Forward genetics a bare minimum for the appearance of specific phenotypes. However, such initial screens were either incomplete as they were missing redundant loci and epigenetic effects, and such screens were difficult to undertake for certain phenotypes that lack directly measurable phenotypes. Additionally a classical genetics approach takes significantly longer. Gregor Mendel experimented with pea plant phenotypes and published his conclusions about genes and inheritance in 1865. Around the early 1900s Thomas Hunt Morgan was mutating Drosophila using radium and attempting to find heritable mutations Alfred Sturtevant later began mapping genes of Drosophila with mutations they had been following. In the 1990s
Plant genetics put forth by Mendel. W.E. Castle discovered that while individual traits may segregate and change over time with selection, that when selection is stopped and environmental effects are taken into account, the genetic ratio stops changing and reach a sort of stasis, the foundation of Population Genetics. This was independently discovered by G.H. Hardy and W. Weinberg, which ultimately gave rise to the concept of Hardy-Weinberg equilibrium published in 1908. For a more thorough exploration of the history of population genetics, see History of Population Genetics by Bob Allard. Around this same time, genetic and plant breeding experiments in maize