Wilhelm Ludvig Johannsen studied plants and helped found the field of genetics, contributing methods and concepts to the study of heredity around the turn of the twentieth century in Denmark. His experiments on heredity and variation in plants influenced the methods and techniques of geneticists, and his distinction between the genotype of an organism-its hereditary disposition-and its phenotype-its observable characteristics-remains at the core of contemporary biology. Johannsen criticized biological explanations that relied on concepts such as vitalism and teleology. For an alternative, he advocated a realist and materialist approach to biology, but one that did not attempt to reduce biological phenomena to the laws of physics and chemistry.
Richard Woltereck first described the concept of Reaktionsnorm (norm of reaction) in his 1909 paper 'Weitere experimentelle Untersuchungen uber Art-veranderung, speziell uber das Wesen quantitativer Artunterschiede bei Daphniden' ('Further investigations of type variation, specifically concerning the nature of quantitative differences between varieties of Daphnia'). This concept refers to the ways in which the environment can alter the development of an organism, and its adult characteristics. Woltereck conceived of the Reaktionsnorm as the full range of potentialities latent in a single genotype, evocable by the environmental circumstances of a developing organism. Biologists used variants of Woltereck's concept of Reaktionsnorm, often called the reaction norm or norm of reaction, throughout the twentieth century in attempts to explain how developmental responses to the environment can evolve, and even alter the tempo and direction of evolutionary change.
Wilhelm Johannsen in Denmark first proposed the distinction between genotype and phenotype in the study of heredity in 1909. This distinction is between the hereditary dispositions of organisms (their genotypes) and the ways in which those dispositions manifest themselves in the physical characteristics of those organisms (their phenotypes). This distinction was an outgrowth of Johannsen's experiments concerning heritable variation in plants, and it influenced his pure line theory of heredity. While the meaning and significance of the genotype-phenotype distinction has been a topic of debate-among Johannsen's contemporaries, later biological theorists, and historians of science-many consider the distinction one of the conceptual pillars of twentieth century genetics. Moreover some have used it to characterize the relationships between studies of development, genetics, and evolution.
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.