Chemical structure

In 1953 James D. Watson and Francis H.C. Crick proposed a three-dimensional structure for DNA based on low-resolution X-ray crystallographic data and on Erwin Chargaff’s observation that, in naturally occurring DNA, the amount of T equals the amount of A and the amount of G equals the amount of C. Watson and Crick, who shared a Nobel Prize in 1962 for their efforts, postulated that two strands of polynucleotides coil around each other, forming a double helix. The two strands, though identical, run in opposite directions as determined by the orientation of the 5′ to 3′ phosphodiester bond. The sugar-phosphate chains run along the outside of the helix, and the bases lie on the inside, where they are linked to complementary bases on the other strand through hydrogen bonds.

The double helical structure of normal DNA takes a right-handed form called the B-helix. The helix makes one complete turn approximately every 10 base pairs. B-DNA has two principal grooves, a wide major groove and a narrow minor groove. Many proteins interact in the space of the major groove, where they make sequence-specific contacts with the bases. In addition, a few proteins are known to make contacts via the minor groove.

Several structural variants of DNA are known. In A-DNA, which forms under conditions of high salt concentration and minimal water, the base pairs are tilted and displaced toward the minor groove. Left-handed Z-DNA forms most readily in strands that contain sequences with alternating purines and pyrimidines. DNA can form triple helices when two strands containing runs of pyrimidines interact with a third strand containing a run of purines.

B-DNA is generally depicted as a smooth helix; however, specific sequences of bases can distort the otherwise regular structure. For example, short tracts of A residues interspersed with short sections of general sequence result in a bent DNA molecule. Inverted base sequences, on the other hand, produce cruciform structures with four-way junctions that are similar to recombination intermediates. Most of these alternative DNA structures have only been characterized in the laboratory, and their cellular significance is unknown.