Psychopathy to Altruism: Neurobiology of the Selfish–Selfless Spectrum

The age-old philosophical, biological, and social debate over the basic nature of humans as being “universally selfish” or “universally good” continues today highlighting sharply divergent views of natural social order. Here we analyze advances in biology, genetics and neuroscience increasing our understanding of the evolution, features and neurocircuitry of the human brain underlying behavior in the selfish–selfless spectrum. First, we examine evolutionary pressures for selection of altruistic traits in species with protracted periods of dependence on parents and communities for subsistence and acquisition of learned behaviors. Evidence supporting the concept that altruistic potential is a common feature in human populations is developed. To go into greater depth in assessing critical features of the social brain, the two extremes of selfish–selfless behavior, callous unemotional psychopaths and zealous altruists who take extreme measures to help others, are compared on behavioral traits, structural/functional neural features, and the relative contributions of genetic inheritance versus acquired cognitive learning to their mindsets. Evidence from population groups ranging from newborns, adopted children, incarcerated juveniles, twins and mindfulness meditators point to the important role of neuroplasticity and the dopaminergic reward systems in forming and reforming neural circuitry in response to personal experience and cultural influences in determining behavior in the selfish–selfless spectrum. The underlying neural circuitry differs between psychopaths and altruists with emotional processing being profoundly muted in psychopaths and significantly enhanced in altruists. But both groups are characterized by the reward system of the brain shaping behavior. Instead of rigid assignment of human nature as being “universally selfish” or “universally good,” both characterizations are partial truths based on the segments of the selfish–selfless spectrum being examined. In addition, individuals and populations can shift in the behavioral spectrum in response to cognitive therapy and social and cultural experience, and approaches such as mindfulness training for introspection and reward-activating compassion are entering the mainstream of clinical care for managing pain, depression, and stress.

[1]  T. Ono,et al.  Selective Medial Prefrontal Cortex Responses During Live Mutual Gaze Interactions in Human Infants: An fNIRS Study , 2015, Brain Topography.

[2]  A. Toga,et al.  Morphological alterations in the prefrontal cortex and the amygdala in unsuccessful psychopaths. , 2010, Journal of abnormal psychology.

[3]  Jean Clobert,et al.  Self-recognition, color signals, and cycles of greenbeard mutualism and altruism. , 2006, Proceedings of the National Academy of Sciences of the United States of America.

[4]  Barbara Tomasino,et al.  Disentangling the neural mechanisms involved in Hinduism- and Buddhism-related meditations , 2014, Brain and Cognition.

[5]  Matthew L. Dixon,et al.  Functional neuroanatomy of meditation: A review and meta-analysis of 78 functional neuroimaging investigations , 2016, Neuroscience & Biobehavioral Reviews.

[6]  J. Fletcher,et al.  TOWARDS A GENERAL THEORY OF GROUP SELECTION , 2013, Evolution; international journal of organic evolution.

[7]  Karen Wynn,et al.  Young infants prefer prosocial to antisocial others. , 2011, Cognitive development.

[8]  Robert D. Hare,et al.  Psychopathy Checklist—Revised , 2011 .

[9]  Christopher J Patrick,et al.  Psychopathic personality traits: heritability and genetic overlap with internalizing and externalizing psychopathology , 2005, Psychological Medicine.

[10]  Rachel E. Kahn,et al.  Callous-unemotional traits robustly predict future criminal offending in young men. , 2013, Law and human behavior.

[11]  A. Loewy,et al.  Central Command Neurons of the Sympathetic Nervous System: Basis of the Fight-or-Flight Response , 1995, Science.

[12]  P. Petrovic,et al.  Reliability and Construct Validity of the Psychopathic Personality Inventory-Revised in a Swedish Non-Criminal Sample – A Multimethod Approach including Psychophysiological Correlates of Empathy for Pain , 2016, PloS one.

[13]  Risto Näätänen,et al.  Learning-induced neural plasticity of speech processing before birth , 2013, Proceedings of the National Academy of Sciences.

