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Dynamical Systems in Neuroscience:

Dynamical Systems in Neuroscience:

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Simple Models 309100080010 msrecovery variable, u600400200v-nullcl<strong>in</strong>e, I=550 pAv-nullcl<strong>in</strong>e, I=0 pAAHP5432112 3 4 5AHP0u-nullcl<strong>in</strong>espike-200-70 -60 -50 -40 -30 -20 -10 0 10 20membrane potential, v (mV)Figure 8.24: Phase portrait of the simple model <strong>in</strong> Fig. 8.23 exhibit<strong>in</strong>g CH fir<strong>in</strong>gpattern.current. There are some subtle differences: The response of an LTS cell to a weak depolariz<strong>in</strong>gcurrent consists of a phasic spike or a doublet with a relatively short latencyfollowed by low-frequency (less than 10 Hz) subthreshold oscillation of membrane potential.Stronger pulses elicit tonic spikes with slow frequency adaptation, decreas<strong>in</strong>gamplitudes and decreas<strong>in</strong>g after-hyperpolarizations, as one can see <strong>in</strong> Fig. 8.11.LTS neurons have more depolarized rest<strong>in</strong>g potentials, lower threshold potentials,and lower <strong>in</strong>put resistances than those of RS neurons. To match the <strong>in</strong> vitro fir<strong>in</strong>gpatterns of LTS <strong>in</strong>terneuron of rat’s barrel cortex <strong>in</strong> Fig. 8.25, we take the simple modelof RS neuron and adjust the rest<strong>in</strong>g and <strong>in</strong>stantaneous threshold potentials v r = −56mV and v t = −42 mV, and the values p = 1 and b = 8 result<strong>in</strong>g <strong>in</strong> the rheobasecurrent of 120 pA and the <strong>in</strong>put resistance of 50 MΩ. To model the decreas<strong>in</strong>g natureof the spike and AHP amplitudes, we assume that the peak of the spike and the afterspikeresett<strong>in</strong>g po<strong>in</strong>t depend on the value of the recovery variable u. This completelyunnecessary cosmetic adjustment has a mild effect on the quantitative behavior of themodel but renders a more “realistic” look to the simulated voltage traces <strong>in</strong> Fig. 8.25.The class of excitability of LTS neurons has not been studied systematically, thoughthe neurons seem to be able to fire periodic spike tra<strong>in</strong>s with a frequency as low as thatof RS neurons (Beierle<strong>in</strong> et al. 2003, Tateno and Rob<strong>in</strong>son, personal communication).The conjecture that they are near saddle-node on <strong>in</strong>variant circle bifurcation, andhence are Class 1 excitable <strong>in</strong>tegrators, seems to be at odds with the observation thattheir membrane potential exhibits slow damped oscillation and that they can fire post<strong>in</strong>hibitoryrebound spikes (Bacci et al. 2003), called low-threshold spikes (hence thename). They are better characterized as be<strong>in</strong>g at the transition from <strong>in</strong>tegrators to

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