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C HAPTE R

Neuron Structure
and Function

Active Lecture Question Slides prepared by


Dr. Alan F. Smith, Mercer University
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Neurons
Vary in structure and properties
Use same basic mechanisms to send signals

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Figure 4.1

Neural Zones
Four functional zones
Signal reception
Dendrites and the cell body (soma)
Incoming signal received and converted to change in
membrane potential

Signal integration
Axon hillock
Strong signal is converted to an action potential (AP)

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Neural Zones
Signal conduction
Axon (some wrapped in myelin sheath)
AP travels down axon

Signal transmission
Axon terminals
Release of neurotransmitter

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Neural Zones

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Figure 4.2

Electrical Signals in Neurons


Neurons have a resting membrane potential (like all
cells)
Membrane potential is negative at rest

Neurons are excitable


Can rapidly change their membrane potential
Depolarization membrane potential becomes less
negative
Repolarization membrane potential returns to resting
value
Hyperpolarization membrane potential becomes more
negative than resting value

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Electrical Signals in Neurons


Changes in membrane potential act as electrical
signals

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Changes in Membrane Potential

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Figure 4.3

Membrane Potential
Factors contributing to membrane potential
Distribution of ions across the membrane
Relative permeability of the ions
Charges of the ions

Goldman equation for the calculation of membrane


potential (Em)

RT PK [ K ]o P Na [ Na ]o P Cl [Cl ]i
Em
ln

F
PK [ K ]i PNa [ Na ]i PCl [Cl ]o
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The Goldman Equation

P
[
K
]

P
[
Na
]

P
[
Cl
]i
RT
K
o
Na
o
Cl
Em
ln

F
PK [ K ]i PNa [ Na ]i PCl [Cl ]o

Em = membrane potential
R = gas constant
T = temperature (Kelvin)
F = Faradays constant
Px = relative permeability of ion
[X] = ion concentration outside or inside membrane
Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

The Goldman Equation

P
[
K
]

P
[
Na
]

P
[
Cl
]i
RT
K
o
Na
o
Cl
Em
ln

F
PK [ K ]i PNa [ Na ]i PCl [Cl ]o

Other ions (Ca++, Mg++, etc.) are ignored in this


simplified form of the equation because their
permeabilities are very low

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Gated Ion Channels


Neurons depolarize or hyperpolarize by selectively
altering permeability
Gated ion channels open or close in response to a
stimulus
Example: neurotransmitter

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Gated Ion Channels


Channels only allow specific ions to pass through
the membrane
Ion moves down its electrochemical gradient
Only relatively small numbers of ions move across

As permeability to a specific ion increases,


membrane potential will approach that ions
equilibrium potential (Nernst equation)

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Changes in Membrane Potential

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Figure 4.4

Signals in the Dendrites and Cell Body


Incoming signal
Example: neurotransmitter

Membrane-bound receptors bind to


neurotransmitter
Receptors transduce the chemical signal to an
electrical signal by changing ion permeability of
membrane
Change in ion permeability causes change in
membrane potential (graded potential)

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Graded Potentials
Vary in magnitude depending on strength of
stimulus
More neurotransmitter more ion channels open
larger magnitude of graded potential

Depolarization
Na+ or Ca2+ channels open

Hyperpolarize
K+ and Cl channels open

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Stimulus Strength and Graded Potentials

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Figure 4.5

Graded Potentials Travel Short Distances


Conduction with decrement
Magnitude of graded potential decreases with
increasing distance from opened ion channel

Decrement due to:


Leakage of charged ions across membrane
Electrical resistance of cytoplasm
Electrical properties of membrane

Electrotonic current spread


Positive charge spreads through cytoplasm causing
depolarization of adjacent membrane

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Conduction with Decrement

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Figure 4.6

Action Potentials Travel Long Distances


Characteristics of Action Potentials:
Triggered by net graded potential at axon hillock
(trigger zone)
Do not degrade over time or distance
Travel long distances along membrane
All-or-none
Must reach threshold potential to fire
Depolarizations below threshold will not initiate an action
potential

