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Chapter 12

*Lecture Outline
*See separate FlexArt PowerPoint slides for all
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without notes.

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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

Introduction to the Nervous System


Learning Objectives:
1) Describe the three general functions of the nervous system.
2) Identify the structural components included in both the CNS
and the PNS.
3) Explain the functional organization of the nervous system.

Introduction to the Nervous System:


General Functions
Nervous system
Bodys primary communication and control system
Integrates and regulates body functions
Uses electrical activity
transmitted along specialized nervous system cells

Introduction to the Nervous System:


General Functions
Nervous system activities
Collects information
specialized nervous structures, receptors
monitor changes in external and internal environment, stimuli
e.g., receptors in the skin detecting information about touch
Processes and evaluates information
then determines if response required

Introduction to the Nervous System:


General Functions
The nervous system activities (continued)
Initiates response to information
initiate response via nerves to effectors
include muscle tissue and glands
e.g., muscle contraction or change in gland secretion

Introduction to the Nervous System:


General Functions
Structural organization: central versus peripheral
nervous system
Central nervous system
anatomic division of the nervous system
includes brain and spinal cord
brain protected in the skull
spinal cord protected in the vertebral canal
Peripheral nervous system
other anatomic division
includes nerves, bundles of neuron processes
includes ganglia, clusters of neuron cell bodies

Introduction to the Nervous System:


General Functions
Functional organization: sensory versus motor
nervous system
Sensory nervous system
also known as afferent nervous system
responsible for receiving sensory information from receptors
transmits information to the CNS
further divided into somatic and visceral sensory

Introduction to the Nervous System:


General Functions
Functional organization: sensory versus motor
nervous system (continued)
Somatic sensory
detects stimuli that we consciously perceive

receptors include:
eyes and nose
tongue and ears
skin
proprioceptors (receptors detecting body position)

Introduction to the Nervous System:


General Functions
Functional organization: sensory versus motor
nervous system (continued)
Visceral sensory
detects stimuli we do not consciously perceive
receptors include:
structures within blood vessels
structures within internal organs
e.g., detecting stretch of organ wall

Introduction to the Nervous System:


General Functions
Functional organization: sensory versus motor
nervous system (continued)
Motor nervous system
also known as efferent nervous system
initiates and transmits motor output from CNS
transmits information to the effectors
may be further divided into the somatic and visceral parts

Introduction to the Nervous System:


General Functions
Functional organization: sensory versus motor
nervous system (continued)
Somatic motor
transmits motor output from CNS to voluntary skeletal muscles
effector consciously controlled
e.g., pressing on accelerator of your car
Autonomic motor
transmits output from CNS without conscious control
transmits to cardiac muscle, smooth muscle, glands

Organization of the Nervous System (Figure 12.1)

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Structural organization

Central
nervous
system (CNS)

Peripheral
nervous
system (PNS)

Brain
Spinal cord

Nerves
Ganglia

Functional organization

Sensory nervous system


detects stimuli and
transmits information from
receptors to the CNS

Somatic sensory

Visceral sensory

Sensory input
that is consciously
perceived from
receptors (e.g.,
eyes, skin, ears)

Sensory input that


is not consciously
perceived from
blood vessels and
internal organs
(e.g., heart)

Motor nervous system


initiates and transmits
information from the CNS
to effectors

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

Introduction to the Nervous System:


General Functions
What are the two primary functional
divisions of the nervous system? How do
they differ?
Sensory nervous system and the motor nervous
system
The sensory nervous system detects stimuli and
transmits information from receptors to the CNS.
The motor nervous system initiates and transmits
information from the CNS to effectors.

Nervous Tissue: Neurons


Learning Objectives:
1) Describe five distinguishing features common to all neurons.
2) Describe the three basic anatomic features common to most
neurons.
3) Identify and describe the structures unique to neurons.
4) Distinguish between fast axonal transport and slow axonal
transport, and give examples of the different substances moved
by each.

Nervous Tissue: Neurons


Learning Objectives: (continued)
5) Name and describe the four structural categories of neurons.
6) Identify the three functional categories of neurons and where
each is primarily located.
7) Describe the structure of a nerve, including the three layers of
connective tissue wrappings.
8) Explain how nerves are classified structurally and functionally.

