Article,

The analysis of variance and the correlations between relatives with respect to deviations from an optimum

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Journal of Genetics, 30 (2): 243--256 (Mar 1, 1935)in "Evolution", collected papers.
DOI: 10.1007/BF02982239

Abstract

The formulae for the mean and variance of squared deviations from an optimum are given for the cases of no dominance and of complete dominance, first assuming no environmental complications but later removing this restriction. The variance in each case is analysed into contributions due to (1) additive gene effects, (2) dominance deviations, (3) epistatic deviations, (4) environmental effects and (5) nonadditive joint effects of heredity and environment. In the case of complete dominance, formulae are developed which apply to epistatic relations in general. These lead to formulae for the correlations between parent and offspring and between two offspring.

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