Wilhelm Johannsen

Wilhelm Johannsen (3 February 1857 – 11 November 1927) was a Danish pharmacist, botanist, plant physiologist, and geneticist. He is best known for coining the terms gene, phenotype and genotype, and for his 1903 "pure line" experiments in genetics.

Biography
Johannsen was born in Copenhagen. While very young, he was apprenticed to a pharmacist and worked in Denmark and Germany beginning in 1872 until passing his pharmacist's exam in 1879. In 1881, he became assistant in the chemistry department at the Carlsberg Laboratory under the chemist Johan Kjeldahl. Johannsen studied the metabolism of dormancy and germination in seeds, tubers and buds. He showed that dormancy could be broken by various anesthetic compounds, such as diethyl ether and chloroform.

In 1892, he was appointed lecturer at Royal Veterinary and Agricultural University and later became professor of botany and plant physiology. He taught plant physiology. His best-known research concerned so-called pure lines of the self-fertile common bean. He was able to show that even in populations homozygous for all traits, i.e. without genetic variation, seed size followed a normal distribution. This was attributable to resource provision to the mother plant and to the position of seeds in pods and of pods on the plant. This led him to coin the terms phenotype and genotype.

Johannsen's findings led him to oppose contemporary Darwinists, most notably Francis Galton and Karl Pearson, who held the occurrence of normal distributed trait variation in populations as proof of gradual genetic variation on which selection could act. Only with the modern synthesis, was it established that variation needed to be heritable to act as the raw material for selection.

He created the terms phenotype and genotype, first using them in his book in German as Elemente der exakten Erblichkeitslehre (Elements of the exact theory of heredity). This book was based in large part on Om arvelighed i samfund og i rene linier ("On heredity in society and in pure lines") and in his book Arvelighedslærens Elementer. It was in this book Johannsen also introduced the term gene. From p. 124: "Dieses "etwas" in den Gameten bezw. in der Zygote, … – kurz, was wir eben Gene nennen wollen – bedingt sind."  (This "something" in the gametes or in the zygote, which has crucial importance for the character of the organism, is usually called by the quite ambiguous term Anlagen [primordium, from the German word Anlage for "plan, arrangement ; rough sketch"].  Many other terms have been suggested, mostly unfortunately in closer connection with certain hypothetical opinions.  The word "pangene", which was introduced by Darwin, is perhaps used most frequently in place of Anlagen.  However, the word "pangene" was not well chosen, as it is a compound word containing the roots pan (the neuter form of Πας all, every) and gen (from γί-γ (ε)ν-ομαι, to become).  Only the meaning of this latter [i.e., gen] comes into consideration here ; just the basic idea – [namely,] that a trait in the developing organism can be determined or is influenced by "something" in the gametes – should find expression. No hypothesis about the nature of this "something" should be postulated or supported by it. For that reason it seems simplest to use in isolation the last syllable gen from Darwin's well-known word, which alone is of interest to us, in order to replace, with it, the poor, ambiguous word Anlage. Thus we will say simply "gene" and "genes" for "pangene" and "pangenes". The word gene is completely free of any hypothesis ; it expresses only the established fact that in any case many traits of the organism are determined by specific, separable, and thus independent "conditions", "foundations", "plans" – in short, precisely what we want to call genes.) This term was coined in opposition to the then common pangene that stemmed from Darwin's theory of pangenesis. The book became one of the founding texts of genetics.

Also in 1905, Johannsen was appointed professor of plant physiology at the University of Copenhagen, becoming vice-chancellor in 1917. In December 1910, Johannsen was invited to give an address before the American Society of Naturalists. This talk was printed in the American Naturalist. In 1911, he was invited to give a series of four lectures at Columbia University.

Johannsen was a corresponding member of the Academy of Natural Sciences of Philadelphia (elected 1915). He was elected to the American Philosophical Society in 1916.