Nervous Tissue: Neurons


Two cell types in nervous tissue
Neurons
basic structural unit of the nervous system
excitable cells that transmit electrical signals
Glial cells
nonexcitable cells that primarily support and protect neurons

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: Neuron Transport


Bidirectional axonal transport
Axons dependent on cell body
for newly synthesized materials
for break down of used materials
Anterograde transport
movement of materials from cell body to synaptic knobs
Retrograde transport
movement of materials from synaptic knobs to cell body

Nervous TissueNeurons: Neuron Transport


Fast axonal transport

Occurs at about 400 mm per day


Involves movement along microtubules
Power from specialized motor proteins that split ATP
Anterograde or retrograde motion possible
anterograde transport of vesicles, organelles, glycoproteins
retrograde transport of used vesicles, potentially harmful agents

Nervous TissueNeurons: Neuron Transport


Slow axonal transport
Occurs at about 0.1 to 3 mm per day
Results from flow of axoplasm
Substances only moved from cell body towards knob
enzymes, cytoskeletal components, new axoplasm

Nervous TissueNeurons: Neuron Transport


Which type of axonal transport is both
anterograde and retrograde? What sorts of
substances are transported by this method?
Fast axonal transport
Anterograde transport of vesicles, organelles,
glycoproteins
Retrograde transport of used vesicles, potentially harmful
agents

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

Functional Classification of Neurons (Figure 12.3)


Copyright The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

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

Epineurium Blood vessels

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

Neurotransmitter molecules released from synaptic knob


Released from synaptic vesicles into cleft
Diffusion of neurotransmitter across cleft
Binding of some neurotransmitters to receptors
Synaptic delay
time between neurotransmitter release and binding
Single postsynaptic neuron
often stimulated by more than one neuron

Synapses
Electrical synapse

Much less common


Presynaptic and postsynaptic neuron physically bound together
Gap junctions present
No delay in passing electrical signal
In limited regions of brain and eyes

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.

Nervous Tissue: Glial Cells


Learning Objectives:
1) List the distinguishing features of glial cells.
2) Describe the structure and function of the four types of glial
cells within the CNS, and the two types of glial cells of the
PNS.
3) Define myelination, and describe the composition and function
of a myelin sheath.
4) Distinguish between the myelination process carried out by
neurolemmocytes and by oligodendrocytes.

Nervous TissueGlial Cells:


General Characteristics
Glial cells (neuroglia)

Nonexcitable cells found in CNS and PNS


Smaller than neurons
Capable of mitosis
Far outnumber neurons
Half volume of nervous system

Nervous TissueGlial Cells:


General Characteristics
Glial cells (continued)
Physically protect and nourish neurons
Provide physical scaffolding for nervous tissue
help guide migrating neurons to their destination
Critical for normal function at neural synapses

Nervous TissueGlial Cells:


Types of Glial Cells
Glial cells of the CNS
Astrocytes
Starlike shape from surface projections
Processes touching capillary walls and neurons
ends termed perivascular feet
Most abundant glial cell in CNS

Nervous TissueGlial Cells:


Types of Glial Cells
Glial cells of the CNS
Astrocytes (continued)
Help form the blood-brain barrier
feet wrap around capillaries in the brain
together form the blood-brain barrier
strictly controls substances entering brain nervous tissue from
blood
protects neurons from toxins
allows nutrients to pass

Nervous TissueGlial Cells:


Types of Glial Cells
Glial cells of the CNS
Astrocytes (continued)
Regulate tissue fluid composition
control movement of substances between blood and interstitial
fluid
e.g., regulate K+ concentration
need constant K+ level for neuron electrical activity
Form a structural network
cytoskeleton strengthening and organizing nervous tissue

Nervous TissueGlial Cells:


Types of Glial Cells
Glial cells of the CNS
Astrocytes (continued)
Assist neuronal development
direct development of neurons in fetal brain
secrete chemicals regulating formation of connections
Occupy the space of dying neurons
space formerly occupied by dead neurons
filled by cells produced by astrocyte division

Nervous TissueGlial Cells:


Types of Glial Cells
Glial cells of the CNS (continued)
Ependymal cells

Line internal cavities of brain and spinal cord


Ciliated simple cuboidal or simple columnar epithelial cells
Slender processes with extensive branching
Form choroid plexus with nearby blood capillaries
helps produce cerebrospinal fluid
liquid that bathes external CNS and fills internal cavities
cilia helping to circulate CSF

Nervous TissueGlial Cells:


Types of Glial Cells
Glial cells of the CNS (continued)
Microglia

Small cells with slender branches


Smallest percentage of CNS glial cells
Phagocytic cells of the immune system
Wander CNS and replicate in infection
Engulf infectious agents
Remove debris from dead or damaged tissue

Nervous TissueGlial Cells:


Types of Glial Cells
Glial cells of the CNS (continued)
Oligodendrocytes

Large cells with slender extensions


Processes ensheathing portions of axons of different neurons
Processes repeatedly wrapping around axon
Insulate axons in a myelin sheath
Prevent passage of ions through axonal membrane
Allow for faster action potential propagation through CNS

Cellular Organization of Nervous Tissue:


CNS Glial Cells (Figure 12.5a)
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Microglial
cell
Neuron

