International
Tables for Crystallography Volume F Crystallography of biological macromolecules Edited by M. G. Rossmann and E. Arnold © International Union of Crystallography 2006 |
International Tables for Crystallography (2006). Vol. F. ch. 23.3, p. 609
Section 23.3.5. Summary
a
Molecular Biology Institute, University of California, Los Angeles, Los Angeles, CA 90095–1570, USA |
Three families of nucleic acid double helix have been found – A, B and Z – with widely different structures and usages. The A and B helices are right-handed and have no limitations on base sequence. Z is left-handed and effectively limited to alternating purines and pyrimidines, with G and C overwhelmingly favoured. B is the biologically significant helix for DNA and is used in genetic coding. A is the helix of preference for RNA because it can accommodate the C2′-OH group of ribose, which produces steric clash in the B helix. The Z helix has, as yet, no well established biological function. A left-handed DNA configuration can be induced in longer DNA segments by negative supercoiling in solution, but it is not clear that this left-handed configuration is identical to the Z-DNA seen in short crystalline oligomers, because of the reversed orientation of backbone strands in Z-DNA.
B-DNA is an inherently malleable or deformable duplex. Its sugar ring conformations are much more variable than those of A-DNA. The base sequence of B-DNA is expressed directly via hydrogen bonds between bases of a pair, and indirectly via hydrogen-bond donors and acceptors along the floor of the major and minor groove. Sequence is also expressed as a differential deformability of different regions of the duplex. The two most obvious parameters affected by base sequence are minor groove width and helix bendability. Certain sequences of B-DNA are not statically bent, but are more bendable under stress than are other sequences. Bending occurs via roll, usually in the direction that compresses the broad major groove. Pyrimidine-purine or Y-R steps are most conducive to roll bending, and purine-purine steps are least bendable, particularly A-tracts of four or more AT base pairs without the weak T-A step. Natural selection has engineered Y-R steps into a DNA sequence where a sharp roll bend is wanted, and short A-tracts into a sequence where bending is not desired.
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