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===Neurobiology=== {{See also|Compassion}} Jorge Moll and [[Jordan Grafman]], neuroscientists at the [[National Institutes of Health]] and LABS-D'Or Hospital Network, provided the first evidence for the neural bases of altruistic giving in normal healthy volunteers, using [[functional magnetic resonance imaging]]. In their research,<ref>{{cite journal | last1=Moll | first1=Jorge | last2=Krueger | first2=Frank | last3=Zahn | first3=Roland | last4=Pardini | first4=Matteo | last5=de Oliveira-Souza | first5=Ricardo | last6=Grafman | first6=Jordan | title=Human fronto–mesolimbic networks guide decisions about charitable donation | journal=Proceedings of the National Academy of Sciences | volume=103 | issue=42 | date=17 October 2006 | issn=0027-8424 | doi=10.1073/pnas.0604475103 | pages=15623–15628 | pmid=17030808 | pmc=1622872 | bibcode=2006PNAS..10315623M | doi-access=free }}</ref> they showed that both pure monetary rewards and charitable donations activated the [[Mesolimbic pathway|mesolimbic]] reward pathway, a primitive part of the brain that usually responds to food and sex. However, when volunteers generously placed the interests of others before their own by making charitable donations, another [[brain circuit]] was also selectively activated: the subgenual cortex/[[septal region]]. These structures are related to social attachment and bonding in other species. The experiment suggested that altruism is not a higher moral faculty overpowering innate selfish desires, but a fundamental, ingrained, and enjoyable trait in the brain.<ref name="brain">{{cite news |url=https://www.washingtonpost.com/wp-dyn/content/article/2007/05/27/AR2007052701056.html |title=If It Feels Good to Be Good, It Might Be Only Natural |newspaper=The Washington Post |date=28 May 2007 |first=Shankar |last=Vedantam |access-date=23 April 2010}}</ref> One brain region, the subgenual [[anterior cingulate cortex]]/[[basal forebrain]], contributes to learning altruistic behavior, especially in people with a propensity for [[empathy]].<ref name="PMID_27528669">{{multiref2 |1={{cite journal |last1 = Lockwood |first1 = Patricia L |last2 = Apps |first2 = Matthew A J |last3 = Valton |first3 = Vincent |last4 = Viding |first4 = Essi |last5 = Roiser |first5 = Jonathan P |year = 2016 |title = Neurocomputational mechanisms of prosocial learning and links to empathy. |journal = Proceedings of the National Academy of Sciences of the United States of America |quote = . fMRI revealed that activity in a posterior portion of the subgenual anterior cingulate cortex/basal forebrain (sgACC) drives learning only when we are acting in a prosocial context |pmid = 27528669 |doi=10.1073/pnas.1603198113 |volume=113 |issue = 35 |pages=9763–8|pmc = 5024617 |bibcode = 2016PNAS..113.9763L |doi-access = free}} |2={{lay source |template=cite news |date=15 August 2016 |title=Finding the Brain's Generosity Center |url=https://neurosciencenews.com/generosity-empathy-neuroscience-4850/ |work=Neuroscience News}} }}</ref><ref name="neurons">{{cite news |last=Svoboda |first=Elizabeth |date=5 September 2013 |title=Scientists Are Finding That We Are Hard-Wired for Giving |url=https://generosityresearch.nd.edu/news/hard-wired-for-giving/ |access-date=7 August 2017 |publisher=University of Notre Dame}}</ref><ref>{{Cite journal |last1=Shepelenko |first1=A. Yu. |last2=Kosonogov |first2=V. V. |date=1 February 2023 |title=Cerebral Support for Making Donation-Related Decision with Altruistic and Egoistic Motives |url=https://link.springer.com/article/10.1007/s11055-023-01413-9 |journal=Neuroscience and Behavioral Physiology |language=en |volume=53 |issue=2 |pages=242–246 |doi=10.1007/s11055-023-01413-9 |issn=1573-899X}}</ref> Bill Harbaugh, a [[University of Oregon]] economist, in an fMRI scanner test conducted with his psychologist colleague Dr. Ulrich Mayr, reached the same conclusions as Jorge Moll and Jordan Grafman about giving to charity, although they were able to divide the study group into two groups: "egoists" and "altruists". One of their discoveries was that, though rarely, even some of the considered "egoists" sometimes gave more than expected because that would help others, leading to the conclusion that there are other factors in charity, such as a person's environment and values.