[14]  U. Bellugi,et al.  Oxytocin and Vasopressin Are Dysregulated in Williams Syndrome, a Genetic Disorder Affecting Social Behavior , 2012, PloS one.

[15]  Jason P. Mitchell,et al.  A neural mechanism of first impressions , 2009, Nature Neuroscience.

[16]  A. D. Higginson,et al.  Altruism in a volatile world , 2018, Nature.

[17]  Richard McElreath,et al.  Culture rather than genes provides greater scope for the evolution of large-scale human prosociality , 2009, Proceedings of the National Academy of Sciences.

[18]  R. Ebstein,et al.  Link Between Vasopressin Receptor AVPR1A Promoter Region Microsatellites and Measures of Social Behavior in Humans , 2005 .

[19]  P. Dechent,et al.  Neural Correlates of Empathy with Pain Show Habituation Effects. An fMRI Study , 2015, PloS one.

[20]  G. Arbanas Diagnostic and Statistical Manual of Mental Disorders (DSM-5) , 2015 .

[21]  William G. Boissonnault Chapter 7 – Review of Systems , 2005 .

[22]  Jennifer L. Skeem,et al.  Psychopathic Personality , 2011, Psychological science in the public interest : a journal of the American Psychological Society.

[23]  Nancy Eisenberg,et al.  The roots of prosocial behavior in children , 1989 .

[24]  Björn N. Persson,et al.  Empathy and universal values explicated by the empathy-altruism hypothesis , 2016, The Journal of social psychology.

[25]  Pascal Belin,et al.  Mother and stranger: an electrophysiological study of voice processing in newborns. , 2011, Cerebral cortex.

[26]  Stephen J. Guter,et al.  Convergence of Genes and Cellular Pathways Dysregulated in Autism Spectrum Disorders , 2014, American journal of human genetics.

[27]  W. Hamilton,et al.  The evolution of cooperation. , 1984, Science.

[28]  E. Kimonis,et al.  Does the self-report Inventory of Callous-Unemotional Traits predict recidivism? , 2016, Psychological assessment.

[29]  Uri Bergmann,et al.  Stochastic resonance as a proposed neurobiological model for Eye Movement Desensitization and Reprocessing (EMDR) therapy. , 2018, Medical hypotheses.

[30]  K. Kiehl,et al.  THE CRIMINAL PSYCHOPATH: HISTORY, NEUROSCIENCE, TREATMENT, AND ECONOMICS. , 2011, Jurimetrics.

[31]  F. Moeller,et al.  Effects of Intranasal Oxytocin on Aggressive Responding in Antisocial Personality Disorder , 2015, The Psychological record.

[32]  Robin I. M. Dunbar,et al.  Kinship and altruism: a cross-cultural experimental study. , 2007, British journal of psychology.

[33]  D. L. Robinson,et al.  Anatomical and pharmacological characterization of catecholamine transients in the medial prefrontal cortex evoked by ventral tegmental area stimulation , 2014, Synapse.

[34]  P. Morgane,et al.  A review of systems and networks of the limbic forebrain/limbic midbrain , 2005, Progress in Neurobiology.

[35]  Ernst Fehr,et al.  Advances in Economics and Econometrics: Theories of Fairness and Reciprocity: Evidence and Economic Applications , 2003 .

[36]  Paul J. Whalen,et al.  The uncertainty of it all , 2007, Trends in Cognitive Sciences.

[37]  S. Sands,et al.  Evidence for alterations in stimulatory G proteins and oxytocin levels in children with autism , 2014, Psychoneuroendocrinology.

[38]  Elinor McKone,et al.  Not just fear and sadness: Meta-analytic evidence of pervasive emotion recognition deficits for facial and vocal expressions in psychopathy , 2012, Neuroscience & Biobehavioral Reviews.

[39]  Margaret A. Sheridan,et al.  Childhood adversity and neural development: Deprivation and threat as distinct dimensions of early experience , 2014, Neuroscience & Biobehavioral Reviews.

[40]  J. VanMeter,et al.  Mediation of the relationship between callous-unemotional traits and proactive aggression by amygdala response to fear among children with conduct problems. , 2014, JAMA psychiatry.

