Bear: Neuroscience: Exploring the Brain. Chapter 4: The Action Potential
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1 Bear: Neuroscience: Exploring the Brain Chapter 4: The Action Potential
2 Introduction Action Potential in the Nervous System Conveys information over distances Cytosol has negative charge relative to extracellular space Neural code - frequency and pattern Action potential Spike Nerve impulse Discharge
3 Hodgkin and Huxley Giant squid axon - squid giant axon Most axons <1 micron, giant squid 1000 microns ( 1mm) Nobel prize 1963 Oscilloscope to visualize an AP
4 Properties of the Action Potential The Ups and Downs of an Action Potential Rising phase, overshoot, falling phase, and undershoot
5 APs - Na+ potentials threshold Generator potential first graded potentials all or none
6 APs - Na+ potentials Puncture foot, stretch membrane of nerve fibers Opens Na+-permeable channels Na+ influx depolarized membrane reaches threshold action potential threshold Generator potential first graded potentials all or none
7 Properties of the Action Potential The Generation of an Action Potential All-or-none : Cross threshold value for action potential Chain reaction threshold
8 What triggers other systems? Visual Auditory Taste Olfactory Motor Anxiety Depression Pleasure
9 Properties of the Action Potential The Generation of Multiple Action Potentials Artificially - inject current into a neuron using a microelectrode
10 Properties of the Action Potential The Generation of Multiple Action Potentials Firing frequency reflects the magnitude of the depolarizing current
11 H and H WERE RIGHT! 20 years PATCH CLAMP gigaseal (resistance HIGH) MOST CHANNELS ON OR OFF
12 The Action Potential, In Theory Depolarization (influx of Na + ) and repolarization (efflux of K + ) Membrane Currents and Conductances Current The net movement of K + across membrane Potassium channel number Proportional to electrical conductances Membrane potassium current Flow and driving force
13 currents
14 The Action Potential, In Theory
15 The Action Potential, In Theory The Ins and Outs of an Action Potential think of driving force difference between membrane potential and equilibrium potential Rising phase: Inward sodium current Falling phase: Outward potassium current
16 threshold LOTS of voltage gated NA+ open
17 rising phase large Na+ driving force overshoot approach E Na
18 Falling phase voltage gated Na+ channels close voltage gated K+ channels open K+ ions exit (big driving force)
19
20 Undershoot voltage gated K+ channels still open Na+ closed heads towards E K (-82 mv)
21 The Action Potential, In Reality The Generation of an Action Potential Hodgkin and Huxley voltage clamp Voltage Clamp: Clamp membrane potential at any chosen value Rising phase transient increase in g Na, influx of Na + ions Falling phase increase in g K, efflux of K + ions Existence of sodium gates in the axonal membrane
22 voltage gated sodium channels 4 domains 6 helices pore twists when voltage changes (charge of amino acids)
23 The Action Potential, In Reality The Voltage-Gated Sodium Channel (Cont d) Structure gating and pore selectivity
24 The Action Potential, In Reality The Voltage-Gated Sodium Channel Patch-clamp method
25 The Action Potential, In Reality The Voltage-Gated Sodium Channel (Cont d) Functional Properties of the Sodium Channel Open with little delay Stay open for about 1msec Cannot be open again by depolarization Absolute refractory period: Channels are inactivated
26 The Action Potential, In Reality The Voltage-Gated Sodium Channel (Cont d) In genetic disease channelopathies E.g., Generalized epilepsy with febrile seizures Toxins as experimental tools Puffer fish: Tetrodotoxin (TTX)- Clogs Na + permeable pore Red Tide: Saxitoxin- Na + Channel-blocking toxin
27 channelopathies Episodic ataxia (uncoordinated movement) Neonatal convulsions (why neonatal?) Startle disease Epilepsy, seizures
28 The Action Potential, In Reality The Voltage-Gated Sodium Channel (Cont d) Varieties of toxins Batrachotoxin (frog): Blocks inactivation so channels remain open Veratridine (lilies): Inactivates channels Aconitine (buttercups): Inactivates channels
29 The Action Potential, In Reality Voltage-Gated Potassium Channels Potassium vs. sodium gates Both open in response to depolarization Potassium gates open later than sodium gates Delayed rectifier Potassium conductance serves to rectify or reset membrane potential Structure: Four separate polypeptide subunits join to form a pore
30 The Action Potential, In Reality Key Properties of the Action Potential Threshold Rising phase Overshoot Falling phase Undershoot Absolute refractory period Relative refractory period
31 Depolarization Repolarization
32 Action Potential Conduction Initiation of synaptic transmission
33 conduction no decrement channels everywhere
34 Action Potential Conduction Propagation of the action potential Down axon to the axon terminal Orthodromic: Action potential travels in one direction Antidromic (experimental): Backward propagation Typical conduction velocity: 10 m/sec Length of action potential: 2 msec
35 action potentials AP travel: muscles 119 m/s touch m/s pain m/s thoughts m/s
36 Action Potential Conduction Factors Influencing Conduction Velocity Spread of action potential along membrane Dependent upon axon structure Path of the positive charge Inside of the axon (faster) Across the axonal membrane (slower) Axonal excitability Axonal diameter (bigger = faster) Number of voltage-gated channels
37 Action Potential Conduction Factors Influencing Conduction Velocity Myelin: Facilitates current flow Layers of myelin sheath Myelinating cells Schwann cells in the PNS Oligodendroglia in CNS
38 Action Potential Conduction Factors Influencing Conduction Velocity Saltatory conduction
39 MS
40 Myelogenesis - Glial cell development When are axons fully myelinated? myelinated = faster speeds
41 Action Potentials, Axons, and Dendrites Spike-initiation zone Sensory nerve endings Axon hillock
42 depolarization (rising and overshoot) repolarization (falling) hyperpolarization (undershoot) 2 ms
43 firing frequency (Hz) max 1000 why?
44 voltage gated sodium channels absolute refractory - Na+ open but either working or INACTIVATED relative refractory working, but K+ interferes depolarization Na+ enters repolarization K+ leaves
45
46 threshold LOTS of voltage gated NA+ open
47 rising phase large Na+ driving force
48 overshoot E Na 62mV
49 Falling phase voltage gated Na+ channels close voltage gated K+ channels open K+ ions exit (big driving force)
50 meanwhile back at the pump maintains concentration gradients heart stomach toxins
51 Undershoot voltage gated K+ channels still open Na+ closed heads towards E K (-82 mv) opening of delayed rectifier K+ channels
52 Rectifying channels Delayed 1 msec Lots of different types Studied w/ patch clamp
53 one-way?? orthodromic antidromic
54 size matters also -channel distribution also duct tape
55 channels NOT evenly distributed node of Ranvier How far does depolarizing wave go? path of least resistance
56 voltage gated channels
57 1. brief delay 2. open 1 ms, then close 3. open again Only at optimal voltage
58 TTX tetrodotoxin Binds to NA channel Pufferfish Red tide - saxitoxin in protozoa
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