Crowding effects of membrane proteins.

In cell membranes, membrane proteins occupy approximately 30% of the total surface area. Crowding effects similar to those in the solution phase are thus to be expected. In addition, there are crowding effects unique to proteins bound to the two-dimensional membranes, such as those exerted on the equilibration of a protein between two membrane orientations and on the redistribution of proteins between different locations in a cell membrane. This article aims to present a theoretical framework for understanding the various crowding effects within membranes. For illustration, the theory is used to analyze previously published experimental and simulation data. It is hoped that the article will encourage quantitative analyses in future experiments and spur systematic investigation of membrane crowding effects.