<ref name="neurons"/> A recent meta-analysis of fMRI studies conducted by Shawn Rhoads, Jo Cutler, and Abigail Marsh analyzed the results of prior studies of generosity in which participants could freely choose to give or not give resources to someone else.<ref>{{cite journal |last1=Rhoads |first1=Shawn A |last2=Cutler |first2=Jo |last3=Marsh |first3=Abigail A |title=A feature-based network analysis and fMRI meta-analysis reveal three distinct types of prosocial decisions |journal=Social Cognitive and Affective Neuroscience |date=30 December 2021 |volume=16 |issue=12 |pages=1214–1233 |doi=10.1093/scan/nsab079 |pmid=34160604 |pmc=8717062 }}</ref> The results of this study confirmed that altruism is supported by distinct mechanisms from giving motivated by reciprocity or by fairness. This study also confirmed that the right ventral striatum is recruited during altruistic giving, as well as the ventromedial prefrontal cortex, bilateral [[anterior cingulate cortex]], and bilateral anterior [[Insular cortex|insula]], which are regions previously implicated in [[empathy]]. [[Abigail Marsh]] has conducted studies of real-world altruists that have also identified an important role for the [[amygdala]] in human altruism. In real-world altruists, such as people who have donated kidneys to strangers, the amygdala is larger than in typical adults. Altruists' amygdalas are also more responsive than those of typical adults to the sight of others' distress, which is thought to reflect an empathic response to distress.<ref name=":0">{{Cite journal |last1=Marsh |first1=Abigail A. |last2=Stoycos |first2=Sarah A. |last3=Brethel-Haurwitz |first3=Kristin M. |last4=Robinson |first4=Paul |last5=VanMeter |first5=John W. |last6=Cardinale |first6=Elise M. |date=21 October 2014 |title=Neural and cognitive characteristics of extraordinary altruists |journal=Proceedings of the National Academy of Sciences|volume=111 |issue=42 |pages=15036–15041 |doi=10.1073/pnas.1408440111 |doi-access=free |issn=0027-8424 |pmc=4210306 |pmid=25225374|bibcode=2014PNAS..11115036M }}</ref><ref>{{Cite journal |last1=Brethel-Haurwitz |first1=Kristin M. |last2=O'Connell |first2=Katherine |last3=Cardinale |first3=Elise M. |last4=Stoianova |first4=Maria |last5=Stoycos |first5=Sarah A. |last6=Lozier |first6=Leah M. |last7=VanMeter |first7=John W. |last8=Marsh |first8=Abigail A. |date=25 October 2017 |title=Amygdala–midbrain connectivity indicates a role for the mammalian parental care system in human altruism |journal=Proceedings of the Royal Society B: Biological Sciences|volume=284 |issue=1865 |pages=20171731 |doi=10.1098/rspb.2017.1731 |issn=0962-8452 |pmc=5666102 |pmid=29070724}}</ref> This structure may also be involved in altruistic choices due to its role in encoding the value of outcomes for others.<ref>{{cite journal |last1=Rhoads |first1=Shawn A |last2=O'Connell |first2=Katherine |last3=Berluti |first3=Kathryn |last4=Ploe |first4=Montana L |last5=Elizabeth |first5=Hannah S |last6=Amormino |first6=Paige |last7=Li |first7=Joanna L |last8=Dutton |first8=Mary Ann |last9=VanMeter |first9=Ashley Skye |last10=Marsh |first10=Abigail A |title=Neural responses underlying extraordinary altruists' generosity for socially distant others |journal=PNAS Nexus |date=3 July 2023 |volume=2 |issue=7 |pages=pgad199 |doi=10.1093/pnasnexus/pgad199 |pmc=10321390 |pmid=37416875 }}</ref> This is consistent with the findings of research in non-human animals, which has identified neurons within the amygdala that specifically encode the value of others' outcomes, activity in which appears to drive altruistic choices in monkeys.<ref>{{Cite journal |last1=Dal Monte |first1=Olga |last2=Chu |first2=Cheng C. J. |last3=Fagan |first3=Nicholas A. |last4=Chang |first4=Steve W. C. |date=April 2020 |title=Specialized medial prefrontal–amygdala coordination in other-regarding decision preference |journal=Nature Neuroscience|volume=23 |issue=4 |pages=565–574 |doi=10.1038/s41593-020-0593-y |pmid=32094970 |pmc=7131896 |issn=1546-1726|hdl=2318/1730693 |hdl-access=free }}</ref><ref>{{cite journal |last1=Putnam |first1=Philip T. |last2=Chu |first2=Cheng-Chi J. |last3=Fagan |first3=Nicholas A. |last4=Dal Monte |first4=Olga |last5=Chang |first5=Steve W.C. |title=Dissociation of vicarious and experienced rewards by coupling frequency within the same neural pathway |journal=Neuron |date=August 2023 |volume=111 |issue=16 |pages=2513–2522.e4 |doi=10.1016/j.neuron.2023.05.020 |pmid=37348507 |pmc=10527039 }}</ref>
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