.  Sapolsky explains that
This page reviews the implications of selection, variation and heredity in a complex adaptive system (CAS).  The mechanism and its emergence are discussed. 
evolution
must tinker leveraging exaptation, initially termed preadaptation refers to the coopting of some function for a new use.   to build in access to any new
This page discusses the mechanisms and effects of emergence underpinning any complex adaptive system (CAS).  Key research is reviewed. 
emergent
strategy, such as development of the von Economo neurons are also called spindle neurons and are:
  • Found only in: Primates, Whales, Dolphins, Elephants;
  • Present in the: Insula, Anterior cingulate - focused on empathy; integrating and repurposing these basic facilities into high level capabilities such as moral disgust. 
  • First neurons destroyed by FTD. 
to support the insula is part of the cerebral cortex folded deep within the lateral sulcus.  It includes: anterior, posterior insula; and is overlaid by the operculum.  It is assumed to participate in consciousness where it has been linked to emotion, salience & body homeostasis functions:
  • Perception,
  • Motor control: Hand-&-eye motor movement, Swallowing, Gastric motility, Speech articulation;
  • Self-awareness,
  • Inter-personal experiences: Disgust at smells, contamination & mutilation which generate visceral responses, that are projected to the amygdala; binding physical and moral aspects of purity (Macbeth effect)
  • Homeostatic regulation of the sympathetic network, parasympathetic network, and immune system.  
's identification of moral provides rules for identifying right from wrong.  It develops in stages with children using play to work out rules of appropriate behavior.  Kohlberg's 1950s experiments using children led him to conclude moral judgement is a cognitive process that develops in three stages.  Sapolsky raises issues with the framework: Its a model, It does not apply to other cultures, Intuition & emotion are as significant as cognition, Moral reasoning doesn't predict moral actions; and notes the capacity of the frontal cortex to regulate emotions and behavior is far more predictive.  The marshmallow test, performed on three to six year olds, actually predicted their subsequent SAT scores at high school, social success and lack of aggression, and forty years on more PFC activation during a frontal task and a lower BMI!  Jonathan Haidt argues people's moral decisions are rationalizations rather than using reasoning. 
disgust is a universal human emotion.  Pinker notes it has its own facial expression and is codified in food taboos.  The mind must be associated with the proximate environment and parents minimize the risk for their omnivorous children by teaching them what foods to eat and what to avoid.  The children's minds are initially receptive to trying all foods but their brains subsequently lock in on the foods they have experienced.  These parental choices are affected by schematic influence on what has been beneficial.  Adolescent's brain developments undermine these constraints enabling intergroup transfers.  Disgust is modulated by the insula cortex which projects signals to the amygdala. 


The Metaphorical Dark Side
Sapolsky stresses the significance of our confusing metaphorical and literal

A glimmer
Sapolsky explains that metaphor can similarly be used to overcome such hatred.  By understanding the emotional importance of metaphorically derived sacred values and ensuring that these are resolved helps undermine the disgust and anger generated by ignoring them: Northern Ireland, Palestine, South Africa.  Leveraging our confusion of literal and metaphorical can be used to bring about the best of behaviors. 

Biology, the criminal justice system, and (oh why not?) free will
Sapolsky asserts the
Matt Taibbi describes the phenotypic alignment of the American justice system.  The result he explains relentlessly grinds the poor and undocumented into resources to be constrained, consumed and ejected.  Even as it supports and aligns the financial infrastructure into a potent weapon capable of targeting any company or nation to extract profits and leave the victim deflated. 

Taibbi uses five scenarios to provide a broad picture of the: activities, crimes, policing, prosecutions, court processes, prisons and deportation network.  The scenarios are: Undocumented people's neighborhoods, Poor neighborhoods, Welfare recipients, Credit card debtors and Financial institutions.

Following our summary of his arguments, RSS comments on them framed by complex adaptive system (CAS) theory.  The alignment of the justice system reflects a set of long term strategies and responses to a powerful global arms race that the US leadership intends to win. 

current criminal justice system
needs to be abolished and replaced with something with similar features but totally different underpinnings.  He wants a radical alteration that better reflects our known biology.  The current legal framework is based on a very shaky assumption:
Critics of neuroscience being relevant to the legal system such as Stephen Morse, note that "We live in a causal universe, which includes human action."  But Sapolsky sees this as Morse trying to hide a non-causal homunculus.  Morse further complains that neuroscience fails to accurately predict our actions.  But Sapolsky argues neurological predictions about individual behaviors are difficult because these behaviors are all multifactorial.  Sapolsky notes that unlike the realm of physics, neuroscience looks at a different type of causation.  Free will becomes the unknown internal forces that would need to be categorized to fully predict our behaviors.  Sapolsky assumes these unknown areas will continue to shrink and consequently so will free will. 

