Gap Junctions Allow Direct Communication Between Neurons and Between Glia
Chemical synapses employing neurotransmitters allow one-way communication at reasonably high speed. However, sometimes signals go from cell to cell electrically, without the intervention of chemical synapses. Electrical synapses depend on gap junction channels that link two cells together (Chapter 20). The effect of gap junction connections is to perfectly coordinate the activities of joined cells. An electrical synapse is usually bidirectional; either neuron can excite the other. Electrical synapses are common in the neocortex and thalamus, for example. The key feature of electrical synapses is their speed. While it takes about 0.5–5 ms for a signal to cross a chemical synapse, transmission across an electrical synapse is almost instantaneous, on the order of a fraction of a millisecond since the cytoplasm is continuous between the cells. In addition, the presynaptic cell (the one sending the signal) does not have to reach a threshold at which it can cause an action potential in the postsynaptic cell. Instead, any electrical current continues into the next cell and causes depolarization in proportion to the current.
Gap junctions form between glial cells as well as between neurons. Astrocytes in the brain are connected to one another through gap junctions, which gives rise to the generation of waves of Ca2+ that propagate through networks of astrocytes in the brain at a speed of 1 µm/sec. Gap junctions also form within individual Schwann cells, forming connections between the layers of myelin formed by a single Schwann cell. These gap junctions are thought to facilitate the passage of metabolites and ions between myelin layers.
An electrical synapse may contain thousands of gap channels, each composed of two hemichannels, one in each apposed cell. Gap junction channels in the neuron have a structure similar to conventional gap junctions (see Figure 20-20). Each hemichannel is an assembly of six copies of the connexin protein. Since there are about 20 mammalian connexin genes, diversity in channel structure and function can arise from the different protein components. The 1.6–2.0-nm channel itself allows the diffusion of molecules up to about 1000 Da in size and has no trouble at all accommodating ions.