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Gap junctions are aggregates of microscopic channels that connect the cytoplasmic compartments of the coupled cells. A gap junction channel is formed of two hemichannels, one contributed by each cell. Hemichannels can be formed in a post-Golgi compartment and transported to the cell surface in vesicles that fuse with the external membrane (Musil and Goodenough, 1993). Hemichannels then find a hemichannel in an apposed membrane and dock to form a cell–cell channel. For a long time, hemichannels were thought not to open to the exterior and to open only when joined to a hemichannel from an apposed membrane. Electrophysiological measurements showed that the hemichannels forming the first cell–cell channel had not been open to the medium before the channel formed (Bukauskas and Weingart, 1993). A teleological argument was that if hemichannels had the same large diameter pore as in a cell channel, they would have provided a deleterious link with the external medium, allowing loss of K+, ATP, and other metabolites and influx of Ca2+ and Na+. Now it is clear that isolated, unapposed hemichannels can open under specific physiological as well as pathological conditions, and metabolites coming out through hemichannels can mediate paracrine as well as autocrine signaling (Saez et al., 2010). The open probability is generally low, and increased opening can contribute to cell death, as presumed in arguing that the hemichannels should not open.

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