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Action Potentials

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Figure 4.7

Integration of Graded Signals


Many graded potentials can be generated
simultaneously
Many receptor sites
Many types of receptors
Some graded potentials are depolarizations, some are
hyperpolarizations

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Integration of Graded Signals


Spatial summation
Graded potentials from different sites influence the net
change

Temporal summation
Graded potentials that occur at slightly different times
influence net change

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Spatial Summation

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Figure 4.8

Temporal Summation

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Figure 4.9

Graded Potentials vs. Action Potentials

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Table 4.1

Action Potentials (AP)


Occur only when membrane potential at axon
hillock reaches threshold
Three phases:
Depolarization
Repolarization
Hyperpolarization

Absolute refractory period


Cell incapable of generating a new AP

Relative refractory period


More difficult to generate new AP
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Action Potentials (AP)

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Figure 4.10

Voltage-Gated Channels
Change shape due to changes in membrane
potential
Closed at resting potential

Positive feedback
Influx of Na+ local depolarization more Na+
channels open more depolarization

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Voltage-Gated Channels

Na+ channels open first (depolarization)


K+ channels open more slowly (repolarization)
Na+ channels close
K+ channels close slowly
relative refractory period caused by open K+ channels

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Action Potentials (AP)

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Figure 4.10

Ion Movement
Relatively small number of ions move into and out
of cell
Single action potential has no measurable affect on
ion concentrations inside and outside cell
Na+/K+ ATPase restores concentration gradients
following repeated action potentials

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Na+ Channels Have Two Gates


Activation gate
Voltage dependent
Opens when membrane reaches threshold

Inactivation gate
Time-dependent
Closes after brief time

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Na+ Channels Have Two Gates

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Figure 4.11

Voltage-Gated Channels and the AP

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Figure 4.12

Action Potentials Travel Long Distances


All-or-none
Occurs or does not occur
All APs are same magnitude

Self propagating
An AP triggers the next AP in adjacent areas of
membrane without degradation

Electronic current spread


Charge spreads along membrane

Regenerative cycle
Ion entry electronic current spread triggering
of AP
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Action Potentials Travel Long Distances

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Figure 4.13

Myelination
Vertebrate neurons are myelinated
Myelin
Insulating layer of lipid-rich Schwann cells wrapped
around axon
Reduce leakage of charge across membrane
Schwann cells are a type of Glial cell
Cells other than neurons that support neuron function

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Myelination
Nodes of Ranvier
Areas of exposed axonal membrane between Schwann
cells

Internodes
The myelinated region

Saltatory conduction
APs leap from node to node
APs occur at nodes of Ranvier, and electrotonic
current spread through internodes
This type of conduction is very rapid

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Myelination

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Figure 4.14

Unidirectional Signals
Action potentials start at the axon hillock and travel
towards the axon terminal
Up-stream Na+ channels (just behind the region
of depolarization) are in the absolute refractory
period
The absolute refractory period prevents backward
(retrograde) transmission and summation of APs
Relatively refractory period also contributes by
requiring a very strong stimulus to cause another AP

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Information Transfer by AP
AP frequency carries information
AP frequency increases with stronger stimuli
Magnitude of each AP does not change

Maximum frequency is limited by the absolute


refractory period
Mammalian nerves can conduct 5001000 action
potentials per second

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Action Potential Frequency

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Figure 4.15

The Synapse
Signal transmission from neuron to another cell
Synapse
Presynaptic cell, synaptic cleft, and postsynaptic cell

Synaptic cleft
Space between the presynaptic and postsynaptic cell

Postsynaptic cell
May be a neuron, muscle cell, or endocrine cell

Neuromuscular junction
Synapse between a motor neuron and a skeletal muscle
cell
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Signal Transmission at a Chemical Synapse

