The dynamical stability of reverberatory neural circuits

The concept of reverberation proposed by Lorente de Nó and Hebb is key to understanding strongly recurrent cortical networks. In particular, synaptic reverberation is now viewed as a likely mechanism for the active maintenance of working memory in the prefrontal cortex. Theoretically, this has spurred a debate as to how such a potentially explosive mechanism can provide stable working-memory function given the synaptic and cellular mechanisms at play in the cerebral cortex. We present here new evidence for the participation of NMDA receptors in the stabilization of persistent delay activity in a biophysical network model of conductance-based neurons. We show that the stability of working-memory function, and the required NMDA/ AMPA ratio at recurrent excitatory synapses, depend on physiological properties of neurons and synaptic interactions, such as the time constants of excitation and inhibition, mutual inhibition between interneurons, differential NMDA receptor participation at excitatory projections to pyramidal neurons and interneurons, or the presence of slow intrinsic ion currents in pyramidal neurons. We review other mechanisms proposed to enhance the dynamical stability of synaptically generated attractor states of a reverberatory circuit. This recent work represents a necessary and significant step towards testing attractor network models by cortical electrophysiology.