Biochemical networks in nervous systems: expanding neuronal information capacity beyond voltage signals

In addition to synaptically mediated signals that are based on changes in membrane potential, neurons also generate and receive many types of signals that involve biochemical pathways, some of which are independent of voltage. Although networks of biochemical pathways have often been thought of as being only neuromodulatory, recent computational and experimental studies have highlighted how these pathways can also integrate and transfer information themselves. Interactions between biochemical pathways involving positive and negative feedback loops allow biochemical signals to exhibit emergent properties, most notably bistability and oscillations. New and evolving techniques, including real-time imaging of second messengers, hold the promise of illuminating information processing that cannot be detected using microelectrodes, and revealing how 'biochemical integration' might contribute to the computational abilities of the nervous system.