B chromosomes are extra chromosomes to the standard complement that occur in many organisms. They can originate in a number of ways including derivation from autosomes and sex chromosomes in intra- and interspecies crosses. Their subsequent molecular evolution resembles that of univalent sex chromosomes, which involves gene silencing, heterochromatinization and the accumulation of repetitive DNA and transposons. B-chromosome frequencies in populations result from a balance between their transmission rates and their e¡ects on host ¢tness. Their long-term evolution is considered to be the outcome of selection on the host genome to eliminate B chromosomes or suppress their e¡ects and on the B chromosome’s ability to escape through the generati...