NOTE 17H: There will also be a slight discrepancy because of deviations from Hardy–Weinberg proportions and linkage equilibrium. If + alleles and – alleles tended to pair up in +– heterozygotes or in +– chromosomes, then there would be less variation in fitness and less change in allele frequency than expected from the Fundamental Theorem. We examine the consequences of such negative associations for the evolution of sex and recombination in Chapter 23.