Plant Phenotypes as Distributions: Johannsen's Beans Revisited

AbstractIn the early twentieth century, Wilhelm Johannsen's breeding experiments on pure lines of beans provided empirical support for his groundbreaking distinction between phenotype and genotype, the foundation stone of classical genetics. In contrast with the controversial history of the genotype concept, the notion of phenotype has remained essentially unrevised since then. The application of the Johannsenian concept of phenotype to modularly built, nonunitary plants, however, needs reexamination. In the first part of this article it is shown that Johannsen's appealing solution for dealing with the multiplicity of nonidentical organs produced by plant individuals (representing individual plant phenotypes by arithmetic means), which has persisted to this day, reflected his intellectual commitment to nineteenth-century typological thinking. Revisitation of Johannsen's results using current statistical tools upholds his major conclusion about the nature of heredity but at the same time falsifies two important ancillary conclusions of his experiments-namely, the alleged homogeneity of pure lines (genotypes) regarding seed weight variability and the lack of transgenerational effects of within-line (within-genotype) seed weight variation. The canonical notion of individual plant phenotypes as arithmetic means should therefore be superseded by a concept of phenotype as a dual property, consisting of central tendency and variability components of organ trait distribution. Phenotype duality offers a unifying framework applicable to all nonunitary organisms.