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Exonucleases remove these unpaired nucleotides and the gaps are filled by DNA synthesis and repair machinery.
He worked as part of the DNA synthesis group at Genentech from 1986–1990, before joining Gilead in 1990 as Director of Organic Chemistry.
Because the binding of the polymerase to the template is the rate-limiting step in DNA synthesis, the overall rate of DNA replication during S phase of the cell cycle is dependent on the processivity of the DNA polymerases performing the replication.