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Crowded Charges in Ion Channels

Ions in water are the liquid of life. Life occurs almost entirely in 'salt water'. Water itself (without ions) is lethal to animal cells and damaging for most proteins. Water must contain the right ions in the right amounts if it is to sustain life. Physical chemistry is the language of electrolyte solutions. Physical chemistry and biology are intertwined. Physical chemists and biologists come from different traditions that separated for several decades as biologists described the molecules of life. Communication is not easy between a fundamentally descriptive tradition and a fundamentally analytical one. Biologists have now learned to study well defined systems with physical techniques, of considerable interest to physical chemists. Physical chemists are increasingly interested in spatially inhomogeneous systems with structures on the atomic scale so common in biology. Physical chemists will find it productive to work on well defined systems built by evolution to be reasonably robust, with input output relations insensitive to environmental insults. This article deals with properties of ion channels that in my view can be dealt with by 'physics as usual', with much the same tools that physical chemists apply to other systems. Indeed, I introduce and use a tool of physicists-a field theory (and boundary conditions) based on an energy variational approach developed by Chun Liu-not too widely used among physical chemists. My goal is to provide the knowledge base, and identify the assumptions, that biologists use in studying ion channels, avoiding jargon. Rather simple models of selectivity and permeation in ion channels work quite well in important cases. Those physical models and cases are the main focus of this review because they demonstrate the strong essential link between the traditional treatments of ions in chemical physics, and the biological function of ion channels.