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Dynamical Systems in Neuroscience:

Dynamical Systems in Neuroscience:

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Introduction 510 ms10 mV-45 mV0 pA-100 pAFigure 1.6: In vitro record<strong>in</strong>g of rebound spikes ofrats bra<strong>in</strong>stem mesV neuron <strong>in</strong> response to a briefhyperpolariz<strong>in</strong>g pulse of current.10ms5msnon-resonant burst10msresonant burst15msnon-resonant burst<strong>in</strong>hibitory burstFigure 1.7: Resonant response of the mesencephalic V neuron of rat bra<strong>in</strong>stem to pulsesof <strong>in</strong>jected current hav<strong>in</strong>g 10 ms period (<strong>in</strong> vitro).the threshold, then how can the neuron <strong>in</strong> Fig. 1.6 fire <strong>in</strong> response to the <strong>in</strong>hibitory<strong>in</strong>put? This phenomenon is also observed <strong>in</strong> the Hodgk<strong>in</strong>-Huxley model, and it iscalled anodal break excitation, rebound spike, or post-<strong>in</strong>hibitory spike. Many biologistssay that rebound responses are due to the activation and <strong>in</strong>activation of certa<strong>in</strong> slowcurrents, which br<strong>in</strong>g the membrane potential over the threshold, or equivalently, lowerthe threshold upon release from the hyperpolarization – a phenomenon called a lowthresholdspike <strong>in</strong> thalamocortical neurons. The problem with this explanation is thatneither the Hodgk<strong>in</strong>-Huxley model nor the neuron <strong>in</strong> the figure have these currents,and even if they did, the hyperpolarization is too short and too weak to affect thecurrents.Another <strong>in</strong>terest<strong>in</strong>g phenomenon is depicted <strong>in</strong> Fig. 1.7. The neuron is stimulatedwith brief pulses of current mimick<strong>in</strong>g an <strong>in</strong>com<strong>in</strong>g burst of three spikes. When thestimulation frequency is high (5 ms period), presumably reflect<strong>in</strong>g a strong <strong>in</strong>put,the neuron does not fire at all. However, stimulation with a lower frequency (10ms period) that resonates with the frequency of subthreshold oscillation of the neuronevokes a spike response, regardless of whether the stimulation is excitatory or <strong>in</strong>hibitory.Stimulation with even lower frequency (15 ms period) cannot elicit spike response aga<strong>in</strong>.Thus, the neuron is sensitive only to the <strong>in</strong>puts hav<strong>in</strong>g resonant frequency. The samepulses applied to a cortical pyramidal neuron evoke a response only <strong>in</strong> the first case(small period), but not <strong>in</strong> the other cases.

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