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Chapter 5:
Inside Outside
K +
ATP Na +
Na+
K+
3 Na +
ADP 2 K+
Inside Outside
K + K+
Na+
Na +
Inside Outside
… how can passive
diffusion of
potassium and
K + K+
sodium lead to
development of
negative membrane
potential?
Na+
Na +
Ki
EK = −61 × log
Ko
Nai
ENa = -61 x log
Nao
Vm −90 to −70
0 mV
Carbonyl oxygens
Figure 5-4
+ + −
+ +
p' K [ K ]o p' Na [ Na ]o p' Cl [Cl ]i
=
Vm 61. log + +
p' K [ K ]i + p' Na [ Na ]i + p' Cl [Cl − −]o-
or
+ +
p' K [ K ]i p' Na [ Na ]i + p' Cl [Cl − ]o
+
Vm = −61. log + +
p' K [ K ]o + p' Na [ Na ]o + p' Cl [Cl − ]i
Vm = membrane potential
P’ = permeability
Figure. 5-5
Normal conditions
Vm −74 ENa+61
EK −94
0
mV
20 mV 135 mV
Vm ENa+61
EK −94
0
mV
0 mV 0 mV
Threshold
Threshold
11 mV
15 mV
−90 mV −90 mV
Inactivation
Gate
Ca++ ions
What is the effect of low plasma
calcium on threshold potential? ++
++ ++
++
Outside
ANS: Threshold potential + + + + + Voltage
Selectivity + Sensor
becomes more negative,
filter +
and thus closer to Vm. This
increases the excitability of +
the cell. +
_ _ _ _ +
Inside Gate
Inactivation
Gate
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC1413606/
Figure 5-2
Figure 5-6
Stimulus
muscle by extrinsic (percutaneous) stimulation.
Have constant amplitude
APs do not summate - information is coded by Time
frequency not amplitude.
Have constant conduction velocity 75
Myelinated
Velocity (m/s)
True for given fiber. Fibers with large diameter (cat)
50
conduct faster than small fibers. As a general
rule: 25 Nonmyelinated
Myelinated fiber diameter (in mm) × 4.5 = (squid)
velocity in m/s. 0
0 3 6 9 12 15
Square root of unmyelinated fiber diameter
= velocity in m/s
0 400 800
Fiber diameter (mm)
Copyright © 2021 by Saunders, an imprint of Elsevier Inc.
Functions of Action
Potentials
• Deliver sensory information to CNS
– APs in sensory nerves are blocked by local anesthetics. This usually produces
analgesia without paralysis. Why no paralysis? LAs are more effective against
small diameter neurons with a large surface area to volume ratio. Hence, small C-
fibers that conduct pain sensations are affected more than large, alpha-
motorneurons.
• Information encoding
– The frequency of APs encodes information (amplitude of AP is constant).
• Rapid transmission over distance (nerve cell APs)
– The speed of transmission depends on fiber size and whether it is myelinated.
Information of lesser importance is carried by slowly conducting unmyelinated
fibers (nonmyelinated c-fibers conduct pain sensations).
• In non-nervous tissues, APs initiate various cellular responses.
– muscle contraction
– secretion (e.g., Epinephrine from chromaffin cells of medulla)
Dow
Na permeability increases
ke
due to opening of Na+ channels
nstr
0
Upstro
memb. potential approaches ENa
oke
(mV)
• During downstroke of action potential:
Resting potential
Na permeability decreases
-90
due to inactivation of Na+ channels
EK
1 ms Membrane
n
• Ion channel - properties ol
tio
ar
iza
iza
lar
Have conducting states and non- tio
po
n
conducting states
De
Transition between states = “gating”
Inactivation
Open Inactivated
• Channels “gate” in response to:
Changes in membrane potential (usually depolarization)
voltage-gated channels. Action potential propagation relies on voltage-gated channels
Occupation of receptor
ligand-gated or receptor operated channels (ROCs). These initiate action potentials
Mechanical forces
mechanosensitive channels - important for hearing
open channels
Inactivated channels do not pass any ions
(nonconducting state).
Number of
By contrast, K channels show activation but
not inactivation.
• The fall in current at the end is K+ channels
deactivation (opposite of activation).
closed
K+ channels
tio
lar
iza
iza
ar
tio
ol
n
p
de
inactivation
open inactivated
deactivation
Figure 5-9
2
3 1 3
2 2
1 2
1
2
3
1
Figure 5-7
Figure 5-10
Copyright © 2021 by Saunders, an imprint of Elsevier Inc.
Voltage Gated Na+ Channels Have
a Selectivity Filter
Figure 5-7
Figure 5-10
d
+ + -
e
----------------++++
• Why does the bipolar action potential +++++++++++++-----
Direction
look like this?
Copyright © 2021 by Saunders, an imprint of Elsevier Inc.
Conduction Velocity of AP (1 of 2)
mV
0
Threshold
-40
-80
0 1 2 3 4 5 msec
ARP RRP
Absolute refractory period—AP not possible due to voltage inactivation
of Na channels
Relative refractory period—greater than normal stimulus required to
elicit AP
Refractory periods limit the maximum frequency of Aps.
Copyright © 2021 by Saunders, an imprint of Elsevier Inc.
Propagation of Action Potential
Opening of Na+ channels generates local current that depolarizes adjacent
membrane, opening more Na+ channels …
Rest
Stimulated
(local depolarization)
Propagation
(current spread)
Figure 5-11
Myelination
• Schwann cells surround the nerve
axon forming a myelin sheath.
Figure 5-16
A, Modified from Leeson TS, Leeson R: Histology. Philadelphia: WB Saunders, 1979
Figure 5-17
Nonmyelinated vs myelinated
Nonmyelinated
Myelinated
• Point of communication
between neurones
Dendrite
Axo-dendritic
• Most synapses involve synapse
transmitter substances. Myellin
mV
• epsp’s and ipsp’s result from increases Presynaptic neuron
in membrane permeability. -70
So, ask yourself:
– to which ions? You need to
-60
consider the equilibrium mV Postsynaptic neuron
potentials. -70
ipsp
IPSP: K or Cl -80
10 ms
[Ca2+]
+++
Ca2+
+++ Na+
Na+,
K+, Ca++
V-G Na+
channels
V-G Ca2+ tx release open APpost
channels tx-R
Diffuse
APpre open
<20 nm>
[Ca]
Depolarization
0.5 ms
Inhibitory
Examples—GABAA, Glycine Inhibitory postsynaptic
– Permeable to anions (Cl-) potential (ipsp)
– Equilibrium potential ~ -90 mV
– Hyperpolarizes post-synaptic cell
– Depresses excitability
Excitatory
Examples—nAChR, Glutamate Excitatory postsynaptic
– Permeable to cations (Na+, K+, and Ca2+) potential (epsp)
– Equilibrium potential ~ 0 mV
– Depolarizes postsynaptic cell
– Enhances excitability
Overshoot
0 mV
Excitability
+
Repolarization
Threshold
Depolarization Resting
-90 mV
potential
Hyperpolarization
Mv •0 Phase 0 (depolarization)
0 − inward Ca + + current
-80
—Parasympathetic mechanism
Mv
0
-20 time
-40
-60
-80
Increasing the permeability to
which ion would cause this
effect?
ANS: K+ Annu. Rev. Physiol. 2001. 63:235–57
Copyright © 2021 by Saunders, an imprint of Elsevier Inc.
SA Node Action Potential (2 of 2)
—Sympathetic mechanism
Mv
0
-20 Time
-40
-60