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Nervous Tissue
Body perceives and responds to multiple sensations
Controls multiple muscle movements
Others movements without voluntary input
e.g., beating of the heart
Nervous System
Controls and interprets all these sensations and muscle movements
receptors include:
eyes and nose
tongue and ears
skin
proprioceptors (receptors detecting body position)
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Structural organization
Central
nervous
system (CNS)
Peripheral
nervous
system (PNS)
Brain
Spinal cord
Nerves
Ganglia
Functional organization
Somatic sensory
Visceral sensory
Sensory input
that is consciously
perceived from
receptors (e.g.,
eyes, skin, ears)
Somatic motor
Motor output
that is consciously
or voluntarily
controlled; effector
is skeletal muscle
Autonomic motor
Motor output that is
not consciously or
is involuntarily
controlled; effectors
are cardiac muscle,
smooth muscle,
and glands
Nervous TissueNeurons:
General Characteristics
Special characteristics of neurons
Excitability
responsive to stimulation
type dependent on its location
most respond only to binding of molecules, neurotransmitters
Conductivity
electrical charges propagated along membrane
can be local and short-lived or self-propagating
Nervous TissueNeurons:
General Characteristics
Special characteristics of neurons (continued)
Secretion
release neurotransmitters in response to electrical charges
given neuron releasing only one type of neurotransmitter
may have excitatory or inhibitory effect on target
Extreme longevity
most formed before birth still present in advanced age
Amitotic
mitotic activity lost in most neurons
not always the case (e.g., occasionally in hippocampus)
Nervous TissueNeurons:
General Characteristics
Name the five distinguishing characteristics
of all neurons.
Excitability, conductivity, secretion, extreme
longevity, amitotic
Nervous TissueNeurons:
General Characteristics
Components of neurons
Cell body
enclosed by plasma membrane
contains cytoplasm surrounding a nucleus
neurons control center
conducts electrical signals to axon
perikaryon, cytoplasm within cell body
Nervous TissueNeurons:
General Characteristics
Components of neurons (continued)
Cell body (continued)
nucleus with prominent nucleolus
free and bound ribosomes termed chromatophilic substance
due to dark staining with basic dyes
gray color of gray matter
due to chromatophilic substance and lack of myelin
Nervous TissueNeurons:
General Characteristics
Components of neurons (continued)
Dendrites
short processes branching off cell body
may have one or many
receive input and transfer it to cell body
more dendrites = more input possible
Nervous TissueNeurons:
General Characteristics
Components of neurons (continued)
Axon
longer process emanating from cell body
makes contact with other neurons, muscle cells, or glands
first part, a triangular region, axon hillock
cytoplasm here termed axoplasm
plasma membrane here termed axolemma
devoid of chromatophilic substance
Nervous TissueNeurons:
General Characteristics
Components of neurons
Axon (continued)
gives rise to side branches, axon collaterals
branch extensively at distal end into telodendria (axon terminals)
at extreme tips, expanded regions, synaptic knobs
knobs containing numerous synaptic vesicles
contain neurotransmitter
Nervous TissueNeurons:
General Characteristics
Cytoskeleton
Composed of microfilaments, intermediate filaments, microtubules
Intermediate filaments, termed neurofilaments
aggregate to form bundles, neurofibrils
provide tensile strength through the neuron
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Dendrite
Perikaryon
Dendrites
Chromatophilic
substance
Nucleolus
Nucleus
Nucleus
Cell body
Chromatophilic
substance
Cell body
Axon hillock
Axon
hillock
Axoplasm
Axolemma
Neurofibrils
Nucleus of
glial cell
Structures in
a Typical
Neuron
(Figure
12.2)
Axon
collateral
Axon (beneath
myelin sheath)
Neurolemmocyte
Neurofibril node
LM 100x
Axon
(b)
Myelin sheath
Telodendria
Synaptic
knobs
Synaptic vesicles
containing
neuro transmitter
Synaptic cleft
Postsynaptic neuron
(or effector)
Synapse
(a)
b: Ed Reschke
Nervous TissueNeurons:
General Characteristics
What are the functions of dendrites, axon,
and neurofibrils?
