August Friedrich Leopold Weismann studied how the traits of organisms developed and evolved in a variety of organisms, mostly insects and aquatic animals, in Germany in the late nineteenth and early twentieth centuries. Weismann proposed the theory of the continuity of germ-plasm, a theory of heredity. Weismann postulated that germ-plasm was the hereditary material in cells, and parents transmitted to their offspring only the germ-plasm present in germ-cells (sperm and egg cells) rather than somatic or body cells. Weismann also promoted Charles Darwin's 1859 theory of the evolution of species. Weismann argued that only changes to the germ cells, and not body cells, could be inherited, a theory that influenced theories of heredity throughout later centuries.
Richard Woltereck was a German zoologist and hydrobiologist who studied aquatic animals and extended the concept of Reaktionsnorm (norm of reaction) to the study of genetics. He also provided some of the first experimental evidence for the early twentieth-century embryological theory of heredity known as cytoplasmic inheritance. Through experiments on the water flea, Daphnia, Woltereck investigated whether variation produced by environmental impacts on development could play a role in heredity and evolution. Woltereck's research emphasized the importance of environment and development in Wilhelm Johannsen's concepts of genotype and phenotype. Biologists throughout the twentieth century used Woltereck's concept of Reaktionsnorm to develop theories and experiments to explain the evolution of adaptive developmental responses to environmental conditions. Later in his career, Woltereck developed a theory of heredity that sought to reconcile embryological concepts, such as regulation and body plans, with Mendelian heredity and Darwinian evolution by natural selection.
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.