Sapolsky suggests that the legal system should abandon its dependence on free will.  But he says it should continue to isolate dangerous people.  And punishment is precluded from being an end in itself.  Sapolsky recalls from game theory investigates the optimal strategies for cooperating.  The iterative Prisoner's Dilemma scenario was shown by Axelrod & Hamilton to reward a Tit for Tat strategy with perfect signals.  An effective way to limit the risks of initiating Tit for Tat in a potential sea of Always Defectors is a genetically encoded signal of cooperation such as the green-beard gene.  Other evolved strategies include genetic algorithms. 
that, while tough to do, punishment supports cooperation and evolved sociality.  As a result
This page reviews the implications of selection, variation and heredity in a complex adaptive system (CAS).  The mechanism and its emergence are discussed. 
evolution
has captured punishment being rewarding for the punisher, by activating the vmPFC is ventromedial prefrontal cortex which is:
  • Focused on the impact of emotion on decision making
  • A participant in limbic system operations 
  • Many human behaviors involve interactions between the vmPFC, the limbic system & the dorsolateral prefrontal cortex.  Part of decision making is for the limbic system to internally simulate (often with the help of the sympathetic nervous system) what alternative outcomes of a decision will feel like with the results of these somatic marker analyses being reported to the vmPFC.  
  • Damage to the vmPFC results in bad decision making: Poor judgement in choosing friends & partners, Failure to respond to negative feedback; because they can't feel the issues. 
, amygdala contains > 12 distinct areas: Central, Lateral.  It receives simple signals from the lower parts of the brain: pain from the PAG; and abstract complex information from the highest areas: Disgust from the insula cortex.  It sends signals to almost every other part of the brain, including to the decision-making circuitry of the frontal lobes.  It has high levels of D(1) dopamine receptors.  During extreme fear the amygdala drives the hippocampus into fear learning.  It outputs directly to subcortical reflexive motor pathways when speed is required.  Its central nucleus projects to the BNST.  It signals the locus ceruleus.  The amygdala:
  • Promotes aggression.  Stimulating the amygdala promotes rage.  It converts anger into aggression and when impaired it impacts the ability to detect angry facial expressions.  
  • Participates in disgust
  • Perceives fear promoting stimuli.  In PTSD sufferers the Amygdala overreacts to mildly fearful stimuli and is slow to calm down and the amygdala expands in size over a period of months.  Fear is processed by the lateral nucleus which serves as the input from various senses, and the central nucleus which outputs to the brain stem (central grey - freezing, lateral hypothalamus - blood pressure, activates paraventricular hypothalamus => crf -> hormone adjustments). 
  • Has lots of receptors for and is highly sensitive to glucocorticoids.  Stress inhibits the GABA interneurons in the basolateral amygdala (BLA) allowing the excitatory glutamate releasing neurons to excite more. 
  • Is sensitive to unsettling/uncertain social situations where it promotes anxiety.  It is also interested in uncertain but potentially painful situations.  The amygdala contributes to social and emotional decision making where the BLA supports rejecting an unacceptable offer, as allowed in the Ultimatum Game, by injecting implicit mistrust and vigilance, generating an anger driven rejection that is used as punishment.  The amygdala is very rapidly excited by subliminal signals from the thalamus of outgroup skin color.  The amygdala subsequently tips social emotions against outgroups unless restrained by the frontal lobe or influenced by subliminal priming to prioritize inclusion.  The fast path from the thalamus rapidly but inaccurately signals its identified a weapon. 
  • Promotes male, but not female, sexual motivation when it is an uncertain potential pleasure. 
  • Responds to the longing for uncertain potential pleasures and fear that the reward will not be worth it if it happens.  The amygdala turns off during orgasm. 
  • Uses but is not directly involved in vision. 
& insula is part of the cerebral cortex folded deep within the lateral sulcus.  It includes: anterior, posterior insula; and is overlaid by the operculum.  It is assumed to participate in consciousness where it has been linked to emotion, salience & body homeostasis functions:
  • Perception,
  • Motor control: Hand-&-eye motor movement, Swallowing, Gastric motility, Speech articulation;
  • Self-awareness,
  • Inter-personal experiences: Disgust at smells, contamination & mutilation which generate visceral responses, that are projected to the amygdala; binding physical and moral aspects of purity (Macbeth effect)
  • Homeostatic regulation of the sympathetic network, parasympathetic network, and immune system.  
.  Sapolsky argues that this 'punishment as virtue' is unacceptable but will be hard to curtail.  He hopes that society will eventually remove labels like evil from terrible crimes that we can see are due to particular unfortunate deployments of neurons, specialized eukaryotic cells include channels which control flows of sodium and potassium ions across the massively extended cell membrane supporting an electro-chemical wave which is then converted into an outgoing chemical signal transmission from synapses which target nearby neuron or muscle cell receptors.  Neurons are supported by glial cells.  Neurons include a:
  • Receptive element - dendrites
  • Transmitting element - axon and synaptic terminals
  • Highly variable DNA schema using transposons. 


