on the evolutionary origins of altruistic behavior: can selection at the individual level be enough?...
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On the evolutionary On the evolutionary origins of altruistic origins of altruistic behavior:behavior: can selection can selection at the individual level be at the individual level be enough?enough?
José A. Cuesta & Anxo José A. Cuesta & Anxo SánchezSánchez
GISC/Matemáticas GISC/Matemáticas Universidad Carlos III de MadridUniversidad Carlos III de Madrid
8th Granada Seminar – Modelling Complex Social Systems8th Granada Seminar – Modelling Complex Social Systems
Cooperation: the basis Cooperation: the basis of human societiesof human societies
Occurs between genetically unrelated Occurs between genetically unrelated individualsindividuals
Anomaly in the animal Anomaly in the animal world:world:
Cooperation: the basis Cooperation: the basis of human societiesof human societies
Shows high division of laborShows high division of labor
Anomaly in the animal Anomaly in the animal world:world:
Cooperation: the basis Cooperation: the basis of human societiesof human societies
Valid for large scale organizations…Valid for large scale organizations…
Anomaly in the animal Anomaly in the animal world:world:
……as well as hunter-gatherer groupsas well as hunter-gatherer groups
Cooperation: the basis Cooperation: the basis of human societiesof human societies
Valid for large scale organizations…Valid for large scale organizations…
Anomaly in the animal Anomaly in the animal world:world:
……or setting up nice events!or setting up nice events!
Cooperation: the basis Cooperation: the basis of human societiesof human societies
Some animals form complex Some animals form complex societies…societies…
……but their individuals are genetically but their individuals are genetically relatedrelated
Altruism: key to Altruism: key to cooperationcooperation
Altruism:Altruism:
fitness-reducing act that benefits othersfitness-reducing act that benefits others
Pure altruism is ruled out by natural Pure altruism is ruled out by natural selection acting on individuals selection acting on individuals á la Darwiná la Darwin
He who was ready to sacrifice his life (…), rather than betray his comrades, would often leave no offspring to inherit his noble nature… Therefore, it seems scarcely possible (…) that the number of men gifted with such virtues (…) would be increased by natural selection, that is, by the survival of the fittest.
Charles Darwin (Descent of Man, 1871)
How did altruism arise?How did altruism arise?
A man who was not impelled by any deep, instinctive feeling, to sacrifice his life for the good of others, yet was roused to such actions by a sense of glory, would by his example excite the same wish for glory in other men, and would strengthen by exercise the noble feeling of admiration. He might thus do far more good to his tribe than by begetting offsprings with a tendency to inherit his own high character.
Charles Darwin (Descent of Man, 1871)
Group selection? Cultural Group selection? Cultural evolution?evolution?
Answers to the puzzle… Answers to the puzzle…
Kin cooperationKin cooperation (Hamilton, 1964)(Hamilton, 1964)
common to animals and humans alikecommon to animals and humans alike Reciprocal altruism in repeated Reciprocal altruism in repeated
interactionsinteractions (Trivers, 1973;(Trivers, 1973; Axelrod & Hamilton, Axelrod & Hamilton, 1981)1981)
primates, specially humansprimates, specially humans Indirect reciprocity (reputation gain)Indirect reciprocity (reputation gain)
(Nowak & Sigmund, 1998)(Nowak & Sigmund, 1998)
primates, specially humansprimates, specially humansNone true altruism: individual benefits in the None true altruism: individual benefits in the long runlong run
… … but only partial!but only partial!