[41]  J. O'Doherty,et al.  Automatic and intentional brain responses during evaluation of trustworthiness of faces , 2002, Nature Neuroscience.

[42]  T. Brugha,et al.  Psychopathy among prisoners in England and Wales. , 2009, International journal of law and psychiatry.

[43]  S. Preston,et al.  Mammalian empathy: behavioural manifestations and neural basis , 2017, Nature Reviews Neuroscience.

[44]  T. Insel,et al.  Facilitation of Affiliation and Pair-Bond Formation by Vasopressin Receptor Gene Transfer into the Ventral Forebrain of a Monogamous Vole , 2001, The Journal of Neuroscience.

[45]  Jan J. V. Busschbach,et al.  Standardised Mindfulness-Based Interventions in Healthcare: An Overview of Systematic Reviews and Meta-Analyses of RCTs , 2015, PloS one.

[46]  Geraint Rees,et al.  Political Orientations Are Correlated with Brain Structure in Young Adults , 2011, Current Biology.

[47]  Larry J Young,et al.  Profound Impairment in Social Recognition and Reduction in Anxiety-Like Behavior in Vasopressin V1a Receptor Knockout Mice , 2004, Neuropsychopharmacology.

[48]  Claus Lamm,et al.  The Neural Substrate of Human Empathy: Effects of Perspective-taking and Cognitive Appraisal , 2007, Journal of Cognitive Neuroscience.

[49]  K. Kiehl,et al.  Psychopathy: developmental perspectives and their implications for treatment. , 2014, Restorative neurology and neuroscience.

[50]  Richard J. Davidson,et al.  Altered anterior insula activation during anticipation and experience of painful stimuli in expert meditators , 2013, NeuroImage.

[51]  M. Iacoboni,et al.  Self‐other resonance, its control and prosocial inclinations: Brain–behavior relationships , 2016, Human brain mapping.

[52]  Claus Lamm,et al.  Meta-analytic evidence for common and distinct neural networks associated with directly experienced pain and empathy for pain , 2011, NeuroImage.

[53]  E. Viding,et al.  Genetic and environmental aetiology of the dimensions of Callous-Unemotional traits , 2015, Psychological Medicine.

[54]  R. Dolan,et al.  Impaired threat prioritisation after selective bilateral amygdala lesions , 2015, Cortex.

[55]  S. Rombouts,et al.  Anterior cingulate cortex grey matter volume abnormalities in adolescents with PTSD after childhood sexual abuse , 2017, European Neuropsychopharmacology.

[56]  Jack C. Rogers,et al.  Cortical and Subcortical Gray Matter Volume in Youths With Conduct Problems: A Meta-analysis. , 2016, JAMA psychiatry.

[57]  L. Young,et al.  Can oxytocin treat autism? , 2015, Science.

[58]  Jorge A. Barraza,et al.  The neurobiology of collective action , 2013, Front. Neurosci..

[59]  R. Hare Psychopathy: a clinical and forensic overview. , 2006, The Psychiatric clinics of North America.

[60]  M. Caldwell,et al.  Reducing violence in serious juvenile offenders using intensive treatment. , 2005, International journal of law and psychiatry.

[61]  M. Olff,et al.  A five-day inpatient EMDR treatment programme for PTSD: pilot study , 2018, European journal of psychotraumatology.

[62]  K. Kiehl,et al.  Neural correlates of reward and loss sensitivity in psychopathy. , 2014, Social cognitive and affective neuroscience.

[63]  M. Posner,et al.  Cognitive and emotional influences in anterior cingulate cortex , 2000, Trends in Cognitive Sciences.

[64]  David R. W. Bachhuber,et al.  Reduced stress and inflammatory responsiveness in experienced meditators compared to a matched healthy control group , 2016, Psychoneuroendocrinology.

[65]  A. Böckler,et al.  Decoding the Charitable Brain: Empathy, Perspective Taking, and Attention Shifts Differentially Predict Altruistic Giving , 2016, The Journal of Neuroscience.

