Situations where individuals have to contribute to joint efforts or share
scarce resources are ubiquitous. Yet, without proper mechanisms to ensure
cooperation, the evolutionary pressure to maximize individual success tends to
create a tragedy of the commons (such as over-fishing or the destruction of our
environment). This contribution addresses a number of related puzzles of human
behavior with an evolutionary game theoretical approach as it has been
successfully used to explain the behavior of other biological species many
times, from bacteria to vertebrates. Our agent-based model distinguishes
individuals applying four different behavioral strategies: non-cooperative
individuals ("defectors"), cooperative individuals abstaining from punishment
efforts (called "cooperators" or "second-order free-riders"), cooperators who
punish non-cooperative behavior ("moralists"), and defectors, who punish other
defectors despite being non-cooperative themselves ("immoralists"). By
considering spatial interactions with neighboring individuals, our model
reveals several interesting effects: First, moralists can fully eliminate
cooperators. This spreading of punishing behavior requires a segregation of
behavioral strategies and solves the "second-order free-rider problem". Second,
the system behavior changes its character significantly even after very long
times ("who laughs last laughs best effect"). Third, the presence of a number
of defectors can largely accelerate the victory of moralists over non-punishing
cooperators. Forth, in order to succeed, moralists may profit from immoralists
in a way that appears like an "unholy collaboration". Our findings suggest that
the consideration of punishment strategies allows to understand the
establishment and spreading of "moral behavior" by means of game-theoretical
concepts. This demonstrates that quantitative biological modeling approaches
are powerful even in domains that have been addressed with non-mathematical
concepts so far. The complex dynamics of certain social behaviors becomes
understandable as result of an evolutionary competition between different
behavioral strategies.Comment: 15 pages, 5 figures; accepted for publication in PLoS Computational
Biology [supplementary material available at
http://www.soms.ethz.ch/research/secondorder-freeriders/ and
http://www.matjazperc.com/plos/moral.html