Similarly he warns that no free will implies that our best acts also occur because we are
Plans are interpreted and implemented by agents.  This page discusses the properties of agents in a complex adaptive system (CAS). 
It then presents examples of agents in different CAS.  The examples include a computer program where modeling and actions are performed by software agents.  These software agents are aggregates. 
The participation of agents in flows is introduced and some implications of this are outlined. 
agents
supported by a culture is how we do and think about things, transmitted by non-genetic means as defined by Frans de Waal.  CAS theory views cultures as operating via memetic schemata evolved by memetic operators to support a cultural superorganism.  Evolutionary psychology asserts that human culture reflects adaptations generated while hunting and gathering.  Dehaene views culture as essentially human, shaped by exaptations and reading, transmitted with support of the neuronal workspace and stabilized by neuronal recycling.  Sapolsky argues that parents must show children how to transform their genetically derived capabilities into a culturally effective toolset.  He is interested in the broad differences across cultures of: Life expectancy, GDP, Death in childbirth, Violence, Chronic bullying, Gender equality, Happiness, Response to cheating, Individualist or collectivist, Enforcing honor, Approach to hierarchy; illustrating how different a person's life will be depending on the culture where they are raised.  Culture:
  • Is deployed during pregnancy & childhood, with parental mediation.  Nutrients, immune messages and hormones all affect the prenatal brain.  Hormones: Testosterone with anti-Mullerian hormone masculinizes the brain by entering target cells and after conversion to estrogen binding to intracellular estrogen receptors; have organizational effects producing lifelong changes.  Parenting style typically produces adults who adopt the same approach.  And mothering style can alter gene regulation in the fetus in ways that transfer epigenetically to future generations!  PMS symptoms vary by culture. 
  • Is also significantly transmitted to children by their peers during play.  So parents try to control their children's peer group.  
  • Is transmitted to children by their neighborhoods, tribes, nations etc. 
  • Influences the parenting style that is considered appropriate. 
  • Can transform dominance into honor.  There are ecological correlates of adopting honor cultures.  Parents in honor cultures are typically authoritarian. 
  • Is strongly adapted across a meta-ethnic frontier according to Turchin.  
  • Across Europe was shaped by the Carolingian empire. 
  • Can provide varying levels of support for innovation.  
  • Produces consciousness according to Dennet. 
of external environmental signals. 

War and Peace
In the final chapter Sapolsky aims to show that our best behaviors are winning out, and often in unlikely circumstances.  He notes progress over 200 years:
  • Every nation has outlawed slavery and most attempt to enforce it. 
  • Most nations have child labor laws and rates of child labor have declined. 
  • Most countries regulate the treatment of animals. 
  • The world is safer with far fewer homicides than in the past. 
  • In the last century there have been: bans on using some weapons, development of a World Court, punishment for crimes against humanity. 
Framing the improvements with Steven Pinker's "The Better Angels of Our Nature: Why Violence Has Declined" he asks why were people so awful in the past.  Pinker argues it is because people had historically been awful.  But citing the disagreement of experts in response to Pinker's comments about war, this claim is also in doubt.  Pinker suggests a 'civilizing process is sociologist Norbert Elias's notion that violence declines when states monopolize force. 
' encouraging commerce and trade, which supports partnering, an 'escalator of reasoning' and a 'rights revolution' helped by the Flynn effect is an increase in average IQ over the last century.  It is well-documented. 
, have together driven our recent improvement.  Again critics doubt Pinker's dependence on improved cognition is the ability to orchestrate thought and action in accordance with internal goals according to Princeton's Jonathan Cohen. 
.  Sapolsky, citing the prior chapters of Behave, argues there must be interaction between reason is dorsolateral prefrontal cortex which is:
  • At the heart of decision making - highly rational, unsentimental 
  • A major agent of working memory
  • The most recently evolved part of the prefrontal cortex.  
  • Mainly interconnected with other parts of the cortex.  
and feeling are models including ratings of situations which are evolutionarily associated with emotions encoded in neural circuits: amygdala, and insula.  

Careful analysis of the data on violence across centuries finds Pinker's conclusions to be questionable.  Violence now happens fast with broad network effects.  World War 2 and World War 1 are found to be key events along with the Rwandan genocide.  Sapolsky concludes that if things have improved they still aren't good. 

Sapolsky looks for ways the insights from Behave might help: