Distributed effects of dopamine modulation in the crustacean pyloric network.

It is now clear that neuromodulators can reconfigure a single motor network to allow the generation of a family of related movements. Using dopamine modulation of the 14-neuron pyloric network from the crustacean stomatogastric ganglion as an example, we describe two major mechanisms by which network output is modulated. First, the baseline electrophysiological properties of the network neurons can be altered. Dopamine can affect the activity of each neuron independently. For example, DA modulates IA in nearly every neuron in the pyloric network, but in opposite directions in different cells. Furthermore, DA usually modulates combinations of ionic currents. In some cases, currents with opposing actions on cell excitability are simultaneously affected, and the net response reflects the sum of these opposing effects. Second, neuromodulators can alter the strength of synaptic interactions within the network, quantitatively "rewiring" the network. Every synapse in the network is affected by DA, with some increased and others decreased in strength. DA acts both pre- and postsynaptically to affect transmission: these actions are frequently opposing in sign, and the net response arises as the sum of these opposing actions. Finally, spike-evoked and graded transmission at the same synapse can be oppositely affected by DA. These results emphasize the distributed nature of modulation in motor networks.