Neuron Structure and Function: Active Lecture Question Slides Prepared by Dr. Alan F. Smith, Mercer University

C HAPTE R
Neuron Structure
and Function
Active Lecture Question Slides prepared by

Dr. Alan F. Smith, Mercer University
Copyright 2008 Pearson Education, Inc., publishing as Benjamin Cummings
Neurons
Vary in structure and properties
Use same basic mechanisms to send signals
Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
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)
Neural Zones
Signal conduction
Axon (some wrapped in myelin sheath)
AP travels down axon
Signal transmission
Axon terminals
Release of neurotransmitter
Neural Zones
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
Electrical Signals in Neurons

Changes in membrane potential act as electrical
signals
Changes in Membrane Potential
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
The Goldman Equation
P
[
K
]
P
[
Na
]
P
[
Cl
]i
RT
K
o
Na
o
Cl
Em
ln
F
Em = membrane potential
R = gas constant
T = temperature (Kelvin)
F = Faradays constant
Px = relative permeability of ion
[X] = ion concentration outside or inside membrane
The Goldman Equation
P
[
K
]
P
[
Na
]
P
[
Cl
]i
RT
K
o
Na
o
Cl
Em
ln
F
Other ions (Ca++, Mg++, etc.) are ignored in this

simplified form of the equation because their
permeabilities are very low
Gated Ion Channels

Neurons depolarize or hyperpolarize by selectively
altering permeability
Gated ion channels open or close in response to a
stimulus
Example: neurotransmitter
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)
Changes in Membrane Potential
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)
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
Stimulus Strength and Graded Potentials
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
Conduction with Decrement
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
Action Potentials
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
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
Spatial Summation
Figure 4.8
Temporal Summation
Figure 4.9
Graded Potentials vs. Action Potentials
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
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
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
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
Na+ Channels Have Two Gates

Activation gate
Voltage dependent
Opens when membrane reaches threshold
Inactivation gate
Time-dependent
Closes after brief time
Na+ Channels Have Two Gates
Figure 4.11
Voltage-Gated Channels and the AP
Figure 4.12

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
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
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
Myelination
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
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
Action Potential Frequency
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
Signal Transmission at a Chemical Synapse
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
Acetylcholine
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
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
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
Structural Diversity of Neurons
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
Neuron Classification Based on Function
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
Neuron Classification Based on Structure
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
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
Types of Glial Cells

Microglia
Remove debris and dead cells from CNS
Ependymal cells
Line fluid-filled cavities of CNS
Glial Cells
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
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
Ion Channel Isoforms
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
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
Conduction Speed of Axons

Two ways to increase speed:
Myelination
Increasing diameter of axon
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

Voltage (V)
Difference in electrical potential
Resistance (R)
Rorce opposing flow of electrical current
Ohms law: V = I R

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
Current Flow In Axons
Figure 4.20a

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)
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
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 )
rm / ri
Length Constant () of Axons
rm /(ri ro )
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
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
Axon Membrane Capacitance
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
Time Constant (t)
Figure 4.23
Giant Axons
Easily visible to naked eye (up to 1 mm diameter)
Not present in mammals
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
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
Axon Diameter and the Length Constant
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
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
Synaptic Transmission
Transfer of electrical signal from presynaptic cell to
postsynaptic cell
Electrical synapse
Gap junction
Chemical synapse
Chemical messenger crosses synaptic cleft
Electrical and Chemical Synapses
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
Structural Diversity of Chemical Synapses
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
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
Neurotransmitters
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
Neurotransmitter Receptor Function

Ionotropic receptors
Ligand-gated ion
channels
Fast
Example: nicotinic
Ach receptor
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
Figure 4.28b
Receptors for Acetylcholine

Cholinergic receptors
Nicotinic receptor
Ionotropic
Muscarinic receptor
Metabotropic
Linked to ion channel function via G-protein
Figure 4.29
Table 4.5
Receptors for Norepinephrine

Adrenergic receptors
Alpha ()
Several isoforms
Metabotropic
Beta ()
Several isoforms
Metabotropic
Receptors for Norepinephrine
Figure 4.31
Adrenergic Receptors
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
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
Post-tetanic Potentiation (PTP)
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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C HAPTE R

Active Lecture Question Slides prepared by

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Electrical Signals in Neurons

Neurons are excitable

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Electrical Signals in Neurons

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Changes in Membrane Potential

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Goldman equation for the calculation of membrane

The Goldman Equation

The Goldman Equation

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

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Gated Ion Channels

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Gated Ion Channels

As permeability to a specific ion increases,

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Changes in Membrane Potential

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Signals in the Dendrites and Cell Body

Membrane-bound receptors bind to

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Stimulus Strength and Graded Potentials

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Graded Potentials Travel Short Distances

Decrement due to:

Electrotonic current spread

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Conduction with Decrement

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Action Potentials Travel Long Distances

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Integration of Graded Signals

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Integration of Graded Signals

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Graded Potentials vs. Action Potentials

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Action Potentials (AP)

Absolute refractory period

Relative refractory period

Action Potentials (AP)

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Na+ channels open first (depolarization)

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Action Potentials (AP)

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Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Na+ Channels Have Two Gates

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Na+ Channels Have Two Gates

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings