neurons - university of edinburgh• communication between neurons is not typically a one-to-one...
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R Cheung MSc Bioelectronics: PGEE11106 1
Neurons
Pyramidal neurons in mouse cerebral cortex expressing green fluorescent protein.
The red staining indicates GABAergic interneurons.
MBL, Woods Hole
R Cheung MSc Bioelectronics: PGEE11106 2
Neuron
• The functional and structural unit of the nervous system
• There are many different types of neurons, but most have
certain structural and functional characteristics in common
Dendrite
Nucleus
Axon
Hillock
Axon
Myelin sheath
Schwann cell
Node of
Ranvier
Axon terminal
R Cheung MSc Bioelectronics: PGEE11106 3
Neurons are excitable cells specialized to conduct information from one part
of the body to another via electrical impulses conducted along their axons.
Function of Neurons
Medium: AXON
Message: ACTION POTENTIAL
Communication Model
Sender Receiver
R Cheung MSc Bioelectronics: PGEE11106 4
Membrane Potentials: Signals
• Neurons use changes in membrane potential to
receive, integrate, and send information.
• Two types of signals are produced by a change
in membrane potential:
– Graded potentials (short-distance)
– Action potentials (long-distance)
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-70 mV
Resting Axon Membrane Potential
The membrane potential is always given as the intracellular potential relative
to the extracellular potential - which is arbitrarily defined as zero.
Resting Neuron :
Membrane is polarized.
Inner, axoplasmic, side
is negatively charged.
All gated-sodium and
potassium channels are
closed.
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Graded Potentials
• Short-lived, local changes in membrane potential
• Currents decrease in magnitude with distance
• Their magnitude varies directly with the strength of the stimulus
• The stronger the stimulus the more the voltage changes, the farther
the current goes, and more likely to initiate action potentials
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Depolarization
stimulus
Spread of Depolarization
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Action Potentials (APs)
• Supra-threshold stimuli cause voltage-gated Na+
channels to open to produce depolarizing currents
• The AP is a brief reversal of membrane potential with a
total amplitude of ~100 mV (from -70mV to +30mV)
• APs do not decrease in strength with distance
All-or-Nothing - action potentials either
happen completely, or not at all
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• Appropriate stimulus applied to
the resting axon triggers nerve
impulse/action potential
• Membrane becomes negative
externally
Signals Carried by Neurons
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High [Na+]
High [K+]
Resting Membrane
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[Na+]
Depolarization and generation
of the nerve impulse
• At the leading edge of the
impulse, ‘fast’ sodium gates open.
The membrane becomes more
permeable to Na+ ions and an
action potential occurs.
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Signals Carried by Neurons
Propagation of the AP As the action potential passes,
slow potassium gates open,
allowing K+ ions to flow out. [Na+]
[K+]
+ - + + + + - -
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[Na+]
Repolarization
Depolarization
The action potential continues
to move along the axon in the
direction of the nerve impulse.
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The Dipole Field due to Current Flow in an Axon
at the Advancing Front of Depolarization.
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Hyperpolarization
• The ‘slow’ K+ gates remain open longer than needed to restore
the resting state.
• This excessive efflux causes hyperpolarization of the
membrane.
• The axon is insensitive to stimulus and depolarization during
this time.
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Role of the Sodium-Potassium Pump
• Repolarization restores the resting electrical
conditions of the axon, but does not restore the
resting ionic conditions.
• Ionic redistribution is accomplished by the sodium-
potassium pump following repolarization.
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Refractory Periods • Absolute Refractory Period: Time between opening and
closing of the Na+ activation gates. The axon cannot respond
to another stimulus.
• Relative Refractory Period: Follows the absolute refractory
period. Na+ gates are closed, K+ gates are open and
repolarization is occurring. Only a strong stimulus can
generate an AP.
stimulus
0
+30 mV
-70 mV
-55 mV
Absolute RP Relative RP
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Axon Conduction Velocities
• Conduction velocities vary widely among
neurons, determined mainly by:
– Axon Diameter - the larger the diameter, the faster
the impulse (less resistance)
– Presence of a Myelin Sheath – myelination
increases impulse speed (Continuous vs. Saltatory
Conduction)
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Saltatory Conduction
• Gaps in the myelin sheath between adjacent Schwann cells are called nodes of Ranvier.
• Voltage-gated Na+ channels are concentrated at these nodes.
• Action potentials are triggered only at the nodes and jump from one node to the next.
• Much faster than conduction along unmyelinated axons.
Axon Myelin sheath
Node of
Ranvier [Na+]
[Na+]
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• As the impulse reaches the axon terminals the signal is relayed
to target cells at specialized junctions known as synapses.
• Arrival of impulse at synapse opens Ca2+ channels.
• Neurotransmitter is released into the synaptic cleft via
exocytosis.
• Neurotransmitter crosses the synaptic cleft and binds to
receptors on the postsynaptic neuron.
• Postsynaptic membrane permeability changes due to opening
of ion channels, causing an excitatory or inhibitory effect.
Information Transfer at Synapse
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Synaptic Transmission
• An AP reaches the axon terminal of
the presynaptic cell and causes V-
gated Ca2+ channels to open.
• Ca2+ rushes in, binds to regulatory
proteins & initiates neurotransmittier
(NT) exocytosis.
• NTs diffuse across the synaptic cleft
and then bind to receptors on the
postsynaptic membrane and initiate
some sort of response on the
postsynaptic cell.
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Neurotransmitter Removal
• NTs are removed from the
synaptic cleft via:
– Enzymatic degradation
– Diffusion
– Reuptake
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• Different neurons can contain different NTs.
• Different postsynaptic cells may contain different receptors.
– Thus, the effects of an NT can vary.
• Some NTs cause cation channels to open, which results in a
graded depolarization.
• Some NTs cause anion channels to open, which results in a
graded hyperpolarization.
Effects of the Neurotransmitter
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• Typically, a single synaptic interaction will not create a graded
depolarization strong enough to migrate to the axon hillock and
induce the firing of an AP
– However, a graded depolarization will bring the membrane
potential closer to threshold. This is referred to as an
excitatory postsynaptic potential.
– Graded hyperpolarizations bring the membrane potential
farther away from threshold and thus are referred to as
inhibitory postsynaptic potentials.
• Whether a transmitter is excitatory or inhibitory depends on
its receptor.
Excitatory and Inhibitory Neurotransmitters
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• Acetylcholine is excitatory because its receptor is
a ligand-gated Na+ channel.
• GABA is inhibitory because its receptor is a
ligand-gated Cl- channel.
• Other transmitters (e.g. vasopressin, dopamine)
have G-protein-linked receptors.
– Effects depend on the signal transduction pathway and
cell type.
Excitatory and Inhibitory Neurotransmitters
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Temporal Summation
• One Excitory Postsynaptic Potential (EPSP) is usually not strong enough to cause an Action Potential. However, EPSPs may be summed:
• Temporal summation - the same presynaptic neuron stimulates the postsynaptic neuron multiple times in a brief period. The depolarization resulting from the combination of all the EPSPs may cause an AP.
• Spatial summation - multiple neurons all stimulate a postsynaptic neuron resulting in a combination of EPSPs which may yield an AP.
0
+30 mV
-70 mV
-55 mV
stimulii
EPSPs
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• Communication between neurons is not typically a one-to-one event.
– Sometimes a single neuron branches and its collaterals synapse on multiple target neurons. This is known as divergence.
– A single postsynaptic neuron may have synapses with as many as 10,000 presynaptic neurons. This is convergence.
Synaptic Organization
Divergence
Convergence
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Videos of Neurons in Action