No Arabic abstract
We examine the dynamics of biological nanomotors within a simple model of a rotor having three ion-binding sites. It is shown that in the presence of an external dc electric field in the plane of the rotor, the loading of the ion from the positive side of a membrane (rotor charging) provides a torque leading to the motor rotation. We derive equations for the proton populations of the sites and solve these equations numerically jointly with the Langevin-type equation for the rotor angle. Using parameters for biological systems, we demonstrate that the sequential loading and unloading of the sites lead to the unidirectional rotation of the motor. The previously unexplained phenomenon of fast direction-switching in the rotation of a bacterial flagellar motor can also be understood within our model.
We analyze the dynamics of rotary biomotors within a simple nano-electromechanical model, consisting of a stator part and a ring-shaped rotor having twelve proton-binding sites. This model is closely related to the membrane-embedded F$_0$ motor of adenosine triphosphate (ATP) synthase, which converts the energy of the transmembrane electrochemical gradient of protons into mechanical motion of the rotor. It is shown that the Coulomb coupling between the negative charge of the empty rotor site and the positive stator charge, located near the periplasmic proton-conducting channel (proton source), plays a dominant role in the torque-generating process. When approaching the source outlet, the rotor site has a proton energy level higher than the energy level of the site, located near the cytoplasmic channel (proton drain). In the first stage of this torque-generating process, the energy of the electrochemical potential is converted into potential energy of the proton-binding sites on the rotor. Afterwards, the tangential component of the Coulomb force produces a mechanical torque. We demonstrate that, at low temperatures, the loaded motor works in the shuttling regime where the energy of the electrochemical potential is consumed without producing any unidirectional rotation. The motor switches to the torque-generating regime at high temperatures, when the Brownian ratchet mechanism turns on. In the presence of a significant external torque, created by ATP hydrolysis, the system operates as a proton pump, which translocates protons against the transmembrane potential gradient. Here we focus on the F$_0$ motor, even though our analysis is applicable to the bacterial flagellar motor.
Shuttle-assisted charge transfer is pivotal for the efficient energy transduction from the food-stuff electrons to protons in the respiratory chain of animal cells and bacteria. The respiratory chain consists of four metalloprotein Complexes (I-IV) embedded in the inner membrane of a mitochondrion. Three of these complexes pump protons across the membrane, fuelled by the energy of food-stuff electrons. Despite extensive biochemical and biophysical studies, the physical mechanism of this proton pumping is still not well understood. Here we present a nanoelectromechanical model of the electron-driven proton pump related to the second loop of the respiratory chain, where a lipid-soluble ubiquinone molecule shuttles between the Complex I and Complex III, carrying two electrons and two protons. We show that the energy of electrons can be converted to the transmembrane proton potential gradient via the electrostatic interaction between electrons and protons on the shuttle. We find that the system can operate either as a proton pump, or, in the reverse regime, as an electron pump. For membranes with various viscosities, we demonstrate that the uphill proton current peaks near the body temperature $T approx 37 ^{circ}$C.
We propose a simple model of cytochrome c oxidase, including four redox centers and four protonable sites, to study the time evolution of electrostatically coupled electron and proton transfers initiated by the injection of a single electron into the enzyme. We derive a system of master equations for electron and proton state probabilities and show that an efficient pumping of protons across the membrane can be obtained for a reasonable set of parameters. All four experimentally observed kinetic phases appear naturally from our model. We also calculate the dependence of the pumping efficiency on the transmembrane voltage at different temperatures and discuss a possible mechanism of the redox-driven proton translocation.
Levy walks (LWs) are spatiotemporally coupled random-walk processes describing superdiffusive heat conduction in solids, propagation of light in disordered optical materials, motion of molecular motors in living cells, or motion of animals, humans, robots, and viruses. We here investigate a key feature of LWs, their response to an external harmonic potential. In this generic setting for confined motion we demonstrate that LWs equilibrate exponentially and may assume a bimodal stationary distribution. We also show that the stationary distribution has a horizontal slope next to a reflecting boundary placed at the origin, in contrast to correlated superdiffusive processes. Our results generalize LWs to confining forces and settle some long-standing puzzles around LWs.
Cilia and flagella often exhibit synchronized behavior; this includes phase locking, as seen in {it Chlamydomonas}, and metachronal wave formation in the respiratory cilia of higher organisms. Since the observations by Gray and Rothschild of phase synchrony of nearby swimming spermatozoa, it has been a working hypothesis that synchrony arises from hydrodynamic interactions between beating filaments. Recent work on the dynamics of physically separated pairs of flagella isolated from the multicellular alga {it Volvox} has shown that hydrodynamic coupling alone is sufficient to produce synchrony. However, the situation is more complex in unicellular organisms bearing few flagella. We show that flagella of {it Chlamydomonas} mutants deficient in filamentary connections between basal bodies display markedly different synchronization from the wild type. We perform micromanipulation on configurations of flagella and conclude that a mechanism, internal to the cell, must provide an additional flagellar coupling. In naturally occurring species with 4, 8, or even 16 flagella, we find diverse symmetries of basal body positioning and of the flagellar apparatus that are coincident with specific gaits of flagellar actuation, suggesting that it is a competition between intracellular coupling and hydrodynamic interactions that ultimately determines the precise form of flagellar coordination in unicellular algae.