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DNA REPLICATION Review of DNA Structure • The chemical side groups of the nitrogen bases form hydrogen bonds, connecting the two strands. – Adenine forms two bonds with thymine hydrogen – Guanine forms three hydrogen bonds with cytosine Fig. 16.6 1 Sugar-Phosphate Backbone • The phosphate group of one nucleotide is attached to the sugar of the next nucleotide in line. • The result is a “backbone” of alternating phosphates and sugars, from which the bases project. • The sugar-phosphate backbones of the double helix strands run antiparallel to each other – Each DNA strand has a 3’ end with a free hydroxyl group – and a 5’ end with a free phosphate group – One strand runs 5’ to 3’ and the other runs 3’ to 5’ DNA Replication • Before a cell divides it must replicate its DNA • Semiconservative – the two parent strands serve as a template for the synthesis of the new complementary strands 2 DNA Replication Old Strand New Strand Replication Origin • Replication begins at special sites called origins of replication – There may be hundreds or thousands of origin sites per chromosome. – Strands separate forming a replication “bubble” with replication forks at each end. – The replication bubbles elongate as the DNA is replicated and eventually fuse. Enzymes Involved • Helicase – unwinds and unzips the DNA helix at the replication forks • Topoisomerases – relieves the torque from the unwinding DNA 3 Enzymes • Primase – constructs RNA primer complementary to the DNA templates • After formation of the primer, DNA polymerase III – elongates the new strand by adding nucleotides to the 3’end (~50 per second) • DNA polymerase I – later replaces RNA primers with DNA nucleotides Elongation • When nucleotide triphosphates are linked to the sugar-phosphate backbone it loses two of its phosphates • Replication always occurs in the 5’ to 3’ direction Elongation • Leading strand – elongates toward the replication fork, continuous • Lagging strand – elongates away from the replication fork – Okazaki fragments: discontinuous short segments 4 • Ligase – enzyme that joins fragments into a single DNA strand Proofreading and Repair • Replication is extremely accurate – 1 error per billion nucleotides • DNA polymerase proofreads each nucleotide against the template and fixes any mismatches – Causes: replication errors, physical and chemical agents When does DNA replicate? • DNA replication only occurs when the cell is preparing to divide • When the cell is NOT dividing DNA is used as a blueprint for protein synthesis 5