Dynamics, Harvard University, Cambridge, MA 02138; 'Department of Organismic and Evolutionary Biology, Harvard University, Cambridge, MA 02138; and °Department of Mathematics, Harvard University, Cambridge, MA 02138 Edited by Brian Skyrms, University of California, Irvine, CA, and approved October 16, 2018 (received for

lam2, Martin A. Nowak2 1 IST Austria, KlostemeubLeg, Austria, 2 Program for Evolutionary Dynamics, Department of Organismic and Evolutionary Biology, Department of Mathematics, Harvard University, Cambridge, Massachusetts, United States of America Abstract A fundamental question in biology is the following: what is the t

lam2, Martin A. Nowak2 11ST Austria, Klosterneuburg, AtIWIld, 2 Program for Evolutionary Dynamics, Department of Organismic and Evolutionary Biology. Department of Mathematics, Harvard University• Cambridge. Massachusetts. United States of America Abstract A fundamental question in biology is the following: what is the t

logy Letters 1 2 3 Fear of shame drives cooperation 4 5 6 Jennifer Jacquetl 2., Christoph Hauert', Arne Traulsen3 and Manfred Milinski4 7 8 Department of Mathematics, University of British Columbia, 1984 Mathematics Road, 9 10 Vancouver, British Columbia, V6T 1Z2 Canada (where the research was carried out) 11

l'"', Carl yeller • Raghavendra Gadagkar -1. Martin A. Nowak a•d•g.s Program for twoltalonary Dynamks. Hanard University, Cambridge, MA 02(38. USA °Department of Mathematics. Emmanuel College, Bosron. MA 02115, USA `Center for Mathematical Sciences and Applicanons. Harvard Unhersky. Cambridge, MA 02138. USA Depaytmcnr o

n A. Nowak' 'IST Austria. Klosteineuburg. A 3400. Austria. and 'Program for Evolutionary Dynamics. Department of Organismic and Evolutionary Biology. Department of Mathematics. Harvard University. Cambridge. MA 02138. USA Submitted to Proceedings of the National Academy of Sciences of the United States of America There a