Astrocyte

Oligodendrocyte

Perivascular
feet
Capillary

Myelinated axon

Ependymal
cells

Myelin sheath (cut)


Ventricle of
brain

(a) CNS glial cells

Nervous TissueGlial Cells:


Types of Glial Cells
Glial cells of the PNS
Satellite cells
Arranged around neuronal cell bodies in a ganglion
Physically separate cell bodies in ganglion from surrounding fluid
Regulate the exchange of nutrients and waste products
e.g., surrounding bodies of sensory neurons in a posterior root
ganglion

Nervous TissueGlial Cells:


Types of Glial Cells
Glial cells of the PNS (continued)
Neurolemmocytes
Also known as Schwann cells
Ensheathe PNS axons to form myelin sheath
Allows for faster action potential propagation
See Table 12.2: Glial Cells

Cellular Organization of Nervous Tissue:


PNS Glial Cells (Figure 12.5b)

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Posterior root ganglion


Neuron cell body

Satellite cells

Neurofibril nodes
Axon

Nucleus

Axon

(b) PNS glial cells

Cell body of
sensory neuron

Neurolemmocyte
Posterior root

Myelin sheath
Neurilemma

Nervous TissueGlial Cells:


Types of Glial Cells
If a person suffers from meningitis (an
inflammation of the coverings around the
brain), which type of glial cell usually
replicates in response to the infection?
Microglia

Nervous TissueGlial Cells:


Types of Glial Cells
Which specific type of glial cell ensheathes
axons in the PNS?

Neurolemmocytes (Schwann cells)

Nervous TissueGlial Cells:


Types of Glial Cells
Clinical View: Tumors of the Central Nervous System

Neoplasm from unregulated cell growth, tumors


Sometimes occur in CNS
Tumors originating from the brain, primary brain tumors
Typically originate in supporting tissues
tissues with capacity to undergo mitosis
from meninges (protective membranes of CNS) or glial cells
Gliomas, glial cell tumors
may be relatively benign
may be malignant, capable of metastasizing

Nervous TissueGlial Cells:


Myelination
Myelination
Process by which part of an axon wrapped in myelin
Myelin, insulating covering around axon
consists of repeating layers of glial cell plasma membrane
has high proportion of lipids
gives glossy appearance and insulates axon
Completed by neurolemmocytes (PNS)
Completed by oligodendrocytes (CNS)

Nervous TissueGlial Cells:


Myelination
Process of myelination

Neurolemmocyte starts to wrap around axon


Its cytoplasm and plasma membrane begin to form layers
Wrapping continues
Layers of plasma membrane form the myelin sheath
Its cytoplasm and nucleus is pushed to the periphery
termed 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)

Copyright The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

PNS

Neurolemmocytes

Neurofibril
node

Neuron
cell body

Neurilemma
Myelin sheath

(a) Myelination by neurolemmocytes

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)

Copyright The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

CNS
Oligodendrocytes

Neurofibril
node
Axons

Myelin sheath
(b) Myelination by oligodendrocytes

Nervous TissueGlial Cells:


Myelination
Unmyelinated axons

Associated with neurolemmocytes


No myelin sheath covers them
Axon in depressed portion of neurolemmocyte
Not wrapped in repeated layers
In CNS,
unmyelinated axons not associated with oligodendrocytes

Unmyelinated Axons (Figure 12.8)


Copyright The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

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

Nervous TissueGlial Cells:


Myelination
What is the function of the myelin sheath?
The myelin sheath provides a protective insulating
covering around the axon. It prevents the passage of
ions through the axonal membrane and allows for
faster action potential propagation.

Nervous TissueGlial Cells:


Myelination
Clinical View: Nervous System Disorders
Affecting Myelin
Multiple Sclerosis

progressive demyelination of neurons in CNS


autoimmune disorder
oligodendrocytes attacked by immune cells
repeated inflammatory events causing scarring and permanent loss of
function
vision problems, muscle weakness and spasms, urinary and bladder
problems, mood problems

Nervous TissueGlial Cells:


Myelination
Clinical View: Nervous System Disorders
Affecting Myelin (continued)
Guillain-Barre syndrome

loss of myelin from peripheral nerves due to inflammation


muscle weakness that begins in distal limbs
advances to involve proximal muscles
no specific infectious agent identified
most function recovered with little medical intervention

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

Skeletal muscle fibers


2 The proximal portion of each severed
axon seals off and swells. The distal
portion of axon and myelin sheath
disintegrate; the neurilemma survives.

Regeneration
of PNS axons
(Figure
12.9)

Endoneurium Sealed, swollen


end of axon

Neurilemma

Neurilemma and endoneurium


form a regeneration tube.

4 Axon regenerates and


remyelination occurs.

