Darwin accepted blending inheritance, but Fleeming Jenkin calculated that as it mixed traits, natural selection could not accumulate useful traits.

达尔文承认混合遗传，但弗莱•认为当性状是混合的情况下，自然选择不能积有益性状。

Haldane applied mathematical analysis to real world examples of natural selection such as the evolution of industrial melanism in peppered moths.

S.霍尔丹发表了一系列的论文，将数学用到真实世界的自然选择例子，如椒粉蛾工业黑变病的进化。