[66]  R. Halleck The cultivation of the will. , 1895 .

[67]  I. Waldman,et al.  The oxytocin receptor gene (OXTR) is associated with autism spectrum disorder: a meta-analysis , 2014, Molecular Psychiatry.

[68]  Ernst Fehr,et al.  Linking Brain Structure and Activation in Temporoparietal Junction to Explain the Neurobiology of Human Altruism , 2012, Neuron.

[69]  V. Calhoun,et al.  Abnormal brain structure in youth who commit homicide , 2014, NeuroImage: Clinical.

[70]  T. Singer,et al.  Short-Term Compassion Training Increases Prosocial Behavior in a Newly Developed Prosocial Game , 2011, PloS one.

[71]  Justin S. Feinstein,et al.  Oxytocin-enforced norm compliance reduces xenophobic outgroup rejection , 2017, Proceedings of the National Academy of Sciences.

[72]  R. Murray,et al.  The impact of childhood adversity on the persistence of psychotic symptoms: a systematic review and meta-analysis , 2015, Psychological Medicine.

[73]  R. Adolphs,et al.  The social brain: neural basis of social knowledge. , 2009, Annual review of psychology.

[74]  Zoe R. Donaldson,et al.  Oxytocin, Vasopressin, and the Neurogenetics of Sociality , 2008, Science.

[75]  Shelley E. Taylor,et al.  Culture, distress, and oxytocin receptor polymorphism (OXTR) interact to influence emotional support seeking , 2010, Proceedings of the National Academy of Sciences.

[76]  J. Bachevalier,et al.  Pervasive alterations of emotional and neuroendocrine responses to an acute stressor after neonatal amygdala lesions in rhesus monkeys , 2013, Psychoneuroendocrinology.

[77]  Joachim G. Frommen,et al.  The evolution of cooperation based on direct fitness benefits , 2016, Philosophical Transactions of the Royal Society B: Biological Sciences.

[78]  Steve Stewart-Williams Human Beings as Evolved Nepotists , 2008, Human nature.

[79]  Michael Davis,et al.  Cortical Afferents to the Extended Amygdala , 1999, Annals of the New York Academy of Sciences.

[80]  Garet P Lahvis Social Reward and Empathy as Proximal Contributions to Altruism: The Camaraderie Effect. , 2016, Current topics in behavioral neurosciences.

[81]  Don M. Gash,et al.  Neuron-based heredity and human evolution , 2015, Front. Neurosci..

[82]  M. Fischer-Shofty,et al.  Intranasal administration of oxytocin increases human aggressive behavior , 2016, Hormones and Behavior.

[83]  B. Herpertz-Dahlmann,et al.  Reduced Cortisol in Boys with Early-Onset Conduct Disorder and Callous-Unemotional Traits , 2013, BioMed research international.

[84]  Preliminary Investigation of a Mindfulness-Based Intervention for Social Anxiety Disorder That Integrates Compassion Meditation and Mindful Exposure. , 2016, Journal of alternative and complementary medicine.

[85]  R. Riemann,et al.  Left or right? Sources of political orientation: the roles of genetic factors, cultural transmission, assortative mating, and personality. , 2012, Journal of personality and social psychology.

[86]  C. Dweck,et al.  Rethinking natural altruism: Simple reciprocal interactions trigger children’s benevolence , 2014, Proceedings of the National Academy of Sciences.

[87]  Clayton A. Robarchek,et al.  Cultures of War and Peace : A Comparative Study of Waorani and Semai , 2006 .

[88]  A. Grace,et al.  Cortico-Basal Ganglia Reward Network: Microcircuitry , 2010, Neuropsychopharmacology.

[89]  Todd A. Hare,et al.  Early developmental emergence of human amygdala–prefrontal connectivity after maternal deprivation , 2013, Proceedings of the National Academy of Sciences.

[90]  D. Reiss,et al.  Heritable and Nonheritable Pathways to Early Callous-Unemotional Behaviors. , 2016, The American journal of psychiatry.