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)
Copyright The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

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

Na+/K+ pump changes


shape (requires energy
from ATP breakdown)

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+

(b) Leak (passive) channels

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)
Copyright The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

Closed

Open

Neurotransmitter
binds to gate

K+

(c) Chemically gated channels

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)
Copyright The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

Closed (resting state)

+ + + +

+ + + +



Inactivation
gate (open) Activation
gate (closed)
Na+

Open (activation state)

Closed (inactivation state)

+ + +
Inactivation
gate (open)
Na+

+ + + +

+ + + +
Inactivation
gate (closed)

+ + + +

(d) Voltage-gated channels (three different states)

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

Copyright The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

Plasma membrane of entire neuron

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.

Ultrastructure of Neurons: Distribution of


Substances and Membrane Potentials
Distribution of substances inside and outside neuron
Essential for neuron function
More prevalent within cytosol
negatively charged phosphate ions (e.g., in ATP)
negatively charged proteins
K+
More prevalent in interstitial fluid
Na+
Cl-

Ultrastructure of Neurons: Distribution of


Substances and Membrane Potentials
Movement of substances and membrane potentials
Ions able to pass through membrane by transport proteins
Phosphate-containing molecules and proteins
generally restricted from crossing
Net movement dependent on electrochemical gradient
combination of the electrical and chemical gradient

Ultrastructure of Neurons: Distribution of


Substances and Membrane Potentials
Movement of substances and membrane potentials
(continued)
Electrical gradient
difference in electrical charge between two areas
cells with differences in total positive and negative charges across
membrane
exhibit electrical gradient at the membrane
inside relatively negative
outside relative positive
difference in charge termed a membrane potential
can be altered to create electrical currents

Ultrastructure of Neurons: Distribution of


Substances and Membrane Potentials
Movement of substances and membrane potentials
(continued)
Chemical concentration gradient
unequal distribution between two areas
each substance with own chemical concentration gradient
e.g., K+ with a higher concentration inside the neuron
e.g., Na+ with a higher concentration outside the neuron

Ultrastructure of Neurons: Distribution of


Substances and Membrane Potentials
What is an electrical gradient? What is a
chemical gradient?
An electrical gradient is a difference in electrical
charge between two areas.
A chemical gradient is an unequal distribution of a
substance between two areas.

Introduction to Neuron Physiology


Learning Objectives:
1) Integrate the concepts of voltage, current, and resistance with
neuron structure and function.
2) Define resting membrane potential, and state its typical value
for neurons.
3) Describe how the resting membrane potential is established
and maintained in neurons.
4) Explain depolarization and hyperpolarization.
5) Compare and contrast graded potentials and action potentials.

Introduction to Neuron Physiology:


Neurons and Ohms Law
Concepts for electrical current in neurons
Neuron activity dependent upon electrical current
Voltage
measure of the amount of difference in electrical charge
measured in volts or millivolts
indicator of relative potential energy
Current
movement of charged particles
greater movement, greater current
can be harnessed to do work

Introduction to Neuron Physiology:


Neurons and Ohms Law
Concepts for electrical current in neurons (continued)
Resistance
opposition to movement of charged particles
Ohms law
current = voltage/resistance
greater current possible with larger voltage and smaller resistance

Introduction to Neuron Physiology:


Neurons and Ohms Law
Electrical currents in neurons
Charged particles, ions
Difference in charge distribution at the plasma membrane
termed membrane potential
Normally resistance present at the plasma membrane

Introduction to Neuron Physiology:


Neurons and Ohms Law
Electrical currents in neurons (continued)
Resistance across membrane altered through ion channels
decreased when ion channels open
increased when ion channels closed
Current generated when ions diffuse across membrane

Introduction to Neuron Physiology:


Resting Membrane Potentials
Resting membrane potential
Membrane potential in a resting, excitable cell
Relative difference in charge across membrane
Measured with a voltmeter
microelectrodes into neuron and interstitial fluid
Negative value, typically -70 mV
More positive ions outside a neuron than in it at rest
A consequence of the plasma membrane permeability to ions

Introduction to Neuron Physiology:


Resting Membrane Potentials
Establishing and maintaining resting membrane
potentials
The role of K+

K+ diffusion the most important factor in specific value of RMP


Dependent on the electrochemical gradient
Outward movement facilitated by steep concentration gradient
Leaves relatively more negatively charged structures inside

Introduction to Neuron Physiology:


Resting Membrane Potentials
Establishing and maintaining resting membrane
potentials
The role of K+ (continued)
Outward movement opposed by electrical gradient
attraction of negative inside of cell for K+
becomes greater as more K+ diffuses out

At equilibrium
chemical concentration gradient equal to electrical gradient opposing
movement

RMP would be -90 mV


if only K+ channels were present

Introduction to Neuron Physiology:


