At the turn of the twentieth century, William Bateson studied organismal variation and heredity of traits within the framework of evolutionary theory in England. Bateson applied Gregor Mendel's work to Charles Darwin's theory of evolution and coined the term genetics for a new biological discipline. By studying variation and advocating Mendelian genetics, Bateson furthered the field of genetics, encouraged the use of experimental methodology to study heredity, and contributed to later theories of genetic inheritance.
Muriel Wheldale Onslow studied flowers in England with genetic and biochemical techniques in the early twentieth century. Working with geneticist William Bateson, Onslow used Mendelian principles and biochemical analysis together to understand the inheritance of flower colors at the beginning of the twentieth century. Onslow's study of snapdragons, or Antirrhinum majus, resulted in her description of epistasis, a phenomenon in which the phenotypic effect of one gene is influenced by one or more other genes. She discovered several biochemicals related to color formation. Onslow's methodology also partly contributed to the establishment of the field of chemical genetics.
In 1868 in England, Charles Darwin proposed his pangenesis theory to describe the units of inheritance between parents and offspring and the processes by which those units control development in offspring. Darwin coined the concept of gemmules, which he said referred to hypothesized minute particles of inheritance thrown off by all cells of the body. The theory suggested that an organism's environment could modify the gemmules in any parts of the body, and that these modified gemmules would congregate in the reproductive organs of parents to be passed on to their offspring. Darwin's theory of pangenesis gradually lost popularity in the 1890s when biologists increasingly abandoned the theory of inheritance of acquired characteristics (IAC), on which the pangenesis theory partially relied. Around the turn of the twentieth century, biologists replaced the theory of pangenesis with germ plasm theory and then with chromosomal theories of inheritance, and they replaced the concept of gemmules with that of genes.