Charles Darwin observed that crossed plants of Linaria vulgaris were taller and more vigorous than self-fertilized plants of the same species. Heterosis, coined by Shull in 1952, refers to the increased performance of F1 hybrid plants compared to the average of their inbred parental lines, in traits like biomass, size, yield and resistance. There are several hypotheses for heterosis, including dominance, overdominance and epistasis models. The dominance model proposes that superior performance is due to dominant alleles masking recessive alleles, while the overdominance model suggests heterozygosity itself provides benefits over either homozygote.