Resting Membrane Potentials
Establishing and maintaining resting membrane
potentials (continued)
The role of Na+
Typical neuron RMP -70 mV
Difference between -90 mV of K+ movement only
due to Na+ movement
Enters cell through Na+ leak channels
Moves down concentration gradient
Also pulled by electrical gradient
Channels present in limited numbers

Introduction to Neuron Physiology:


Resting Membrane Potentials
Establishing and maintaining resting membrane
potentials (continued)
The role of Na+/K+ Pumps
Play relatively small role in establishing RMP
Three Na+ pumped out for two K+ pumped in
More significant role in maintaining gradients of K + and Na+
following diffusion as part of neurons electric current
Ions pumped back up concentration gradient by pump
Two-thirds of a neurons energy expenditure

Establishing and Maintaining the Resting Membrane Potential


(RMP) (Figure 12.12)
Copyright The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

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

Introduction to Neuron Physiology:


Resting Membrane Potentials
The resting membrane potential is
primarily established by what two
structures embedded in the plasma
membrane?
Sodium ion and potassium ion leak channels

Introduction to Neuron Physiology:


Changing the Membrane Potential
Depolarization and hyperpolarization

Opening of chemically gated channels or voltage-gated channels


Causes change in ion flow across membrane
Alters resting membrane potential
Depolarization
inside of cell becomes more positive than RMP
e.g., from -70 mV to -60 mV
occurs when gated channels open
movement of Na+ into neuron
causes inside of neuron to become more positive

Introduction to Neuron Physiology:


Changing the Membrane Potential
Depolarization and hyperpolarization (continued)
Hyperpolarization
inside of cell becomes more negative
e.g., from -70 mV to -80 mV
may result from opening of gated K+ channels
may result from opening of gated Cl- channels
loss of positive ion (K+) or gain of negatively charged ion (Cl-)

Changing the Resting Membrane Potential: Resting


Membrane Potential (Figure 12.13a)
Copyright The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

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

Changing the Resting Membrane Potential:


Depolarization (Figure 12.13b)
Copyright The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

60 mV
+

+ + + + + +


+ + + + + +

Gated Na+ channel


Cl

+ +

+ + + + +

+ + + +

+ +

Na+
K+
(b) Depolarization: Na+ flows in

e.g., 60 mV

Changing the Resting Membrane Potential:


Hyperpolarization (Figure 12.13c)
Copyright The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

80 mV
+

Na+

+ + + + + +


+ + + + + +
Gated K+
channel

Gated Cl
channel

K+
Cl

+ +

+ + + + +

+ + + +

+ +

e.g., 80 mV
(c) Hyperpolarization: K+ flows out or Cl flows in

Introduction to Neuron Physiology:


Changing the Membrane Potential
Graded potentials versus action potentials
Graded potentials

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

Introduction to Neuron Physiology:


Changing the Membrane Potential
Graded potentials versus action potentials
Graded potentials (continued)
May result in depolarization or hyperpolarization
depends on channel that opens
Degree dependent on stimulus magnitude
larger stimulus opening more chemically gated channels
flow of more ions
Decreases in intensity with distance along the membrane
Short-lived
lasts until local ion current ceases

Introduction to Neuron Physiology:


Changing the Membrane Potential
Graded potentials versus action potentials
Action potentials

Generated within the initial segment


Propagated along axon
Due to opening of voltage-gated channels
Threshold value
minimum voltage change to open voltage-gated channel
any value below this, a subthreshold value

Introduction to Neuron Physiology:


Changing the Membrane Potential
Graded potentials versus action potentials
Action potentials (continued)
If threshold value reached
channels open and membrane potential reversed
if Na+ channel opens, enters the neuron
makes inside relatively positive
flow of local current to adjacent areas

Introduction to Neuron Physiology:


Changing the Membrane Potential
Graded potentials versus action potentials
Action potentials
If threshold value reached (continued)
opening of voltage-gated channels in these areas
successive opening down the axon
followed by sequential opening of voltage-gated K + channels
movement of K+ out of neuron returns membrane to RMP

Introduction to Neuron Physiology:


Changing the Membrane Potential
Graded potentials versus action potentials
Action potentials: (continued)
involve temporary reversal of polarity across plasma membrane
inside becomes relatively positive
followed by a return to RMP

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

Introduction to Neuron Physiology:


Changing the Membrane Potential
How does depolarization and
hyperpolarization occur in a neuron?
Depolarization occurs when cation or voltage-gated
Na+ channels open, allowing positively charged Na+ to
move into the neuron.
Hyperpolarization occurs when gated K+ channels
open and allow K+ to move out of the neuron, or when
gated Cl- channels open and allow Cl- to move into the
cell.