[91]  T. Johnstone,et al.  Regulation of the Neural Circuitry of Emotion by Compassion Meditation: Effects of Meditative Expertise , 2008, PloS one.

[92]  J. Kennedy,et al.  Childhood aggression, callous-unemotional traits and oxytocin genes , 2011, European Child & Adolescent Psychiatry.

[93]  Tobias Grossmann,et al.  The role of medial prefrontal cortex in early social cognition , 2013, Front. Hum. Neurosci..

[94]  James H Fowler,et al.  Altruistic Punishment and the Origin of Cooperation , 2005, Proceedings of the National Academy of Sciences of the United States of America.

[95]  D. Shohamy,et al.  Intrinsic connectivity between the hippocampus, nucleus accumbens, and ventral tegmental area in humans , 2013, Hippocampus.

[96]  J. Hamlin,et al.  The early emergence of sociomoral evaluation: infants prefer prosocial others. , 2018, Current opinion in psychology.

[97]  N. Hadjikhani,et al.  Seeing Fearful Body Expressions Activates the Fusiform Cortex and Amygdala , 2003, Current Biology.

[98]  D. Balliet,et al.  Trust, Punishment, and Cooperation Across 18 Societies , 2013, Perspectives on psychological science : a journal of the Association for Psychological Science.

[99]  C. S. Carter,et al.  Oxytocin pathways and the evolution of human behavior. , 2014, Annual review of psychology.

[100]  Stephanie L. Brown,et al.  Connecting prosocial behavior to improved physical health: Contributions from the neurobiology of parenting , 2015, Neuroscience & Biobehavioral Reviews.

[101]  F. Sweep,et al.  Voluntary activation of the sympathetic nervous system and attenuation of the innate immune response in humans , 2014, Proceedings of the National Academy of Sciences.

[102]  Nancy L Pedersen,et al.  Genetic variation in the vasopressin receptor 1a gene (AVPR1A) associates with pair-bonding behavior in humans , 2008, Proceedings of the National Academy of Sciences.

[103]  J. Craig,et al.  The motivations and experiences of living kidney donors: a thematic synthesis. , 2012, American journal of kidney diseases : the official journal of the National Kidney Foundation.

[104]  C. Nelson,et al.  Early experience and brain development. , 2017, Wiley interdisciplinary reviews. Cognitive science.

[105]  E. Tucker-Drob,et al.  Hair and Salivary Testosterone, Hair Cortisol, and Externalizing Behaviors in Adolescents , 2018, Psychological science.

[106]  C. Daniel Batson,et al.  Is Empathic Emotion a Source of Altruistic Motivation , 1981 .

[107]  P. Hatemi,et al.  The genetics of politics: discovery, challenges, and progress. , 2012, Trends in genetics : TIG.

[108]  P. Pye-Smith The Descent of Man, and Selection in Relation to Sex , 1871, Nature.

[109]  Lisa Feldman Barrett,et al.  Amygdala and fusiform gyrus temporal dynamics: Responses to negative facial expressions , 2008, BMC Neuroscience.

[110]  Jean Decety,et al.  Impulsive-antisocial dimension of psychopathy linked to enlargement and abnormal functional connectivity of the striatum. , 2017, Biological psychiatry. Cognitive neuroscience and neuroimaging.

[111]  Tania Singer,et al.  Differential pattern of functional brain plasticity after compassion and empathy training. , 2014, Social cognitive and affective neuroscience.

[112]  A. Christophe,et al.  Newborns' Cry Melody Is Shaped by Their Native Language , 2009, Current Biology.

[113]  L. Negi,et al.  Effect of compassion meditation on neuroendocrine, innate immune and behavioral responses to psychosocial stress , 2009, Psychoneuroendocrinology.

[114]  M. Caldwell,et al.  Treatment-Related Changes in Behavioral Outcomes of Psychopathy Facets in Adolescent Offenders , 2011, Law and human behavior.

[115]  Ernst Fehr,et al.  Theories of Fairness and Reciprocity , 2001 .

[116]  James H Fowler,et al.  Friendships Moderate an Association Between a Dopamine Gene Variant and Political Ideology , 2008, The journal of politics.

