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The barrier can be restored with Nup98 containing either its own cohesive repeats or even unrelated cohesive repeats. Noncohesive or only Megestrol Acetate partially cohesive FG repeats, even when providing binding sites for NTRs, cannot build a proper barrier. This deficiency cannot be overcome by increasing the concentration of?the noncohesive FG domains within the pores. These findings?demonstrate that NPC selectivity cannot be explained by NTR?FG interactions alone. Instead, inter-FG cohesion appears to be equally important. The virtual gate model suggested that brushes of noninteracting FG domains are sufficient to exclude inert macromolecules from NPC passage (Rout et?al., 2003). The model relates to the fact that brushes (e.g., of PEG chains) can block macromolecules from binding to surfaces. Why then does a multivalently crosslinked FG hydrogel represent a far more efficient NPC barrier? We see several related explanations. First, a repellence of inert material by noninteracting linear polymers would come down to a volume exclusion effect. Even if one assumes the FG repeats to reach a local concentration of 200?mg/ml, they would exclude only ?1/5 of the available volume. In an FG hydrogel, however, the complete gel volume (comprising also the entrapped solvent) will exclude inert material larger than the mesh size. selleck kinase inhibitor Second, one could argue that noncohesive FG domains show an enhanced barrier effect when anchored (as brushes) to?a surface. Brushes, however, are effective obstacles only near their anchor points, where Brownian motion cannot displace them. This applies to distances from the grafting surface of no more than a persistence length, which is a measure of the polymer's stiffness. This length is Ruxolitinib order as high as 200?mg/ml, inert molecules would diffuse through such a ��noncohesive FG cloud�� at rates that are not fundamentally different from their diffusion rates in the cytoplasm. In other words, noncohesive FG brushes might hinder inert materials from an encounter with the grafting surface, but they are rather ineffective in suppressing NPC passage, which occurs mostly parallel to and in considerable distance (up to 20?nm) from the grafting surface. In contrast, meshes formed by inter-FG repeat cohesion pose effective barriers independently of any anchoring.