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Figure 4.16

Amount of Neurotransmitter Released


[Ca2+]i is affected by AP frequency
More open voltage-gated Ca2+ channels [Ca2+]i

Factors that lower intracellular [Ca2+]i


Binding with intracellular buffers [Ca2+]i
Ca2+ ATPases [Ca2+]i

High AP frequency Ca2+ influx is greater than


removal [Ca2+]i many synaptic vesicles
release their contents high [neurotransmitter] in
synapse

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Acetylcholine

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Figure 4.17

Postsynaptic Cells
Postsynaptic cells have specific receptors for
neurotransmitters
Example: nicotinic ACh receptors
Similar to specific hormone receptors on target cells
Binding of neurotransmitter to receptor alters ion
permeability of postsynaptic cell
Change in membrane potential of postsynaptic cell

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Transmission of Signal Strength at Synapse


Response of postsynaptic cell influenced by amount
of neurotransmitter in synapse and number of
receptors
Amount of neurotransmitter
Rate of release rate of removal
Release determined by frequency of APs
Removal determined by
Passive diffusion out of synapse
Degradation by synaptic enzymes
Uptake by surrounding cells

Number of receptors
Density of receptors on postsynaptic cell
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Diversity of Neurons
All neurons have three functions:
Receive and integrate incoming signals
Conduct the signal along the neuron
Transmit the signal to other cells

Neurons differ in their ability to receive incoming


signals
Different receptors

Neurons differ in mechanism of signal conduction


and synaptic transmission
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Structural Diversity of Neurons

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Figure 4.18a

Functional Classes of Neurons


Afferent neurons (sensory)
Conduct action potentials towards the central
nervous system

Efferent neurons (motor)


Conduct action potentials from the central nervous
system to the organs

Interneurons
Conduct action potentials between neurons in the
central nervous system

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Neuron Classification Based on Function

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Figure 4.18b

Structural Classes of Neurons


Multipolar
Many dendrites
One axon

Bipolar
One dendrite (may have branches)
One axon

Unipolar
Single process extending from cell body
May split to form afferent and efferent branches

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Neuron Classification Based on Structure

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Figure 4.18c

Glial Cells
More abundant than neurons
90% of cells in human brain are glial cells

Do not generate or conduct APs


Do not form synapses with neurons

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Types of Glial Cells


Five main types of glial cells in vertebrates
Schwann cell
Forms myelin on motor and sensory neurons of PNS

Oligodendrocyte
Forms myelin on neurons in CNS

Astrocyte
Transport nutrients, remove debris in CNS

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Types of Glial Cells


Microglia
Remove debris and dead cells from CNS

Ependymal cells
Line fluid-filled cavities of CNS

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Glial Cells

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Figure 4.19

Diversity of Signal Conduction


Diverse mechanisms of signal conduction

Electrotonic
Action potentials
Saltatory conduction
Chemical and electrical synapses

Additional diversity in AP physiology:


Shape and speed of action potential due to
properties of Na+ and K+ channels
Function of channels
Number of channels
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Ion Channel Isoforms


Channel isoforms encoded by different genes
Voltage-gated K+ channels are highly diverse
18 genes encode for 50 isoforms in mammals

Voltage-gated Na+ channels are less diverse


11 isoforms in mammals

Each isoform has distinct functional characteristics

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Ion Channel Isoforms

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Table 4.2

Channel Density
Density of voltage-gated Na + channels affects signal
conduction
Increased density of channels lowers threshold
Increased density of channels shortens relative
refractory period

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Voltage-Gated Ca2+ Channels


Presence of voltage-gated Ca2+ channels affects AP
Open at the same time or instead of voltage-gated Na +
channels
Ca2+ enters the cell causing depolarization
Ca2+ influx is slower and more sustained than Na +
influx
Slower maximal frequency of APs due to longer
refractory period
Voltage-gated Ca2+ channels play key role in function
of cardiac muscle

