The highly concentrated liquid-crystalline phase of DNA is columnar hexagonal

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Abstract
THE DNA molecule is extremely compacted in bacteria, in cell nuclei, sperm heads and virus capsids. These interactions between DNA molecules are important to our understanding of chromatin condensation. DNA forms multiple liquid-crystalline phases whose nature depends on the polymer concentration1–12, and it has been suggested that the highly concentrated phase of 50-nm DNA molecules is two-dimensionally ordered and smectic-like13. We rule out this smectic hypothesis and demonstrate by polarizing microscopy, electron microscopy and X-ray diffraction that this phase is characterized by a columnar longitudinal order and a hexagonal lateral order, with intermolecular distances ranging from 2.8 to 4.0 nm depending on the DNA concentration.