Introduction to Neuron Physiology:


Changing the Membrane Potential
How does a graded potential differ from an
action potential in terms of the types of
channels involved and where it occurs?
Graded potentials involve chemically gated channels
at dendrites and the cell body.
Action potentials involved voltage-gated channels at
the axon.

Physiologic Events in the Neuron Segments


Learning Objectives:
1) Define a postsynaptic potential.
2) Compare and contrast the action of excitatory and inhibitory
neurotransmitters in developing postsynaptic potentials
(graded potentials) in the receptive segment.
3) Graph and explain an excitatory postsynaptic potential (EPSP)
and an inhibitory postsynaptic potential (IPSP).
4) Define summation, and describe the two types of summation
that can occur in the initial segment.
5) Describe the initiation and propagation of depolarization and
repolarization, which constitutes a nerve signal.

Physiologic Events in the Neuron Segments


Learning Objectives: (continued)
6) Graph and explain the electrical changes that occur in an axon.
7) Define refractory period, and explain the difference between
the absolute refractory period and relative refractory period
associated with transmitting an action potential.
8) Describe events that occur when the propagated action
potential reaches the transmissive segment
9) Explain the role of Ca2+ in neurotransmitter release.

Physiologic Events in the Neuron Segments:


Receptive Segment
Postsynaptic potentials

Graded potentials that occur in postsynaptic neurons


Occur after release of neurotransmitter from presynaptic neuron
Opening of gated channels after binding of neurotransmitters
Results in small local potential

Physiologic Events in the Neuron Segments:


Receptive Segment
Postsynaptic potentials (continued)
Postsynaptic neuron
able to bind many neurotransmitters at once
numerous postsynaptic potentials generated at once
Type of graded potential formed depends on neurotransmitter
excitatory or inhibitory neurotransmitter

Physiologic Events in the Neuron Segments:


Receptive Segment
Generation of EPSPs
Sequence of events
1) Excitatory neurotransmitter crosses synaptic cleft.
binds to receptor
opens a chemically gated cation channel
2)
More Na+ moves into neuron than K+ moves out.

Physiologic Events in the Neuron Segments:


Receptive Segment
Generation of EPSPs
Sequence of events (continued)
3) Inside becomes slightly more positive.
less negative state called excitatory postsynaptic potential
(EPSP)
4) Local current of Na+ becomes weaker
decreases in intensity with distance traveled

Physiologic Events in the Neuron Segments:


Receptive Segment
Generation of EPSPs (continued)
Degree of change in RMP
dependent on amount of neurotransmitter bound per unit time
More excitatory neurotransmitter released
more cation channels open
greater change in the positive direction
e.g., from -70 mV to -65 mV

Release of Excitatory Neurotransmitter and


Generation of EPSP (Figure 12.15a)
Copyright The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

Axons of
presynaptic
neuron

Postsynaptic
neuron

Release of excitatory neurotransmitter and generation of EPSP


1 Excitatory neurotransmitter released from presynaptic neuron binds to
receptors, which are chemically gated cation channels, causing them to open.
Axons of
presynaptic
neuron

Excitatory
neurotransmitter

Postsynaptic neuron
0

Chemically gated cation channel


2 Na+ flows into neuron.
Na+
3 Inside of neuron becomes
more positive (less negative);
called EPSP (e.g., 68 mV).

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

Physiologic Events in the Neuron Segments:


Receptive Segment
Generation of IPSPs
Sequence of events
1) Inhibitory neurotransmitter crosses synaptic cleft.
binds to chemically gated K+ channel or Cl- channel
depends on neurotransmitter and channels present
2) If neurotransmitter binds K+ channel, K+ moves out of neuron.
If neurotransmitter binds Cl-channel, Cl- flows into neuron.

Physiologic Events in the Neuron Segments:


Receptive Segment
Generation of IPSPs
Sequence of events (continued)
3) Inside of the cell becomes slightly more negative
more negative state termed inhibitory postsynaptic potential
(IPSP)
4) Local current of ions becomes weaker.
decreases in intensity with distance traveled toward initial
segment

Release of Inhibitory Neurotransmitter and


Generation of IPSP (Figure 12.15b)
Copyright The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

Release of inhibitory neurotransmitter and generation of IPSP

1
Axons of
presynaptic
neuron

Inhibitory neurotransmitter binds to either chemically gated K+


channels or chemically gated Cl channels, causing them to open.
Inhibitory
neurotransmitter

Postsynaptic neuron
0

Chemically gated K+ channel


Chemically gated Cl channel

20

2 Either K+ flows out of, or Cl


flows into, the neuron,
depending on the type of
channel stimulated.

