Living cells display a remarkable capacity to compartmentalize their functional biochemistry spatially . A particularly fascinating example is the cell nucleus. Exchange of macromolecules between the nucleus and the surrounding cytoplasm does not involve crossing a lipid bilayer membrane. Instead, large protein channels known as nuclear pores cross the nuclear envelope and regulate the passage of other proteins and RNA molecules. Together with associated soluble proteins, the nuclear pores constitute an important transport system. Beyond simply gating diffusion, this system is able to generate substantial concentration gradients, at the energetic expense of guanosine triphosphate (GTP) hydrolysis.  Abstracting the biological paradigm, we examine this transport system as a thermodynamic machine of solution demixing. Building on the construct of free energy transduction and biochemical kinetics, we find conditions for stable operation and optimization of the concentration gradients as a function of dissipation in the form of entropy production . In contrast to conventional engineering approaches to demixing such as reverse osmosis, the biological system operates continuously, without application of cyclic changes in pressure or other intrinsic thermodynamic parameters .
Living cells display a remarkable capacity to compartmentalize their functional biochemistry . A particularly fascinating example is the cell nucleus. Exchange of macromolecules between the nucleus and the surrounding cytoplasm does not involve crossing a lipid bilayer membrane. Instead, large protein channels known as nuclear pores cross the nuclear envelope and regulate the passage of other proteins and RNA molecules. Beyond simply gating diffusion, the system of nuclear pores and associated transport receptors is able to generate substantial concentration gradients, at the energetic expense of guanosine triphosphate (GTP) hydrolysis. In contrast to conventional approaches to demixing such as reverse osmosis or dialysis, the biological system operates continuously, without application of cyclic changes in pressure or solution exchange. Abstracting the biological paradigm, we examine this transport system as a thermodynamic machine of solution demixing. Building on the construct of free energy transduction and biochemical kinetics, we find conditions for stable operation and optimization of the concentration gradients as a function of dissipation in the form of entropy production .
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spatially
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Together with associated soluble proteins, the nuclear pores constitute an important transport system.
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this system
the system of nuclear pores and associated transport receptors
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In contrast to conventional approaches to demixing such as reverse osmosis or dialysis, the biological system operates continuously, without application of cyclic changes in pressure or solution exchange.
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. In contrast to conventional engineering approaches to demixing such as reverse osmosis, the biological system operates continuously, without application of cyclic changes in pressure or other intrinsic thermodynamic parameters