Dendrites conduct electrical signals toward the cell
body. They receive input that they transfer to the
cell body.
The axon is used to make contact with other neurons,
muscle cells, or gland cells.
Neurofibrils give tensile support to neurons.
Nervous TissueNeurons:
Classification of Neurons
Structural classification
Structural classification of neurons
according to number of neuron processes
Multipolar neurons
most common type
have many dendrites and a single axon
Bipolar neurons
have two processes extending from cell body
one dendrite and one axon
limited, e.g., in retina of the eye
Nervous TissueNeurons:
Classification of Neurons
Structural classification (continued)
Unipolar neurons
have single short neuron process
emerges from cell and branches like a T
also called pseudounipolar
start out as bipolar neurons during development
axons with peripheral process (dendrites to cell body)
axons with central process (cell body into CNS)
Nervous TissueNeurons:
Classification of Neurons
Structural classification (continued)
Anaxonic neurons
have dendrites and no axons
produce local electrical changes but no action potentials
Structural
Classification
of Neurons
(Table
12.1)
Nervous TissueNeurons:
Classification of Neurons
Functional classification
Sensory neurons (afferent neurons)
neurons of the sensory nervous system
conduct input from somatic and visceral receptors
most unipolar, few bipolar
cell bodies usually in posterior root ganglia, outside CNS
Motor neurons (efferent neurons)
neurons of the motor nervous system
conduct motor output to somatic and visceral effectors
all multipolar
most cell bodies in CNS
Nervous TissueNeurons:
Classification of Neurons
Functional classification (continued)
Interneurons (association neurons)
entirely within the CNS
receive stimulation from many other neurons
receive, process, and store information
decide how body responds to stimuli
facilitate communication between sensory and motor neurons
99% of neurons
generally multipolar
Posterior root
ganglion
Cell body of sensory
neuron
Sensory
input
Skin receptors
Sensory neuron
Motor
output
Interneuron
Motor neuron
Skeletal
muscle
Spinal cord
Nervous TissueNeurons:
Classification of Neurons
How are the different processes that extend
from a cell body used to structurally classify
neurons?
Multipolar neurons: many dendrites and single
axon
Bipolar neurons: one dendrite and one axon
Unipolar neurons: single short neuron process
which branches like a T
Anaxonic neurons: dendrites and no axon
Nervous TissueNeurons:
Relationship of Neurons and Nerves
Nerve
Cablelike bundle of parallel axons
Macroscopic structure
Epineurium
thick layer of dense irregular connective tissue
encloses the entire nerve
provides support and protection
Nervous TissueNeurons:
Relationship of Neurons and Nerves
Nerve (continued)
Perineurium
layer of dense irregular connective tissue
wraps bundles of axons, fascicles
supports blood vessels
Endoneurium
delicate layer of areolar connective tissue
separates and electrically insulates each axon
has capillaries that supply the axon
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Perineurium
Fascicle
Blood vessels
Endoneurium
Neurolemmocyte
Epineurium
Nerve
Structure of
a Nerve and
Ganglion
(Figure
12.4)
Axon
Perineurium
Fascicle
Endoneurium
Axon
M
SE
Neurolemmocyte
(b)
(a)
Blood vessels
Ganglion
Cell bodies
Nerve
Axons
Axons
(c)
b: Dr. Richard Kessel & Dr. Randy Kardon/Tissues & Organs/Visuals Unlimited
0x
45
Nervous TissueNeurons:
Relationship of Neurons and Nerves
Classification of nerves
Structural classification
Cranial nerves
extend from brain
Spinal nerves
extend from spinal cord
Nervous TissueNeurons:
Relationship of Neurons and Nerves
Classification of nerves (continued)
Functional classification
Sensory nerves
contain only sensory neurons
Motor nerves
contain primarily motor neurons
Mixed nerves
contain both sensory and motor neurons
most named nerves in this category
individual neurons transmitting one type of information
Nervous TissueNeurons:
Relationship of Neurons and Nerves
What are the three connective tissue
wrappings in a nerve, and what specific
structure does each ensheathe?