Strong reciprocityStrong reciprocity (Gintis, 2000; Fehr, Fischbacher & Gächter, 2002)(Gintis, 2000; Fehr, Fischbacher & Gächter, 2002)
typically human (primates?)typically human (primates?)– altruistic rewarding:altruistic rewarding: predispositionpredisposition to to
reward others for cooperative behaviorreward others for cooperative behavior– altruistic punishment:altruistic punishment: propensitypropensity to to
impose sanctions on non-cooperatorsimpose sanctions on non-cooperators Strong reciprocators bear the cost of Strong reciprocators bear the cost of
altruistic actsaltruistic acts even if they gain no benefits even if they gain no benefits
Hammerstein (ed.), Genetic and cultural evolution of cooperation (Dahlem Workshop Report 90, MIT, 2003)
ExperimentsExperiments
experimentexperimenterer
subject 1subject 1 subject 2subject 2
2-3 month’s income2-3 month’s income
Altruistic punishment: Altruistic punishment: the Ultimatum Game the Ultimatum Game(Güth, Schmittberger & Schwarze, 1982)(Güth, Schmittberger & Schwarze, 1982)
experimentexperimenterer
proposeproposerr
respondresponderer
M M euroseuros
M-uM-u uuOKOK
uuM-uM-u
NNOO
00 00
Experimental resultsExperimental results
Proposers offer substantial amounts (50% Proposers offer substantial amounts (50% is a typical modal offer)is a typical modal offer)
Responders reject offers below 25% with Responders reject offers below 25% with high probabilityhigh probability
Universal behavior throughout the worldUniversal behavior throughout the world Large degree of variability of offers Large degree of variability of offers
among societies (26 - 58%)among societies (26 - 58%)
Proposer’s optimal strategy: Proposer’s optimal strategy: offer the offer the minimumminimum
Responder’s optimal strategy: Responder’s optimal strategy: accept accept anythinganything
Results:Results:
Experimental resultsExperimental results
Extraordinary amount of dataExtraordinary amount of data
Camerer, Behavioral Game Theory (Princeton University Press, 2003)
Henrich et al. (eds.), Foundations of Human Sociality : Economic Experiments and Ethnographic Evidence from Fifteen Small-Scale Societies (Oxford University Press, 2004)
Experimental resultsExperimental results
Paciotti Paciotti et al.et al., , American ScientistAmerican Scientist 9393, 58 (2005), 58 (2005)
Ultimatum game as an interpretingtool
Altruistic rewardingAltruistic rewarding(Fehr, Kirchsteiger & Riedl, 1993)(Fehr, Kirchsteiger & Riedl, 1993)
experimentexperimenterer
trustetrusterr
trustetrusteee
MMMM-u-u +2u+2u-v-v+2v+2v
Experimental resultsExperimental results
More than 50% of trustees transfer money More than 50% of trustees transfer money back, the more the higher the truster’s back, the more the higher the truster’s transfertransfer
Players often exchangePlayers often exchange M M Universal behaviour throughout the worldUniversal behaviour throughout the worldHomo economicus vs. Homo reciprocansReview in Fehr & Fischbacher, Review in Fehr & Fischbacher, NatureNature 425, 785 (2003) 425, 785 (2003)
Both players gain the maximum by Both players gain the maximum by exchanging exchanging MM The trustee gains more by returning The trustee gains more by returning nothingnothing
Results:Results:So truster’s best strategy is to transfer So truster’s best strategy is to transfer nothingnothing
Strong reciprocityStrong reciprocity
Kin cooperationKin cooperation selfish genes selfish genes ReciprocityReciprocity tit-for-tattit-for-tat strategy strategy
(Axelrod’s experiments)(Axelrod’s experiments) Reputation gainReputation gain altruism as sexual altruism as sexual
ornament ornament (handicap principle)(handicap principle) Strong reciprocity sense of Strong reciprocity sense of fairnessfairness
requires “other-regarding” capabilitiesrequires “other-regarding” capabilities
Explanations for altruistic behaviour:Explanations for altruistic behaviour:
Strong reciprocityStrong reciprocity
In experiments of public goods, the In experiments of public goods, the presence of a few cheaters quickly presence of a few cheaters quickly deteriorates cooperationdeteriorates cooperation
The introduction of a few The introduction of a few altruistic altruistic punishers punishers forces long term cooperation forces long term cooperation
Strong reciprocity seems a crucial element Strong reciprocity seems a crucial element to ensure cooperation in human societiesto ensure cooperation in human societies
Importance of strong reciprocity:Importance of strong reciprocity:
Strong reciprocityStrong reciprocity
Standard answer: Standard answer: cultural evolution cultural evolution through group selection through group selection
Evidences in favour:Evidences in favour:– Strong variability between cultures in Strong variability between cultures in
altruistic patternsaltruistic patterns– Increase of altruistic behaviour in Increase of altruistic behaviour in
children as they growchildren as they grow– Simulations of group selection modelsSimulations of group selection models
Evolutionary origins of strong Evolutionary origins of strong reciprocity:reciprocity:
Our model: setupOur model: setup
............ NN playersplayers
player player ii
ttii : threshold (minimum share : threshold (minimum share player player ii accepts) accepts)