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Conduction Speed of Axons


Two ways to increase speed:
Myelination
Increasing diameter of axon

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Table 4.3

Cable Properties of Axons


Similar physical principals govern current flow
through axons and undersea telephone cables
Current (I)
Amount of charge moving past a point at a given
time
A function of the voltage (V) drop across circuit and
the resistance (R) of circuit

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Cable Properties of Axons


Voltage (V)
Difference in electrical potential

Resistance (R)
Rorce opposing flow of electrical current

Ohms law: V = I R

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Cable Properties of Axons


An axon behaves like an electrical circuit
Ions moving through voltage-gated channels cause
current across membrane
Current spreads electrotonically along axon
Some current leaks out of axon and flows
backwards along outside of axon, completing
circuit

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Current Flow In Axons

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Figure 4.20a

Cable Properties of Axons


Each area of axon consists of an electrical circuit
Three resistors:
Extracellular fluid (Re)
Membrane (Rm)
Intracellular fluid (Ri)

A capacitor (Cm)
Stores electrical charge;
Two conducting materials (ICF and ECF)
Insulating layer (phospholipids)

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Cable Properties of Axons

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Figure 4.20b,c

Voltage Decreases with Distance


Change in membrane potential (voltage) during AP
decreases over distance due to resistance
Conduction with decrement
Higher resistance of intracellular and extracellular
fluids causes greater decrease in voltage along axon
Lower resistance of membrane causes greater
decrease in voltage along axon
K+ leak channels (always open)
Some + charge leaks out
Number of K+ leak channels will affect current loss and
voltage decrease along axon
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Length Constant () of Axons


Distance over which membrane potential will
decrease to 37% (1/e) of its original value
Variables affecting length constant:
Resistance of cell membrane (rm)
Resistance of intracellular fluid (ri)
Resistance of extracellular fluid (ro)
ro is usually low and constant; and is often ignored

is largest when rm is high and ri is low

rm /(ri ro )
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rm / ri

Length Constant () of Axons

rm /(ri ro )

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rm / ri

Figure 4.21

and the Speed of Conduction


Axonal conduction is a combination of electrotonic
current flow and ions flowing through voltage-gated
channels during AP
Electrotonic current flow much faster than opening
of voltage-gated channels
Electronic current flow decreases over distance

Higher allows more electrotonic current flow and


faster speed of conduction

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Axon Membrane Capacitance


Capacitance
Quantity of charge needed to create a potential
difference between two surfaces of a capacitor

Depends on three features of the capacitor:


Material properties
Generally the same in cells (lipid bilayer)

Area of two conducting surfaces


Larger area increases capacitance

Thickness of insulating layer


Greater thickness decreases capacitance
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Axon Membrane Capacitance

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Figure 4.20b and Figure 4.22

Time Constant (t)


Time over which membrane potential will decay to
37% of its maximal value
How well does the membrane hold its charge?

Variables affecting time constant:


Resistance of cell membrane (rm)
Capacitance of the cell membrane (cm)
= rmcm

Low rm or cm result in low


Capacitor becomes full faster
Faster depolarization
Faster conduction
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Time Constant (t)

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Figure 4.23

Giant Axons
Easily visible to naked eye (up to 1 mm diameter)
Not present in mammals

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Figure 4.24

Giant Axons Have High Conduction Speed


rm inversely proportional to surface area
Large diameter axons have greater surface area and
more leak channels; therefore low resistance

ri inversely proportional to volume


Large diameter axons have greater volume; therefore
low resistance

As axon diameter increases, rm and ri both decrease

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Giant Axons Have High Conduction Speed


Low rm reduces the length constant and decreases
conduction speed
Low ri increases the length constant and increases
conduction speed
Do not cancel each other out: rm is proportional to
radius, ri is proportional to radius2
Net effect of increasing axon radius is to increase
speed of conduction

rm / ri
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Axon Diameter and the Length Constant

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Figure 4.25

Myelinated Neurons in Vertebrates


Disadvantage of large axons
Take up a lot of space which
Limits number of neurons that can be packed into
nervous system