The epineurium encloses the entire nerve.
The perineurium encloses bundles of axons.
The endoneurium encloses individual axons.
Synapses
Learning Objectives:
1) Define a synapse.
2) Describe the essential structural and functional differences
between a chemical synapse and an electrical synapse.
Synapses
Synapse
Where neuron functionally connected to neuron or effector
Two types: chemical and electrical
Synapses
Chemical synapse
Most common
Composed of presynaptic neuron, signal producer
Composed of postsynaptic neuron, signal receiver
Between axon and any portion of postsynaptic neuron
most commonly with a dendrite
Knob almost touches the postsynaptic neuron
narrow fluid filled gap, the synaptic cleft
Synapses
Transmission at chemical synapse
Synapses
Electrical synapse
Synapses
What is the mode of transmission in a
chemical synapse?
Molecules stored in synaptic vesicles are released
from the synaptic knob of a presynaptic neuron into
the synaptic cleft. Some neurotransmitter diffuses
across the cleft and binds receptors on the
postsynaptic membrane.
Microglial
cell
Neuron
Astrocyte
Oligodendrocyte
Perivascular
feet
Capillary
Myelinated axon
Ependymal
cells
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Satellite cells
Neurofibril nodes
Axon
Nucleus
Axon
Cell body of
sensory neuron
Neurolemmocyte
Posterior root
Myelin sheath
Neurilemma
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1
Neurolemmocyte
starts to wrap around
a portion of an axon.
Axon
Neurolemmocyte
Nucleus
Myelination
of PNS Axons
(Figure
12.6)
Neurolemmocyte
cytoplasm and
plasma membrane
begin to form
consecutive layers
around the axon as
wrapping continues.
The overlapping
inner layers of the
neurolemmocyte
plasma membrane
form the myelin
sheath.
Direction of
wrapping
Cytoplasm of the
neurolemmocyte
Myelin sheath
Eventually, the
neurolemmocyte
cytoplasm and
nucleus are pushed
to the periphery of
the cell as the myelin
sheath is formed.
Myelin sheath
Neurolemmocyte
nucleus
Neurilemma
Neurolemmocyte
in the PNS
Can myelinate only
1 mm of single axon
Takes many to
myelinate entire axon
Gaps between
neurolemmocytes
neurofibril
nodes, or nodes
of Ranvier
(Figure 12.7a)
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PNS
Neurolemmocytes
Neurofibril
node
Neuron
cell body
Neurilemma
Myelin sheath
Axon
Oligodendrocyte
in the CNS
Can myelinate 1 mm
of many axons
Extensions wrapping
around axons
No neurilemma
formed
Neurofibril nodes
between adjacent
wraps
(Figure 12.7b)
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CNS
Oligodendrocytes
Neurofibril
node
Axons
Myelin sheath
(b) Myelination by oligodendrocytes
Unmyelinated axons
1
Unmyelinated
axons
Neurolemmocyte
Neurolemmocyte starts
to envelop multiple
axons.
Axons
Unmyelinated
axon
Neurolemmocyte
(a)
Neurolemmocyte
nucleus
Myelin sheath
Myelinated axon
TEM 60,000x
2 The unmyelinated
axons are enveloped by
the neurolemmocyte,
but there are no myelin
sheath wraps around
each axon.
(b)
b: Donald Fawcett/Visuals Unlimited
Neurilemma
Axon Regeneration
Learning Objectives:
1) Identify factors that influence regeneration of PNS axons, and
explain why axon regeneration in the CNS is limited.
2) Describe the events of Wallerian degeneration and axon
regrowth.
Axon Regeneration
Factors influencing axon regeneration
PNS axons
vulnerable to cuts, trauma
Regeneration possible if
cell body intact
enough neurilemma remains
Regeneration success more likely if
amount of damage less extensive
smaller distance between site of damage and structure it innervates
Axon Regeneration
Steps of axon regeneration
1) Axon severed by trauma
2) Sealing off and swelling of proximal portion of severed axon
disintegration of distal axon and myelin sheath
termed Wallerian degeneration
survival of neurilemma
Axon Regeneration
Steps of axon regeneration (continued)
3) Formation of regeneration tube
neurilemma and remaining endoneurium
4) Axon regeneration and remyelination
guided by regeneration tube
nerve growth factor released by neurolemmocytes
5) Innervation restored
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1 Trauma severs axon.
