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Living cells display a remarkable capacity to compartmentalize theirfunctional biochemistry. A particularly fascinating example is the cellnucleus. Exchange of macromolecules between the nucleus and the surroundingcytoplasm does not involve traversing a lipid bilayer membrane. Instead, largeprotein channels known as nuclear pores cross the nuclear envelope and regulatethe passage of other proteins and RNA molecules. Beyond simply gatingdiffusion, the system of nuclear pores and associated transport receptors isable to generate substantial concentration gradients, at the energetic expenseof guanosine triphosphate (GTP) hydrolysis. In contrast to conventionalapproaches to demixing such as reverse osmosis or dialysis, the biologicalsystem operates continuously, without application of cyclic changes in pressureor solvent exchange. Abstracting the biological paradigm, we examine thistransport system as a thermodynamic machine of solution demixing. Building onthe construct of free energy transduction and biochemical kinetics, we findconditions for stable operation and optimization of the concentration gradientsas a function of dissipation in the form of entropy production.

A thermodynamic paradigm for solution demixing inspired by nuclear transport in living cells