MSc - Lomonosov Moscow State University, 1978 Candidate of Biology - Research Computing Center, Pushchino, 1984 I worked on multilocus theory of sympatric speciation, the evolution of sex, recombination, and mutation rates, theory of negative selection against deleterious mutations, on properties of spontaneous and induced mutation in Drosophila, and on detection of positive selection and of epistasis from the data on the evolution of genotypes.