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Dendrites are the main recipients of synaptic inputs and are important sites that determine neurons' input-output functions. This review focuses on thin neocortical dendrites, which receive the vast majority of synaptic inputs in cortex but also have specialized electrogenic properties. We present a simplified working-model biophysical scheme of pyramidal neurons that attempts to capture the essence of their dendritic function, including the ability to behave under plausible conditions as dynamic computational subunits. We emphasize the electrogenic capabilities of NMDA receptors (NMDARs) because these transmitter-gated channels seem to provide the major nonlinear depolarizing drive in thin dendrites, even allowing full-blown NMDA spikes. We show how apparent discrepancies in experimental findings can be reconciled and discuss the current status of dendritic spikes in vivo; a dominant NMDAR contribution would indicate that the input-output relations of thin dendrites are dynamically set by network activity and cannot be fully predicted by purely reductionist approaches.
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Download all Supplemental Material as one PDF. Includes links to Supplemental Videos 1-4 (videos embedded below), Supplemental Figures 1-6, and a Supplemental Discussion. Videos show time evolution of I-V curves and membrane potential (Vm) of models, following a brief pulse of glutamate, with AMPAR conductances yielding just subthreshold and just-suprathreshold responses. Pale orange line in top panel and X in bottom right panel indicate threshold when bistable state exists. Download video files to view with QuickTime Player, dragging slider back and forth to vary speed and to replay key portions such as when threshold declines until it sinks below voltage (so technically, in this situation, voltage doesn’t cross above threshold, rather, threshold crosses below voltage). In some cases, the threshold (orange line and X) first appears transiently below Vm, and then rapidly rises above Vm as the AMPAR conductance continues to fall; the net current is too small to charge the membrane capacitance fast enough to move the membrane into the up-state at that point (e.g. Supplemental Videos 2, 4). Matlab code available on request. Supplemental Videos 1 and 2. Highest gmax NMDA = 5 × leak (as in Figure 3b, dark green curve or Supplemental Figure 4c). Supplemental Video 1: gmax AMPA = 2.1 × leak (subthreshold). Download video file (MPG) Supplemental Video 2 : gmax AMPA = 2.2 × leak (suprathreshold). Download video file (MPG) Would see small spike "riding on" big subthreshold response (if superimposed). Supplemental Videos 3 and 4. Highest gmax NMDA = 6 × leak (as in Figure 3a, green curve, or Supplemental Figure 4d). Supplemental Video 3: gmax AMPA = 1.37 × leak (subthreshold). Download video file (MPG) Supplemental Video 4 : gmax AMPA = 1.38 × leak (suprathreshold). Download video file (MPG) Would see medium spike "riding on" medium subthreshold response (if superimposed).