[117]  M. Raichle,et al.  The anterior cingulate cortex mediates processing selection in the Stroop attentional conflict paradigm. , 1990, Proceedings of the National Academy of Sciences of the United States of America.

[118]  G. Wilkinson,et al.  Non-kin cooperation in bats , 2016, Philosophical Transactions of the Royal Society B: Biological Sciences.

[119]  P. Whalen Fear, Vigilance, and Ambiguity , 1998 .

[120]  M. A. Preis,et al.  Empathy for pain: The effects of prior experience and sex , 2012, European journal of pain.

[121]  StephaniC. Wang,et al.  Nonsocial Functions of Hypothalamic Oxytocin , 2013, ISRN neuroscience.

[122]  K. Kendrick,et al.  Oxytocin Modulates Social Distance between Males and Females , 2012, The Journal of Neuroscience.

[123]  S. Preston,et al.  The origins of altruism in offspring care. , 2013, Psychological bulletin.

[124]  M. Jung,et al.  Differential regulation of observational fear and neural oscillations by serotonin and dopamine in the mouse anterior cingulate cortex , 2014, Psychopharmacology.

[125]  Peter Kirsch,et al.  Beyond revenge: Neural and genetic bases of altruistic punishment , 2011, NeuroImage.

[126]  U. Fischbacher,et al.  Oxytocin increases trust in humans , 2005, Nature.

[127]  T. Moffitt,et al.  Predictors and outcomes of joint trajectories of callous-unemotional traits and conduct problems in childhood. , 2011, Journal of abnormal psychology.

[128]  W. McArdle,et al.  Environmental risk, Oxytocin Receptor Gene (OXTR) methylation and youth callous-unemotional traits: A 13-year longitudinal study , 2014, Molecular Psychiatry.

[129]  Mriganka Sur,et al.  Genes, circuits, and precision therapies for autism and related neurodevelopmental disorders , 2015, Science.

[130]  E. Leibenluft,et al.  The Inventory of Callous-Unemotional Traits (ICU) in Children: Reliability and Heritability , 2016, Behavior Genetics.

[131]  C. Daniel Batson,et al.  More evidence that empathy is a source of altruistic motivation. , 1982 .

[132]  J. Harlow Recovery from the passage of an iron bar through the head , 1993 .

[133]  César F. Lima,et al.  Reduced Laughter Contagion in Boys at Risk for Psychopathy , 2017, Current Biology.

[134]  A. Raine,et al.  Successful and Unsuccessful Psychopathy , 2020 .

[135]  Dustin A. Pardini,et al.  Lower Amygdala Volume in Men is Associated with Childhood Aggression, Early Psychopathic Traits, and Future Violence , 2014, Biological Psychiatry.

[136]  J. Herman Neural pathways of stress integration: relevance to alcohol abuse. , 2012 .

[137]  S. Matusall Social behavior in the “Age of Empathy”?—A social scientist's perspective on current trends in the behavioral sciences , 2013, Front. Hum. Neurosci..

[138]  Hanna Damasio,et al.  Impairment of social and moral behavior related to early damage in human prefrontal cortex , 1999, Nature Neuroscience.

[139]  F. Zufall,et al.  Social motivation is reduced in vasopressin 1b receptor null mice despite normal performance in an olfactory discrimination task , 2004, Hormones and Behavior.

[140]  Wen Zhou,et al.  Oxytocin modulates the racial bias in neural responses to others’ suffering , 2013, Biological Psychology.

[141]  M. Dadds,et al.  Polymorphisms in the oxytocin receptor gene are associated with the development of psychopathy , 2013, Development and Psychopathology.

[142]  Evangelos Kranakis Towards a General Theory , 1986 .

[143]  J. W. Whiting,et al.  Children of Six Cultures: A Psycho-Cultural Analysis , 1975 .

[144]  Anneke Buffone,et al.  The Neurogenetics of Nice , 2012, Psychological science.

[145]  R. Lewontin ‘The Selfish Gene’ , 1977, Nature.

