Johannsen's criticism of the chromosome theory

The genotype theory of Wilhelm Johannsen (1857-1927) was an important contribution to the founding of classical genetics. This theory built on Johannsen's experimental demonstration that hereditary change is discontinuous, not continuous as had been widely assumed. Johannsen is also known for his criticism of traditional Darwinian evolution by natural selection, as well as his criticism of the classical Mendelian chromosome theory of heredity. He has often been seen as one of the anti-Darwinians that caused the "eclipse of Darwinism" in the early 20th century, before it was saved by the Modern Synthesis. This article focuses on Johannsen's criticism of the chromosome theory. He was indeed skeptical of the notion of the chromosomes as the sole carriers of heredity, but he praised the mapping of Mendelian genes on the chromosomes as a major step forward. Johannsen objected that these genes could not account for the whole of heredity, and that the stability of the genotype depended on much more than the stability of Mendelian genes. For Johannsen, the genotype, as a property of the whole organism, was the fundamental and empirically well-established entity.