Endoneurium
Neurilemma
Regeneration
of PNS axons
(Figure
12.9)
Neurilemma
5 Innervation to effector
is restored.
Axon Regeneration
CNS axon regeneration
Extremely limited
growth-inhibiting molecules secreted by oligodendrocytes
larger number of axons crowded within the CNS
regrowth obstructed by scars from astrocytes and connective tissue
Axon Regeneration
What are the two primary factors
determining the effectiveness of PNS axon
regeneration?
The amount of damage and distance between site of
damage and innervated structure.
Ultrastructure of Neurons
Learning Objectives:
1) Distinguish between a pump and a channel, and identify the
pumps and channels located along the entire neuron plasma
membrane.
2) List and describe the four functional neuron segments,
including the distribution of channels and pumps in each.
3) Describe the distribution of substances between the inside and
the outside of a neuron.
Pumps
Type of transport protein
Move substances against concentration gradient
Require energy
e.g., sodium-potassium and calcium pumps in plasma membr ane
(Figure 12.10a)
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Interstitial
fluid
Cytosol
Breakdown of ATP
(releases energy)
ATP binding
site
K+
ATP
ADP
Na+
Na+
Na+/K+
pump
(a) Sodium-potassium (Na+/K+) pump
Channels
Move substances down
concentration gradient
Leak channels
always open for
continuous diffusion
e.g., sodium ion and
potassium ion channels
(Figure 12.10b)
Copyright The McGraw-Hill Companies, Inc. Permission required for reproduction or display.
Interstitial
fluid
Cytosol
Na+
Channels (continued)
Chemically gated channels
normally closed
allow specific type of ion to diffuse when open
e.g., chemically gated K+ channels
(Figure 12.10c)
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Closed
Open
Neurotransmitter
binds to gate
K+
Channels (continued)
Voltage-gated channels
normally closed
open in response to changes in electrical charge across membrane
allow specific type of ion to diffuse
e.g., voltage gated Na+ channels
(Figure 12.10d)
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+ + + +
+ + + +
Inactivation
gate (open) Activation
gate (closed)
Na+
+ + +
Inactivation
gate (open)
Na+
+ + + +
+ + + +
Inactivation
gate (closed)
+ + + +
Activation
gate (open)
Activation
gate (open)
Ultrastructure of Neurons:
Pumps and Channels
Three states of voltage-gated Na+ channels
Most gated channels either closed or open
Voltage gated Na+ channels unique
have two gates (activation gate and inactivation gate)
have three states
Ultrastructure of Neurons:
Pumps and Channels
Three states of voltage-gated Na+ channels
1) Resting state
inactivation gate open
activation gate closed
entry of Na+ prevented
2) Activation state
inactivation gate open
activation gate open (in response to voltage change)
Na+ moving through the channel
Ultrastructure of Neurons:
Pumps and Channels
Three states of voltage-gated Na+ channels
(continued)
3) Inactivation state
activation gate open
inactivation gate temporarily closed
entry of Na+ prevented
4) Resting state reestablished
inactivation gate open
activation gate closed
Ultrastructure of Neurons:
Pumps and Channels
Distribution of pumps and channels
Entire plasma membrane of neuron
Na+ leak channels
K+ leak channels
present in greater numbers than Na+ leak channels
easier for K+ to move through
Na+/K+ pumps
important in maintaining resting membrane potential
Ultrastructure of Neurons:
Pumps and Channels
Distribution of pumps and channels (continued)
Membrane of functional segments in a neuron
Receptive segment
includes dendrites and cell body
chemically gated channels here (cation channels, K +, Cl-)
no significant voltage-gated channels
Initial segment
composed of axon hillock
contains voltage-gated Na+ and K+ channels
Ultrastructure of Neurons:
Pumps and Channels
Distribution of pumps and channels
Membrane of functional segments in a neuron
(continued)
Conductive segment
length of the axon and its branches
contains voltage-gated Na+ and K+ channels
Transmissive segment
includes synaptic knobs
contains voltage-gated Ca2+ channels and pumps
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Receptive segment
Chemically
gated cation
channel
Distribution
of Pumps and
Channels in
the Plasma
Membrane of
a Neuron
(Figure
12.11)
Chemically
gated K+
channel
Chemically
gated Cl
channel
(b)
Cell body
Dendrites
Initial segment
Voltage-gated Voltage-gated
Na+ channel
K+ channel
Axon hillock
Na+/K+
pump
Na+ leak
channel
(c)
K+ leak
channel
Conductive segment
Voltage-gated Voltage-gated
+
Na channel
K+ channel
Entire neuron
Axon
(d)
Transmissive segment
Voltage-gated
Ca2+ channel
(a)
Ca2+ pump
Synaptic bulb
(e)
Ultrastructure of Neurons:
Pumps and Channels
What is the difference between a chemically
gated channel and a voltage-gated channel
in terms of how they function?