[146]  John L. Bradshaw,et al.  Shared pain: From empathy to synaesthesia , 2010, Neuroscience & Biobehavioral Reviews.

[147]  Bradford C. Dickerson,et al.  Amygdala Volume and Social Network Size in Humans , 2010, Nature Neuroscience.

[148]  B. Quinn,et al.  Prolonged institutional rearing is associated with atypically large amygdala volume and difficulties in emotion regulation. , 2010, Developmental science.

[149]  Karen O'Quin,et al.  Influence of self-reported distress and empathy on egoistic versus altruistic motivation to help. , 1983 .

[150]  P. Schyns,et al.  A mechanism for impaired fear recognition after amygdala damage , 2005, Nature.

[151]  V. Calhoun,et al.  Differentiating emotional processing and attention in psychopathy with functional neuroimaging , 2017, Cognitive, affective & behavioral neuroscience.

[152]  Peter Kirsch,et al.  Oxytocin Modulates Neural Circuitry for Social Cognition and Fear in Humans , 2005, The Journal of Neuroscience.

[153]  Reginald B. Adams,et al.  Voting behavior is reflected in amygdala response across cultures. , 2010, Social cognitive and affective neuroscience.

[154]  Christina M. Karns,et al.  The Cultivation of Pure Altruism via Gratitude: A Functional MRI Study of Change with Gratitude Practice , 2017, Front. Hum. Neurosci..

[155]  J. Field,et al.  Cooperation between non-relatives in a primitively eusocial paper wasp, Polistes dominula , 2016, Philosophical Transactions of the Royal Society B: Biological Sciences.

[156]  Hanna Damasio,et al.  Neural correlates of gratitude , 2015, Front. Psychol..

[157]  W. Malarkey,et al.  Marital behavior, oxytocin, vasopressin, and wound healing , 2010, Psychoneuroendocrinology.

[158]  Joanne P. Smith-Darden,et al.  The Development of Severe and Chronic Violence Among Youth: The Role of Psychopathic Traits and Reward Processing , 2017, Child psychiatry and human development.

[159]  Y. Bloch,et al.  The reliability, concurrent validity and association with salivary oxytocin of the self-report version of the Inventory of Callous-Unemotional Traits in adolescents with conduct disorder , 2017, Psychiatry Research.

[160]  M. Dadds,et al.  Nasal Oxytocin for Social Deficits in Childhood Autism: A Randomized Controlled Trial , 2013, Journal of Autism and Developmental Disorders.

[161]  Scott A. Huettel,et al.  Differential Reward Learning for Self and Others Predicts Self-Reported Altruism , 2014, PloS one.

[162]  U. Bellugi,et al.  The social phenotype of Williams syndrome , 2013, Current Opinion in Neurobiology.

[163]  D. Katz,et al.  Learning-Related Plasticity of Temporal Coding in Simultaneously Recorded Amygdala–Cortical Ensembles , 2008, The Journal of Neuroscience.

[164]  A. Endress,et al.  The Social Sense: Susceptibility to Others’ Beliefs in Human Infants and Adults , 2010, Science.

[165]  D. Hinton,et al.  Loving-kindness and compassion meditation: potential for psychological interventions. , 2011, Clinical psychology review.

[166]  Eric M. Hansen,et al.  "I've also experienced loss and fear": Effects of prior similar experience on empathy. , 2009, Scandinavian journal of psychology.

[167]  K. Fuxe,et al.  Role of dopamine receptor mechanisms in the amygdaloid modulation of fear and anxiety: Structural and functional analysis , 2010, Progress in Neurobiology.

[168]  Ale Smidts,et al.  The Herding Hormone , 2012, Psychological science.

[169]  Giuseppe Pagnoni,et al.  Effects of intranasal oxytocin and vasopressin on cooperative behavior and associated brain activity in men , 2012, Psychoneuroendocrinology.

[170]  John W. VanMeter,et al.  Neural and cognitive characteristics of extraordinary altruists , 2014, Proceedings of the National Academy of Sciences.

