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330 Force spectroscopy of single DNA and RNA molecules Mark C Williams* and Ioulia Rouzina? Experiments in which single molecules of RNA and DNA are The resolution of this method is about 5 pN, but it can also stretched, and the resulting force as a function of extension is be used to measure forces in the nanonewton range. This measured have yielded new information about the physical, wide range of techniques has been used to study several chemical and biological properties of these important molecules. regimes of DNA stretching behavior in detail and has also The behavior of both single-stranded and double-stranded been extended to study DNA–protein and DNA–drug nucleic acids under changing solution conditions, such as ionic interactions. In addition, high-resolution force measure- strength, pH and temperature, has been studied in detail. There ments have been obtained from the stretching of single has also been progress in using these techniques to study both RNA molecules, suggesting that these methods can be the kinetics and equilibrium thermodynamics of DNA–protein extended to the study of sequence-specific RNA–protein interactions. These studies generate unique insights into the interactions. functions of these proteins in the cell. Stretching nucleic acids at low forces: DNA Addresses elasticity, and DNA and RNA unzipping *Department of Physics and Center for Interdisciplinary Research on The force/extension curve of double-stranded (ds) DNA Complex Systems, Northeastern University, 111 Dana Research can be fit to the following equation: Center, Boston, MA 02115, USA; e-mail: mark@neu.edu ?Department of Biochemistry, Molecular Biology and Biophysics, FP 1 ? 1 ? b University of Minnesota, St Paul, MN 55108, USA ds = ? ?