Chemically gated channels open in response to
binding of a neurotransmitter.
Voltage-gated channels open in response to changes in
electrical charge.
At equilibrium
chemical concentration gradient equal to electrical gradient opposing
movement
70mV
+ + + + + + + + + +
+ + + + + + + + + +
70 mV
Voltmeter
Na+ leak
channel
K+ leak
channel
Na+/K+
pump
Cl
Na+
Interstitial fluid
Greater concentration
of Na+ and Cl
Microelectrode
+ + +
+ +
+ +
+ +
+ + +
+ + +
Plasma membrane
70 mV
Cytosol
Greater concentration
of K+, Pi, and proteins
K+
ADP
Pi
Protein
ATP
70 mV
+
++
++
+++ ++++
+++ ++++
Gated Na+
channel
Gated K+
channel
Na+
Interstitial
fluid
Gated Cl
channel
Cl
+ +
+ + + + +
+ + + + +
+ +
Plasma
membrane
Cytosol
K+
(a) Resting membrane potential
e.g., 70 mV
60 mV
+
+ + + + + +
+ + + + + +
+ +
+ + + + +
+ + + +
+ +
Na+
K+
(b) Depolarization: Na+ flows in
e.g., 60 mV
80 mV
+
Na+
+ + + + + +
+ + + + + +
Gated K+
channel
Gated Cl
channel
K+
Cl
+ +
+ + + + +
+ + + +
+ +
e.g., 80 mV
(c) Hyperpolarization: K+ flows out or Cl flows in
Local potentials
Occur in the receptive segment of a neuron
Due to opening of chemically gated channels
Temporarily allow passage of small amount of specific ion
Local current established
ions moving parallel to plasma membrane
experiences resistance from contents of cytosol
eventually becomes weaker and ceases
are self-propagated
maintain intensity as move to synaptic knob
obey the all or none law
if threshold reached, action potential sent
if not reached, no action potential sent
See Table 12.3: Graded Potential Versus Action Potential
Axons of
presynaptic
neuron
Postsynaptic
neuron
Excitatory
neurotransmitter
Postsynaptic neuron
0
Synaptic vesicles
containing excitatory
neurotransmitter
20
Voltage (mV)
Synaptic
knob
40
EPSP
Threshold
60
4 EPSP propagates
toward axon hillock.
Stimulus
70
Resting
membrane
potential
80
Time (msec)
(a)
Synaptic cleft
1
Axons of
presynaptic
neuron
Postsynaptic neuron
0
20
K+
Cl
3
Inside of neuron
becomes more negative;
called IPSP (e.g., 72 mV).
Voltage (mV)
Synaptic vesicles
containing inhibitory
neurotransmitter
40
Threshold
60
Cl
Stimulus
4
IPSP propagates
toward axon
hillock.
