Micromechanics of chromatin and chromosomes

The enzymes that transcribe, recombine, package, and duplicate the eukaryotic genome all are highly processive and capable of generating large forces. Understanding chromosome function therefore will require analysis of mechanics as well as biochemistry. Here we review development of new biophysical-biochemical techniques for studying the mechanical properties of isolated chromatin fibers and chromosomes. We also discuss microscopy-based experiments on cells that visualize chromosome structure and dynamics. Experiments on chromatin tell us about its flexibility and fluctuation, as well as quantifying the forces generated during chromatin assembly. Experiments on whole chromosomes provide insight into the higher-order organization of chromatin; for example, recent experiments have shown that the mitotic chromosome is held together by isolated chromatin-chromatin links and not a large, mechanically contiguous non-DNA "scaffold".