Figure 1. Delayed increase in intrinsic excitability following muscarinic receptor activation by endogenous acetylcholine.
(A) Diagram of experiment protocol; orange bar indicates timing of light pulses that trigger ACh release via ChR2. (B) Example step responses in control conditions (black trace) and when ACh was released 2 s before and during the step response (orange trace). Blue arrowhead indicates timing of first AP in the control step response that was accelerated when ACh was released (Quantified in G). (C) Blockade of muscarinic cholinergic receptors with 10 M atropine abolished the increase in excitability triggered by ACh release. Black traces acquired in control conditions; orange traces illustrate responses with coincident light stimulation to the same depolarizing step. (D) Summary of the average number of APs evoked by depolarizing steps in five experiments similar to C. *p=0.0106, T = 4.526; n.s., p=0.847, T = 0.206. paired t-test. (E) Example trace showing that ACh release in the absence of a depolarizing step does not modulate the membrane potential. Inset, in the same cell, pairing ACh release with depolarizing step increases the number of APs triggered. (F) Comparison of mean membrane potential in control conditions (black bar) and after different durations of light pulse trains only (orange bars; p>0.05; paired t-test). Experiments conducted on neurons in which coincident light trains increased the number of APs triggered by depolarizing steps. (G) Summary plot of the average time during similar step responses before an AP was accelerated (shorter latency for the Nth AP when ACh was released, compare B for example traces). (H) Example illustrating different light/step timing protocols. Response shown (orange trace) is from ‘post’ protocol. (I) Coincident ACh release is required to increase neuronal excitability. Plot of the number of APs evoked the same depolarizing step in control conditions (black bar) and when a 6 s light pulse train was applied 7 s before (Pre), 2 s before (Coinc) or 2 s after (Post) the depolarizing step. **p=0.00334, T = 5.8373; paired t-test, Bonferroni corrected. (J) Coincident ACh release and depolarizing step stimuli reveals an afterdepolarization potential (ADP; red arrowhead) that is abolished by 10 M atropine. Action potentials truncated. (K) Summary plot of the dependence of the ADP on coincident ACh release. Same light train/depolarizing step timing as G. *p=0.0219, T = 3.977; paired t-test, Bonferroni corrected. Plots are mean ± SEM unless noted throughout study.