[171]  A. Damasio,et al.  The return of Phineas Gage: clues about the brain from the skull of a famous patient. , 1994, Science.

[172]  F. Warneken,et al.  The shadow of the future: 5-year-olds, but not 3-year-olds, adjust their sharing in anticipation of reciprocation. , 2015, Journal of experimental child psychology.

[173]  Phan Luu,et al.  The Anterior Cingulate Cortex : Regulating Actions in Context , 2004 .

[174]  R. Adolphs,et al.  Impaired recognition of emotion in facial expressions following bilateral damage to the human amygdala , 1994, Nature.

[175]  R. Herringa,et al.  Paradoxical Prefrontal–Amygdala Recruitment to Angry and Happy Expressions in Pediatric Posttraumatic Stress Disorder , 2016, Neuropsychopharmacology.

[176]  A. Gardner,et al.  The genetical theory of multilevel selection , 2015, Journal of evolutionary biology.

[177]  Simone Ullrich,et al.  Prevalence and correlates of psychopathic traits in the household population of Great Britain. , 2009, International journal of law and psychiatry.

[178]  D. Picard,et al.  Infants' preference for prosocial behaviors: A literature review. , 2016, Infant behavior & development.

[179]  P. Chauvel,et al.  Reversible antisocial behavior in ventromedial prefrontal lobe epilepsy , 2013, Epilepsy & Behavior.

[180]  C. Heyes,et al.  The cultural evolution of mind reading , 2014, Science.

[181]  Danielle S. Bassett,et al.  A validated network of effective amygdala connectivity , 2007, NeuroImage.

[182]  Paul Babiak,et al.  Snakes in Suits: When Psychopaths Go to Work , 2015 .

[183]  A. Raine,et al.  Successful and unsuccessful psychopaths: a neurobiological model. , 2010, Behavioral sciences & the law.

[184]  D. Tranel,et al.  Acquired Personality Disturbances Associated With Bilateral Damage to the Ventromedial Prefrontal Region , 2000, Developmental neuropsychology.

[185]  A. Lo,et al.  Group Selection as Behavioral Adaptation to Systematic Risk , 2014, PloS one.

[186]  David H. Skuse,et al.  Dopaminergic-neuropeptide interactions in the social brain , 2009, Trends in Cognitive Sciences.

[187]  A. Simmons,et al.  Red Brain, Blue Brain: Evaluative Processes Differ in Democrats and Republicans , 2013, PloS one.

[188]  Youngkyoo Jung,et al.  Mindfulness Meditation-Based Pain Relief Employs Different Neural Mechanisms Than Placebo and Sham Mindfulness Meditation-Induced Analgesia , 2015, The Journal of Neuroscience.

[189]  Brian W. Haas,et al.  Functional connectivity during affective mentalizing in criminal offenders with psychotic disorders: Associations with clinical symptoms , 2018, Psychiatry Research: Neuroimaging.

[190]  Dan J Stein,et al.  Impaired acquisition of classically conditioned fear-potentiated startle reflexes in humans with focal bilateral basolateral amygdala damage. , 2015, Social cognitive and affective neuroscience.

[191]  A. Griffin,et al.  Social semantics : altruism , cooperation , mutualism , strong reciprocity and group selection , 2007 .

[192]  Mark Mayford,et al.  Empathic Fear Responses in Mice Are Triggered by Recognition of a Shared Experience , 2013, PloS one.

[193]  E. Viding,et al.  Can Positive Parental Reinforcement Counter Genetic Risk for Callous-Unemotional Behavior? , 2016, The American journal of psychiatry.

[194]  Axel Ockenfels,et al.  Similarity increases altruistic punishment in humans , 2013, Proceedings of the National Academy of Sciences.

[195]  L. Young,et al.  Vasopressin-dependent neural circuits underlying pair bond formation in the monogamous prairie vole , 2004, Neuroscience.

[196]  Samuel M. McClure,et al.  Dissociating affective evaluation and social cognitive processes in the ventral medial prefrontal cortex , 2007, Cognitive, affective & behavioral neuroscience.