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

Physiologic Events in the Neuron Segments:


Receptive Segment
Simultaneous release
Excitatory and inhibitory neurotransmitters
may be simultaneously released from different neurons
Varied frequency of releasing neurotransmitter
Result: many EPSPs, many IPSPs, or both

Several Presynaptic Neurons


with a Postsynaptic Neuron(Figure 12.16)
Copyright The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

EPSP

Postsynaptic
neuron

Synaptic
knob

Presynaptic
axons

Axons of presynaptic
neuron
Dendrites
Cell body of
postsynaptic neuron

Myelin
sheath

Axon

Axons of presynaptic neuron


b: Science VU/Lewis-Everhart-Zeevi/Visuals Unlimited

SEM 80,000x

IPSP

Physiologic Events in the Neuron Segments:


Receptive Segment
How are EPSP graded potentials established
in the receptive segment of a neuron?
Excitatory neurotransmitters bind chemically gated
cation channels. More Na+ moves into neuron than
K+ moves out. The inside of the cell becomes
slightly more positive.

Physiologic Events in the Neuron Segments:


Initial Segment
Summation
Addition of graded postsynaptic potentials (IPSPs and EPSPs)
Occurs at the initial segment
Determines if threshold membrane potential is reached
-55 mV, +15 mV from RMP
If threshold reached
voltage-gated channels open
action potential generated that travels along axon

Physiologic Events in the Neuron Segments:


Initial Segment
Summation (continued)
IPSPs negate effects of EPSPs
Thousands of EPSPs required to reach threshold
must arrive at nearly the same time
Spatial summation
release of neurotransmitter from multiple presynaptic neurons
action potential initiated if enough EPSPs generated

Physiologic Events in the Neuron Segments:


Initial Segment
Summation (continued)
Temporal summation
repeated release of excitatory neurotransmitter at same location
effects added if occur within small timeframe
action potential initiated if threshold reached

Copyright The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

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

Axons of presynaptic neurons (P), (P1P5)


(a) Spatial summation

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

(b) Temporal summation

Axon

Physiologic Events in the Neuron Segments:


Initial Segment
All or none law

If threshold reached, action potential propagated


If threshold not reached, not propagated
Same intensity of response to values greater than threshold
Similar to what occurs with a gun
with sufficient pressure on trigger, gun fired
with insufficient pressure on trigger, not fired
travels at same velocity even if pressure is greater than needed

Physiologic Events in the Neuron Segments:


Initial Segment
What is the significance of the threshold
membrane potential in the initial segment of
a neuron?
If the threshold membrane potential is reached, it
initiates an action potential. If it is not reached, no
action potential is generated.

Physiologic Events in the Neuron Segments:


Conductive Segment
Nerve signal

The propagation of an action potential


Involves depolarization and repolarization
Threshold reached in initial segment
Initiates action potential along the axon (conductive segment)

Physiologic Events in the Neuron Segments:


Conductive Segment
Depolarization and its propagation
Positive change in membrane potential
Occurs only at plasma membrane
via voltage-gated Na+ channels
Channels triggered to open when threshold reached
Rapid entry of Na+
Inside of axon made positive
Channels open briefly before closing
change from activation state to temporary inactivation state

Physiologic Events in the Neuron Segments:


Conductive Segment
Depolarization and its propagation (continued)
Propagation of depolarization
Sequential opening of voltage-gated Na+ channels along the axon
Flow of Na+ into cell
causes adjacent regions to also reach threshold
triggers voltage-gated Na+ channels in these areas
Process repeated rapidly down synaptic knob
does not go backwards
voltage-gated Na+ channels here in inactivated state

Physiologic Events in the Neuron Segments:


Conductive Segment
Depolarization and its propagation (continued)
Local anesthetics

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

Propagation of Action Potential Down an Axon (Figure 12.18a)


Copyright The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

++ +
+

Nerve signal:
propagation of
action potential

++
+ + ++ + + + + + ++ + + + ++ + + + + + ++ + + + ++ + + + + + ++ + +
Axon
+ ++ +
hillock
+ ++ +
++ ++ + + + + + ++ + + + ++ + + + + + ++ + + + ++ + + + + + ++ + +
+
+
Repolarization Depolarization
+
+
+ +

(a)

Propagation of Action Potential: Depolarization (Figure 12.18b)


Copyright The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

Depolarization: Consecutive voltage-gated Na+ channels go through the


following stages: open, closed (inactivation state), closed (resting state)

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)

Physiologic Events in the Neuron Segments:


Conductive Segment
Repolarization and its propagation
Return to resting membrane potential

Voltage-gated K+ channels normally closed


Stimulated to open by threshold
Not open until depolarization has ended
Exit of K+, making inside of axon negative
Return to RMP (-70 mV)
Triggers voltage-gated Na+ channels to return to resting state

Physiologic Events in the Neuron Segments:


Conductive Segment
Repolarization and its propagation (continued)
Propagation of repolarization
Opening of voltage-gated K+ channels adjacent
Open sequentially along length of axon

