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If we reverse the order to f→f+/a→A, then the allele f+ already exists in the background state and the connection is completed by the mutation of the a allele to the A allele which contains a novel output regulatory domain.
For a→A/X mutation pairs (i.e. the A regulatory allele already exists in the background state before a cis mutation occurs in the m or f alleles), polymorphism occurs at the D locus, while for X/a→A mutation pairs (i.e. cis mutation occurs in the m or f allele before mutation to the A allele), polymorphism occurs at both the A and D loci.
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