Current Fluctuations in Biological Transport Systems Far from Equilibrium

Abstract

A new theoretical approach to transport fluctuations around stable steady states in discrete biological transport systems [1],[2],[3] is used for the investigation of general fluctuation properties at nonequilibrium. An expression for the complex frequency dependent admittance at nonequilibrium is derived by calculation of the, linear current response of the transport systems to small disturbances in the applied external voltage. It is shown that the Nyquist or fluctuation dissipation theorem, by which at equilibrium the macroscopic admittance or linear response can be expressed in terms of fluctuation properties of the system, breaks down at nonequilibrium. The spectral density of current fluctuations is decomposed into one term containing the macroscopic admittance and a second term which is bilinear in current. This second term is generated by microscopic disturbances, which cannot be excited by external macroscopic perturbations. At special examples it is demonstrated that this second term is decisive for the occurrence of excess noise e.g. the 1/f2-Lorentzian noise generated by the opening and closing of nerve channels in biological membranes. Furthermore the quadratic dependence on current (e.g. 1/f noise) comes out to be typical of excess noise.