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1Molecular Biology and Physics, Princeton University, Princeton, New Jersey; 2Physiology, Technion Medical School, Haifa, Israel; and 3Max-Planck Institut für Medizinische Forschung, Heidelberg, Germany
Submitted 4 January 2008; accepted in final form 5 March 2008
Glutamatergic inputs clustered over
20–40 µm can elicit local N-methyl-D-aspartate (NMDA) spike/plateau potentials in terminal dendrites of cortical pyramidal neurons, inspiring the notion that a single terminal dendrite can function as a decision-making computational subunit. A typical terminal basal dendrite is
100–200 µm long: could it function as multiple decision-making subunits? We test this by sequential focal stimulation of multiple sites along terminal basal dendrites of layer 5 pyramidal neurons in rat somatosensory cortical brain slices, using iontophoresis or uncaging of brief glutamate pulses. There was an approximately sevenfold spatial gradient in average spike/plateau amplitude measured at the soma, from
3 mV for distal inputs to
23 mV for proximal inputs. Spike/plateaus were NMDA receptor (NMDAR) conductance-dominated at all locations. Large Ca2+ transients accompanied spike/plateaus over a
10- to 40-µm zone around the input site; smaller Ca2+ transients extended approximately uniformly to the dendritic tip. Spike/plateau duration grew with increasing glutamate and depolarization; high Ca2+ zone size grew with spike/plateau duration. The minimum high Ca2+ zone half-width (just above NMDA spike threshold) increased from distal (
10 µm) to proximal locations (
25 µm), as did the NMDA spike glutamate threshold. Depolarization reduced glutamate thresholds. Simulations exploring multi-site interactions based on this demonstrate that if appropriately timed and localized inputs occur in vivo, a single basal dendrite could correspond to a cascade of multiple co-operating dynamic decision-making subunits able to retain information for hundreds of milliseconds, with increasing influence on neural output from distal to proximal. Dendritic NMDA spike/plateaus are thus well-suited to support graded persistent firing.
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