70
80
IPSP
Time (msec)
(b)
Resting
membrane
potential
EPSP
Postsynaptic
neuron
Synaptic
knob
Presynaptic
axons
Axons of presynaptic
neuron
Dendrites
Cell body of
postsynaptic neuron
Myelin
sheath
Axon
SEM 80,000x
IPSP
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Spatial summation
Membrane
potential (mV)
+30
Spatial
Summation
at the Axon
Hillock
(Figure
12.7a)
Initial segment
Action
potential
55
P1
P4
P2 P3
P5
Threshold
70
Time (m sec)
Dendrites
Cell body of postsynaptic neuron
P1
P2
Myelin
sheath
P3
EPSPs
P4
P5
Axon
Axon hillock
Copyright The McGraw-Hill Companies, Inc. Permission required for reproduction or display.
Temporal summation
+30
Membrane
potential (mV)
Temporal
Summation at
the Axon
Hillock
(Figure
12.7b)
Action
potential
0
P2
Threshold
55
70
Time (m sec)
Axon of presynaptic
neuron (P2)
Postsynaptic neuron
P2
EPSPs
Axon
E.g., lidocaine
Inhibit action of voltage-gated Na+ channels
Block nerve signal
Pain signal blocked from reaching CNS
Application of ice
reduces pain sensation
slows transmission of sensory action potentials
++ +
+
Nerve signal:
propagation of
action potential
++
+ + ++ + + + + + ++ + + + ++ + + + + + ++ + + + ++ + + + + + ++ + +
Axon
+ ++ +
hillock
+ ++ +
++ ++ + + + + + ++ + + + ++ + + + + + ++ + + + ++ + + + + + ++ + +
+
+
Repolarization Depolarization
+
+
+ +
(a)
Interstitial
fluid
Na+
+ + + + + + + + + + + + + +
Cytosol
Closed
(resting state)
(b)
Closed
(inactivation
state)
+ + + + + + + + +
+ 55
mv
Open
(activation state)
+30
mv
As threshold is reached
Na+ channels open and Na+
diffuses in; polarity reversed
70
mv
Closed
(resting
state)
K+
+ + + + + + + + + + + +
Closed
70
mv
+ + + + + + + + + +
+ +
Open
+30 + + + + +
mv
Closed
+30
+10
0
mV
10
30
50
Threshold
70
90
Resting
membrane
potential
0
1
Time (msec)
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Neuromuscular junction
Transmissive
Segment:
Release of
Neurotransmitter
(Figure
12.21a)
+
+ + ++ +
1
++
++ + + +
++
+
Synaptic vesicles
merge with synaptic
knob plasma membrane
and neurotransmitter is
released by exocytosis.
Neurotransmitter
Synaptic knob
(a)
Synaptic vesicle
(contains
neurotransmitter)
Neurotransmitter
crosses synaptic cleft
and attaches to receptors
on a muscle, as shown.
(Or to receptors of a
neuron or gland.)
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Neurofibril node
Na+
K+
Myelin sheath
Diffusion of Na+
through axoplasm
+ + + + +
+ + + + +
+ + +
+ + +
+ + +
+ + + + +
+ + + + +
+ + +
+ + +
+ + +
Action
potential
Repolarization
Depolarization
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Types of circuits
Converging circuit
input that converges at a single
postsynaptic neuron
e.g., multiple sensory neurons
synapsing on neurons in
salivary nucleus
causes salivary nucleus to alter
activity of salivary glands
inputs originating from more
than one stimulus
multiple inputs leading to
single output: saliva production
(Figure 12.24a)
Input
Input
Input
Input
Output
(a) Converging
circuit
Types of circuits
(continued)
Copyright The McGraw-Hill Companies, Inc. Permission required for reproduction or display.
Diverging circuit
spreads information
from one presynaptic
neuron to several
postsynaptic neurons
e.g., neurons in the brain
controlling movements
of skeletal muscles
single or few inputs
leading to multiple
outputs
(Figure 12.24b)
Input
Output
Output
Output
(b) Diverging
circuit
Output
Output
Copyright The McGraw-Hill Companies, Inc. Permission required for reproduction or display.
Input
Output
(c) Reverberating
circuit
Copyright The McGraw-Hill Companies, Inc. Permission required for reproduction or display.
Input
Output
(d) Parallel-after-discharge
circuit