Chapter 5

CHAPTER
4
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
Figure 4.1
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
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
  
RT P [ K ]  P [ Na ]  P [Cl ]i
Em  ln K

o Na

o Cl

Em = membrane potential
R = gas constant
T = temperature (Kelvin)
F = Faraday’s constant
Px = relative permeability of ion
[X] = ion concentration outside or inside membrane

The Goldman Equation
  
RT P [ K ]  P [ Na ]  P [Cl ]i
Em  ln K

o Na

o Cl

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 ion’s
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 500–1000 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 absolute
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 principles 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)
 Force opposing flow of electrical current
Ohm’s 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  r ) o   rm / ri
Length Constant () of Axons
  rm /( ri  r ) o   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)
 t = rmcm
 Low rm or cm result in low t
 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 Postsynaptic signal can be different

presynaptic
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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CHAPTER

Active Lecture Question Slides prepared by

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

 Vary in structure and properties

Four functional zones

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

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

 Neurons have a resting membrane potential (like all

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

 Changes in membrane potential act as electrical

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

Factors contributing to membrane potential

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

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

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

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

 Neurons depolarize or hyperpolarize by selectively

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

 Channels only allow specific ions to pass through

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

 Vary in magnitude depending on strength of

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

 Conduction with decrement

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 Characteristics of Action Potentials:

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 Many graded potentials can be generated

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

 Occur only when membrane potential at axon

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 Change shape due to changes in membrane

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 Na+ channels open first (depolarization)

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 Relatively small number of ions move into and out

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

 Vertebrate neurons are myelinated

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

 Action potentials start at the axon hillock and travel

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 AP frequency carries information

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 Signal transmission from neuron to another cell

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 [Ca2+]i is affected by AP frequency

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 Postsynaptic cells have specific receptors for

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 Response of postsynaptic cell influenced by

All neurons have three functions:

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Afferent neurons (sensory)

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

 More abundant than neurons

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Five main types of glial cells in vertebrates

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