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Action Potential & Propagation
DENT/OBHS 131 Neuroscience 2009
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Ionic basis of APs action potential:
faithfully transmit information along the membrane (axon) of excitable cells allow rapid communication between distant parts of a neuron
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Learning Objectives Describe the roles of both sodium and potassium ions / voltage-gated channels before, during and after the action potential Understand how the resistive & capacitive properties of neurons influence electrical signaling Compare and contrast local circuit and saltatory propagation of action potentials
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How many distinct ion channels are necessary for the AP?
1 2 3 4 5
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3 phases of the action potential
Resting i.e. RMP Depolarization reversal of membrane potential Repolarization return of membrane potential to RMP
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General rule relationship between:
ENa +67 membrane potential (mV) relationship between: membrane potential ion equilibrium potentials if the membrane becomes more permeable to one ion over other ions then the membrane potential will move towards the equilibrium potential for that ion (basis of AP) RMP ECl -90 EK -98
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Depolarization rapid opening of Na-selective channels
entry of Na “down” its electrochemical gradient 1. membrane more permeable to Na than K 2. membrane potential moves (rapidly) towards ENa 3. because ENa is positive, the AP overshoots zero 4. At the peak of the AP Na is the primary ion determining the membrane potential
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Repolarization closure (inactivation) of Na-selective channels
slower opening of K-selective channel 1. membrane more permeable to K than Na 2. K moves out of cell 3. membrane potential moves towards EK
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2 independent channels selective agents block the 2 components
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Voltage-gated ion channels
the opening and closing of AP Na and K channels are controlled by changes in the membrane potential
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What triggers an AP? all-or-none threshold
AP are not graded potentials threshold in order for an AP to occur the membrane must be depolarized beyond a threshold level inward Na overcomes resting outward K movement electrical stimulation synaptic activation
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APs are regenerative activation of Na channels is cyclical
initial depolarization opening of Na channels Na entry etc..
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Learning Objective #1 Describe the roles of both sodium and potassium ions / voltage-gated channels before, during and after the action potential
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AP review
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Learning Objective #2 Understand how the resistive & capacitive properties of neurons influence electrical signaling
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How does an AP move? Propagation
Aps are conducted along excitable cell membranes away from their point of origin e.g. down the axon from cell soma to terminal
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Resistance ≈ how far it can get
membrane resistance (rm) axon / dendrite diameter (d) axial, or internal, resistance (ri) rm ri ength constant = “leaky pipe”
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Fat axons are fastest!
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Capacitance ≈speed “bulk” solutions IN and OUT are neutral
the transmembrane potential difference exists within a narrow band just across the membrane a capacitor separates / stores charge to change membrane potential must add or remove charge this takes time
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Summary Capacitance - speed (time constant)
Resistance - distance (length constant) How does neuron deal with these properties in order to have efficient AP propagation?
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Unmyelinated axons local circuit propagation slow
of the membrane during the AP is not restricted to a single spot the inward current carried by Na ions during the AP depolarizes adjacent portions of the membrane beyond threshold and the regenerative AP travels along the membrane
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Refractory period following a single AP a second AP cannot be generated at the same site for some time (absolute versus relative) Na channels need to recover from inactivation open K channels oppose inward Na movement
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Myelination local circuit propagation is slow (< 2 m/s)
In motor neurons propagation is fast 100 m/s Schwann cell / oligodendrocyte envelop axons / layer of insulation increase membrane resistance less leaky eliminate capacitance less discharge Nodes of Ranvier discontinuity in myelin sheath (every few 200+ m)
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Saltatory conduction APs are only generated at Nodes of Ranvier
high density of Na / K channels current flows rapidly between nodes little current leakage between nodes AP “jumps” down fiber as successive nodal membrane capacitances are discharged
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Learning Objective #3 Compare and contrast local circuit and saltatory propagation of action potentials
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propagation review Press button
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How can AP rise so fast (< 1 ms)?
m= rmcm
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Membrane time constant
changing the membrane potential takes time charging a capacitor is not instantaneous inject current V record voltage I m = rmcm ≈ 50 ms axon/dendrite
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