ffii : fitness (accumulated capital) : fitness (accumulated capital)
MM monetary units ( monetary units (MM=100) =100)
A.S. & J. A. Cuesta, A.S. & J. A. Cuesta, J. Theor. Biol., J. Theor. Biol., in press (2005)in press (2005)
Our model: game stepOur model: game step
............ NN playersplayers
proposeproposerr
responderesponderr
tt
pp
ffpp
ttrr
ffrr
ttp p
ttrr
≥≥
+M-+M-ttpp
+t+tpp
ttp p ttrr <<
Our model: every Our model: every ss stepssteps
............ NN playersplayers
minimuminimum m
fitnessfitness
maximumaximum fitnessm fitness
ttmimi
nn
ffminmin
ttmaxmax
ffmaxmax
new new playerplayer
t’t’maxmax
ffmaxmax
mutation: mutation: t’t’maxmax==
ttmaxmax
(prob.=1/3(prob.=1/3))
+1 +1 -1 -1
Implemented featuresImplemented features
Degree of altruism inherited from Degree of altruism inherited from progenitorprogenitor (with some error) (with some error)
No learning from experienceNo learning from experience No influence from other playersNo influence from other players (no (no
culture)culture) No groupsNo groups Other-regarding behaviour with Other-regarding behaviour with
optimizationoptimization (players offer the minimum (players offer the minimum they would accept)they would accept)
Fitness Fitness (accumulated capital) (accumulated capital) is inheritedis inherited
Why are Why are reciprocators reciprocators selected?selected? Higher threshold implies lower payoffs but Higher threshold implies lower payoffs but
higher acceptance rateshigher acceptance rates for agent’s own for agent’s own offersoffers
The presence of altruistic punishers The presence of altruistic punishers increases increases rejection of selfish agents’ offersrejection of selfish agents’ offers
Mutations are smallMutations are small (darwinian), so (darwinian), so newborn altruistic agents are just a little newborn altruistic agents are just a little more altruisticmore altruistic
A quantitative description is still lackingA quantitative description is still lacking
Two thresholds (s Two thresholds (s small)small)
N =1000, 106 games, s =1, uniform initial condition
acceptaccept
offeroffer
Two thresholds (s Two thresholds (s middle)middle)
N =1000, 108 games, s =1000, ti = ai =1 initial condition
acceptaccept
offeroffer
Two thresholds (s Two thresholds (s large)large)
N =1000, 109 games, s =105, ti = ai =1 initial condition
acceptaccept
offeroffer
DiscussionDiscussion
Altruistic punishment is not necessarily a Altruistic punishment is not necessarily a losing strategy when there are many losing strategy when there are many agentsagents
Thus it may be established by individual Thus it may be established by individual selection aloneselection alone
Results reproduce the observed variabilityResults reproduce the observed variability
One or two levels: not important for the One or two levels: not important for the arising of altruistic punishmentarising of altruistic punishment
Mutation rate might be relevantMutation rate might be relevant
Related resultsRelated results
Stag-hunt game:
• Two equilibria, S and R
• Round-robin gaming: S or Rselected from x0
(x fraction of R)
• s games between death-birth
S R
C. P. Roca, J. A. Cuesta & A. S., in progress (2005)C. P. Roca, J. A. Cuesta & A. S., in progress (2005)
DiscussionDiscussion In general, evolutionary game theory In general, evolutionary game theory
studies a limit situation: studies a limit situation: s infinite! s infinite! (every player plays every other one (every player plays every other one before selection)before selection)
Number of games per player Poisson Number of games per player Poisson distributeddistributed
Fluctuations may keep players with Fluctuations may keep players with smaller ‘mean-field’ fitness alivesmaller ‘mean-field’ fitness alive
Further discussionFurther discussion Thresholds observed in neural activity Thresholds observed in neural activity
measurements measurements (Sanfey et al., 2003) (Sanfey et al., 2003) (suggest that this threshold is an (suggest that this threshold is an inheritable feature) inheritable feature)
Correlation with other-regarding Correlation with other-regarding behavior: also in different primates behavior: also in different primates (capuchin monkeys) (capuchin monkeys) (Brosnan & de (Brosnan & de Waal, 2003)Waal, 2003)
Main conclusionsMain conclusions
Altruism Altruism (altruistic punishment, as (altruistic punishment, as implemented in the Ultimatum Game)implemented in the Ultimatum Game) need need not be an evolutionary losing strategy, not be an evolutionary losing strategy, so it so it may have arisen through standard may have arisen through standard individual selectionindividual selection
A new perspective in evolutionary game A new perspective in evolutionary game theory: theory: more general dynamics, dictated more general dynamics, dictated by the specific applicationby the specific application
Adam Smith on Adam Smith on altruismaltruismHow selfish soever man may be supposed, there are evidently some principles in his nature, which interest him in the fortunes of others, and render their happiness necessary to him, though he derives nothing from it, except the pleasure of seeing it.
Adam Smith (The Theory of Moral Sentiments, 1854, Chapter 1)