The Full Breadth of Mendel’s Genetics

Gregor Mendel’s “Experiments on Plant Hybrids” (1865/1866), published 150 years ago, is without doubt one of the most brilliant works in biology. Curiously, Mendel’s later studies on Hieracium (hawkweed) are usually seen as a frustrating failure, because it is assumed that they were intended to confirm the segregation ratios he found in Pisum. Had this been his intention, such a confirmation would have failed, since, unknown to Mendel, Hieracium species mostly reproduce by means of clonal seeds (apomixis). Here we show that this assumption arises from a misunderstanding that could be explained by a missing page in Mendel’s first letter to Carl Nägeli. Mendel’s writings clearly indicate his interest in “constant hybrids,” hybrids which do not segregate, and which were “essentially different” from “variable hybrids” such as in Pisum. After the Pisum studies, Mendel worked mainly on Hieracium for 7 years where he found constant hybrids and some great surprises. He also continued to explore variable hybrids; both variable and constant hybrids were of interest to Mendel with respect to inheritance and to species evolution. Mendel considered that their similarities and differences might provide deep insights and that their differing behaviors were “individual manifestations of a higher more fundamental law.”