Bauer Wolfgang Rudolf
Department of Internal Medicine I, University Hospital of Würzburg, Oberdürrbacher Straße 6, D-97080 Würzburg, Germany.
Comprehensive Heart Failure Centre, Am Schwarzenberg 15, A15, D-97080 Würzburg, Germany.
Entropy (Basel). 2020 Mar 25;22(4):376. doi: 10.3390/e22040376.
Understanding the function and control of channel transport is of paramount importance for cell physiology and nanotechnology. In particular, if several species are involved, the mechanisms of selectivity, competition, cooperation, pumping, and its modulation need to be understood. What lacks is a rigorous mathematical approach within the framework of stochastic thermodynamics, which explains the impact of interparticle in-channel interactions on the transport properties of the respective species. To achieve this, stochastic channel transport of two species is considered in a model, which different from mean field approaches, explicitly conserves the spatial correlation of the species within the channel by analysis of the stochastic dynamics within a state space, the elements of which are the channel's spatial occupation states. The interparticle interactions determine the stochastic transitions between these states. Local flow and entropy production in this state space reveal the respective particle flows through the channel and the intensity of the Brownian ratchet like rectifying forces, which these species exert mutually on each other, together with its thermodynamic effectiveness and costs. Perfect coupling of transport of the two species is realized by an attractive empty channel and strong repulsive forces between particles of the same species. This confines the state space to a subspace with circular topology, in which the concentration gradients as thermodynamic driving forces act in series, and channel flow of both species becomes equivalent. For opposing concentration gradients, this makes the species with the stronger gradient the driving, positive entropy producing one; the other is driven and produces negative entropy. Gradients equal in magnitude make all flows vanish, and thermodynamic equilibrium occurs. A differential interparticle interaction with less repulsive forces within particles of one species but maintenance of this interaction for the other species adds a bypass path to this circular subspace. On this path, which is not involved in coupling of the two species, a leak flow of the species with less repulsive interparticle interaction emerges, which is directed parallel to its concentration gradient and, hence, produces positive entropy here. Different from the situation with perfect coupling, appropriate strong opposing concentration gradients may simultaneously parallelize the flow of their respective species, which makes each species produce positive entropy. The rectifying potential of the species with the bypass option is diminished. This implies the existence of a gradient of the other species, above which its flow and gradient are parallel for any gradient of the less coupled species. The opposite holds for the less coupled species. Its flow may always be rectified and turned anti-parallel to its gradient by a sufficiently strong opposing gradient of the other one.
理解通道运输的功能和控制对于细胞生理学和纳米技术至关重要。特别是,如果涉及多个物种,则需要了解选择性、竞争、合作、泵送及其调节机制。目前缺乏的是一种在随机热力学框架内的严格数学方法,该方法能够解释通道内粒子间相互作用对各物种传输特性的影响。为了实现这一点,在一个模型中考虑了两种物种的随机通道运输,该模型不同于平均场方法,通过分析状态空间内的随机动力学来明确保留通道内物种的空间相关性,状态空间的元素是通道的空间占据状态。粒子间相互作用决定了这些状态之间的随机转变。该状态空间中的局部流和熵产生揭示了通过通道的各粒子流以及类似布朗棘轮的整流力的强度,这些物种相互施加这种力,以及其热力学效率和成本。通过有吸引力的空通道和相同物种粒子之间的强排斥力实现了两种物种运输的完美耦合。这将状态空间限制在一个具有圆形拓扑结构的子空间中,其中作为热力学驱动力的浓度梯度串联起作用,并且两种物种的通道流变得等效。对于相反的浓度梯度,这使得具有较强梯度的物种成为驱动物种,产生正熵;另一种则被驱动并产生负熵。大小相等的梯度会使所有流消失,从而达到热力学平衡。一种物种粒子间排斥力较小但另一种物种保持这种相互作用的微分粒子间相互作用为这个圆形子空间增加了一条旁路路径。在这条不参与两种物种耦合的路径上,出现了粒子间排斥力较小的物种的泄漏流,其方向与其浓度梯度平行,因此在这里产生正熵。与完美耦合的情况不同,适当的强相反浓度梯度可能会同时使各自物种的流平行,这使得每个物种都产生正熵。具有旁路选项的物种的整流潜力会降低。这意味着存在另一种物种的梯度,高于该梯度时,对于耦合较弱的物种的任何梯度,其流和梯度都是平行的。对于耦合较弱的物种则相反。通过另一种物种足够强的相反梯度,其流可能总是被整流并转向与其梯度反平行。