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Deletion Mapping -

Many of the spontaneous r- II mutants isolated by Benzer failed to recombine with two or more mutants that recombine with each other.

They behaved as though they contained multisite mutations or defects that extended over a segment of the r- II locus. Some of them appeared to span the entire rII, others only a short segment.

Benzer proposed that these multisite mutations resulted from the deletion or loss of segments of the DNA.

As they failed to undergo back mutation, Benzers proposal has subsequently been verified directly by analysis of several deletion mutations by electron microscopy.