I work on Darwinian adaptation. Natural selection explains the appearance of design in the living world, but at what level is this design expected to manifest – gene, individual, society – and what is its function? Social evolution provides a window on this problem, by pitting the interests of genes, individuals and societies against each other. I develop general theory on the topics of inclusive fitness and multilevel selection, and also tailor general theory to the biology of particular species to facilitate empirical testing. I work on a wide range of biological systems, including viruses, bacteria, protozoa, crustaceans, insects, fish and humans. Visit my webpage at http://synergy.st-andrews.ac.uk/gardner/ for more information.