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Indirect reciprocity is cooperation in which helping one individual affects the help received from a different individual, often through reputation. For example, individual A helps B; C observes the act and later helps A. B need never repay A. In direct reciprocity, by contrast, the same pair exchanges help over repeated encounters.

Martin A. Nowak and Karl Sigmund's 1998 model assigned individuals an image score reflecting earlier helping. Potential donors used that information when deciding whom to help, and successful strategies became more frequent. In the model, let $c$ be the donor's cost, $b$ the recipient's benefit, and $q$ the probability of knowing a recipient's reputation. An evolutionarily stable strategy based on helping required $q>c/b$. With $c=1$ and $b=4$, the threshold is $q>0.25$; merely observing that help is beneficial does not meet this information requirement.^[ref]

The condition belongs to this reputation model. It is not a universal numerical threshold for every system described as indirect reciprocity.

Karl Sigmund applies mathematics to the evolution of cooperation. His work in evolutionary game theory examines how strategies spread when their success depends on interactions with other individuals.^[ref]

With Martin A. Nowak, Sigmund developed a 1998 model of indirect reciprocity: an individual may help someone because that person's previous behaviour has established a favourable reputation, even if the same pair never meets again. The analysis combined mathematical conditions for stability with computer simulations of changing strategy frequencies.^[ref]

For example, A helps B, C learns of the act, and C later helps A. The return to A passes through another individual instead of requiring repayment by B. The scientific question is which information and decision rules allow such behaviour to persist against alternatives that avoid the cost of helping. A reputation mechanism makes those rules explicit; it does not establish that every apparently generous action is explained by the same process.^[ref]

Martin A. Nowak works on mathematical models of evolution, including the evolution of cooperation. His 2006 synthesis separated five mechanisms: kin selection, direct reciprocity, indirect reciprocity, network reciprocity and group selection. Each mechanism changes which interactions contribute to an individual's evolutionary success.^[ref]

A simple direct reciprocity calculation shows how repeated encounters can change incentives. Suppose cooperation costs $c$, benefits the partner by $b>c$, and an encounter continues with probability $w$. Two cooperating reciprocators obtain expected payoff $(b-c)/(1-w)$. An unconditional defector against a strategy that stops helping after a defection obtains $b$ initially and no later benefit. The first payoff exceeds the second precisely when $w>c/b$. With $b=3$, $c=1$ and $w=1/2$, these payoffs are four and three.

This calculation depends on the specified repeated-interaction model and retaliation rule. It identifies a mechanism by which cooperation can be favored, without asserting that a single inequality describes every biological system. His institutional research profile also lists mathematical work on infection and cancer.^[ref]

Portrait source: www.oeb.harvard.edu/people/martin-nowak .