Membrane and circuit properties of lateral septum neurons: relationships with hippocampal rhythms

The lateral septum receives the most important afferents from the hippocampus, has been proposed to contribute to theta (theta) rhythm generation. Our aim was to study the membrane and circuital properties of lateral septum neurons and their relationship with hippocampal rhythms. Extra- and intracellular recordings (n=81) were obtained in urethane-anesthetized rats. Two neuronal populations were found, one of them with hippocampal theta; rhythm dependence (theta-D, 68%), and the other group independent of hippocampal theta; rhythm (theta-I, 32%). Other differences were spontaneous firing rate (theta-D=13.20+/-2.09, theta-I=6.99+/-1.18; p<0.005) with a bursting pattern in the theta-D group and single discharges in the theta-I group. Intracellular recordings showed higher synaptic activity in theta-D than in theta-I neurons. Both groups showed fast spikes while only theta-D neurons had high-threshold and low-threshold slow spikes. theta-D neurons had theta-oscillations in their membrane potential. Stimulation of the contralateral lateral septum resets the hippocampal theta rhythm and the theta rhythm recorded intracellularly in the lateral septum neurons. Some neurons (10.6%) showed rhythmic oscillations lasting a few seconds, at a higher frequency than those of the simultaneously recorded hippocampal EEG. This high frequency appeared spontaneously or could be evoked by stimulations of the fornix and reticularis pontis oralis nucleus (RPO). Homologous high frequency oscillations appeared in the simultaneously recorded hippocampal EEG, synchronized with the neuronal activity, during fornix stimulation. RPO stimulation evoked increments (57%) or decrements (43%) in the firing rate of lateral septum neurons. Thus, they could participate in different septal networks to modulate the theta rhythm. The marked functional relationship between lateral septum neurons and hippocampal theta rhythm supports the proposal that lateral septum represents a feedback system for the improvement of medial septum theta rhythm. The hypothalamic projections could be a way to introduce theta and higher rhythms into this structure that control many biological rhythms.