Large volume of cytoplasm makes them expensive


to produce and maintain

Myelin enables rapid signal conduction in compact


space

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Myelin Increases Conduction Speed


Increased membrane resistance
Insulators decrease current loss through leak
channels, increasing the length constant

Decreased membrane capacitance


Increased thickness of insulating layer reduces
capacitance, decreasing the time constant

High length constant and low time constant increase


conduction speed
Nodes of Ranvier are needed to boost depolarization

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Synaptic Transmission
Transfer of electrical signal from presynaptic cell to
postsynaptic cell
Electrical synapse
Gap junction

Chemical synapse
Chemical messenger crosses synaptic cleft

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Electrical and Chemical Synapses

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Figure 4.26

Electrical and Chemical Synapses


Electrical synapse

Chemical synapse

Rare in complex animals

Common in complex animals

Common in simple animals

Rare in simple animals

Fast

Slow

Bi-directional

Unidirectional

Postsynaptic signal is similar to


presynaptic

Postsynaptic signal can be different

Excitatory

Excitatory or inhibitory

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Structural Diversity of Chemical Synapses

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Figure 4.27

Neurotransmitters
Characteristics of neurotransmitters
Synthesized in neurons
Released at presynaptic cell following
depolarization
Bind to a postsynaptic receptor and cause an effect

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Neurotransmitters
More than 50 known substances
Categories

Amino acids
Neuropeptides
Biogenic amines
Acetylcholine
Miscellaneous (gases, purines, etc.)

A single neuron can produce and release more than


one neurotransmitter
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Neurotransmitters

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Table 4.4

Neurotransmitter Action
Inhibitory neurotransmitters
Cause hyperpolarization of membrane
Inhibitory postsynaptic potential (IPSP)

Make postsynaptic cell less likely to generate


an AP

Excitatory neurotransmitters
Cause depolarization of membrane
Excitatory postsynaptic potential (EPSP)

Make postsynaptic cell more likely to generate


an AP
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Neurotransmitter Receptor Function


Ionotropic receptors
Ligand-gated ion
channels
Fast
Example: nicotinic
Ach receptor

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Figure 4.28a

Neurotransmitter Receptor Function


Metabotropic receptors
Receptor changes shape
Formation of second
messenger
Alters opening of ion
channel
Slow
May lead to long-term
changes via other cellular
functions

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Figure 4.28b

Receptors for Acetylcholine


Cholinergic receptors
Nicotinic receptor
Ionotropic

Muscarinic receptor
Metabotropic
Linked to ion channel function via G-protein

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Receptors for Acetylcholine

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Figure 4.29

Receptors for Acetylcholine

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Table 4.5

Receptors for Norepinephrine


Adrenergic receptors
Alpha ()
Several isoforms
Metabotropic
Linked to ion channel function via G-protein

Beta ()
Several isoforms
Metabotropic
Linked to ion channel function via G-protein

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Receptors for Norepinephrine

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Figure 4.31

Adrenergic Receptors

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Table 4.6

Synaptic Plasticity
Change in synaptic function in response to patterns
of use
Synaptic facilitation
Repeated APs result in increased Ca2+ in terminal
Increased neurotransmitter release

Synaptic depression
Repeated APs deplete neurotransmitter in terminal
Decreased neurotransmitter release

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Synaptic Plasticity
Post-tetanic potentiation (PTP)
After train of high frequency APs there is increased
neurotransmitter release
Exact mechanism unknown, but believed to involve
changes in Ca2+ in terminal

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Post-tetanic Potentiation (PTP)

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Figure 4.32

Evolution of Neurons
Only metazoans have neurons
Other organisms have electrical signaling
Algae have giant cells that can generate APs using
Ca2+ activated Cl channels
Plants have APs involving Ca2+ that travel through the
xylem and phloem
Paramecium can change direction as a result of APs
produced by Ca2+ channels

Only metazoans have voltage-gated Na+ channels

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