The differences between the leaf characters of two species of golden-rod, Solidago rugosa and S. sempervirens, have been shown to be determined by a minimum of 21 genes. It is pointed out that a minimum of about twice as many genes are probably in operation in the control of all the morphologically distinguishable differences between these two species. A method has been devised to estimate the maximum number of gene differences common to two morphological character differences between related strains or interfertile species from the standard deviations and correlations of the character pairs in the F1 and F2 hybrids.
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Charles et al. (1943) studied this question.