Macrodipoles. Unusual electric properties of biological macromolecules.

The wide range of different effects induced by electric fields in biological macromolecules is clearly due to the unusual quality and quantity of their electric parameters. A general concept for a quantitative description of the polarizability of macromolecules remains to be established. In the case of DNA, experimental data indicate the existence of an effective polarization length N(p); at chain lengths N < N(p) the polarizability increases with N(2), whereas saturation is approached at N > or = N(p). The polarization length decreases with increasing ionic strength in close analogy to the Debye length, but is approximately 10 times larger than the Debye length. The dynamics of DNA polarization at high field strengths has been observed in the ns time range and is consistent with biased field induced ion dissociation. In the range of chain lengths from approximately 400 to approximately 850 base pairs DNA molecules exhibit permanent dipole moments, which are in a preferentially perpendicular direction to the end-to-end-vector, leading to a positive electric dichroism. These results are consistent with a "frozen" ensemble of bent DNA configurations and provide evidence for the existence of slow, non-elastic bending transitions. The electric parameters of proteins are usually dominated by a permanent anisotropy of the charge distribution, corresponding to permanent dipole moments of the order of several hundred Debye up to about 1500 Debye. Relatively small dipole moments of protein monomers add up to millions of Debye, when these proteins are in a vectorial organization in membrane patches, as found for bacteriorhodopsin and Na (+)K (+)-ATPase . In these cases the dipole vector may support vectorial ion transport. It is remarkable that the dipole moments of proteins usually show a relatively small dependence on the salt concentration; a rational for these observations is provided by a dipole potential at the plane of shear for rotational diffusion, which is defined in close analogy to the zeta-potential for translational diffusion. Symmetry breaking leading to huge electric dipole moments may be expected for mixed lipid vesicles: according to model calculations the phase separation of lipid components with and without net charges may lead to very high dipole moments; the expectation has been verified experimentally for vesicles containing DMPA and DMPC. The state of these systems should be extremely sensitive to electric fields. In summary, there is an unusual wide variation of electric parameters associated with biological macromolecules and with biomolecular assemblies, which is the basis for the complexity of different phenomena induced by electric fields in biological systems.