Propagation of Action Potential: Repolarization (Figure 12.18c)


Copyright The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

Repolarization: Consecutive voltage-gated K + channels go through the


following stages: open and closed

K+
+ + + + + + + + + + + +


Closed

70
mv

+ + + + + + + + + +

+ +
Open

+30 + + + + +
mv
Closed

K+ channels open and


K+ diffuses out; RMP
(70 mv) is reestablished
(c)

Physiologic Events in the Neuron Segments:


Conductive Segment
Hyperpolarization and return to resting
membrane potential
Voltage-gated K+ channels
open longer than time needed to reestablish RMP
Inside of neuron briefly more negative than RMP
hyperpolarized
Closure of K+ channels
RMP reestablished by Na+/K+ pumps

Physiologic Events in the Neuron Segments:


Conductive Segment
Refractory period
Brief time period after action potential initiated
During absolute refractory period
no amount of stimulus able to generate a second action potential
Na+ channels opened then closed in inactivated state
remain closed until potential almost to resting potential
ensures that action potential moves in one direction only

Physiologic Events in the Neuron Segments:


Conductive Segment
Refractory period (continued)
During relative refractory period
with greater stimulation, action potential possible
Na+ returned to resting state
neuron hyperpolarized
due to extended time K+ channels remain open

Events of an Action Potential (Figure 12.19)


Copyright The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

The unstimulated axon has a resting membrane


potential of 70 mV.

Graded potentials reach axon hillock and are added together.

Depolarization occurs when the threshold (55 mV) is


reached; voltage-gated Na+ channels open and Na+ enters
rapidly, reversing the polarity from negative to positive
(55 mV
+30 mV).

Repolarization occurs due to closure of voltage-gated


Na+ channels (inactivation state) and opening of
voltage-gated K+ channels. K+ moves out of the cell into
the IF and polarity is reversed from positive to negative
(+30 mV
70 mV).

Hyperpolarization occurs when voltage-gated K+ channels


stay open longer than the time needed to reach the resting
membrane potential; during this time the membrane potential
is less than the resting membrane potential of 70 mV.

Voltage-gated K+ channels are closed, and the plasma


membrane has returned to resting conditions by activity
of Na+/K+ pumps.

+30
+10
0

mV

10
30

50

Threshold

70
90

Resting
membrane
potential
0

1
Time (msec)

Physiologic Events in the Neuron Segments:


Conductive Segment
What type of channels are sequentially
opened in the propagation of the action
depolarization? In the propagation of
repolarization?
Voltage-gated Na+ channels
Voltage-gated K+ channels

Physiologic Events in the Neuron Segments:


Transmissive Segment
Activity at the synaptic knob
Calcium concentration gradient established by pumps
more calcium outside synaptic knob than in
Voltage-gated Ca2+ channels
triggered by propagated action potential
movement of calcium ions into synaptic knob

Physiologic Events in the Neuron Segments:


Transmissive Segment
Activity at the synaptic knob (continued)
Binding of calcium to proteins of synaptic vesicles
Triggers fusion of synaptic vesicles with neuron plasma membrane
Neurotransmitters released into synaptic cleft by exocytosis
facilitated by numerous proteins
diffuse across cleft
bind to specific receptors on cell to be stimulated
See Figure 12.22: Events of Neuron Physiology

Copyright The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

Neuromuscular junction

Transmissive
Segment:
Release of
Neurotransmitter
(Figure
12.21a)

Voltage-gated Ca2+ channel


Synaptic cleft
Receptor
Ca2+
Voltage-gated Ca2+
channels open and
Ca2+ enters the
synaptic knob
and binds with proteins
of synaptic vesicles.

+
+ + ++ +
1

++

Action potential reaches


synaptic knob.

++ + + +
++
+

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.)

Physiologic Events in the Neuron Segments:


Transmissive Segment
What is the sequence of events from the
arrival of an action potential at the synaptic
knob until the release of neurotransmitter
into the synaptic cleft?
Voltage-gated calcium ion channels are triggered to
open by an action potential. Calcium ions move into
the synaptic knob. They bind to proteins of synaptic
vesicles, resulting in fusion of vesicles with neuron
plasma membrane. Neurotransmitter is released.

Velocity of a Nerve Signal


Learning Objectives:
1) Compare and contrast continuous conduction and saltatory
conduction in the mechanism and velocity of transmission of
an action potential.
2) Identify the criteria used to distinguish the groups of nerve
fibers.

Velocity of a Nerve Signal


Factors influencing velocity of nerve signal
Diameter of axon
larger diameter, faster the velocity of the signal
Myelination of axon
more important factor
faster velocity in myelinated axons

Velocity of a Nerve Signal: Propagation


Continuous conduction
Occurs in unmyelinated axons
Sequential opening of voltage-gated Na+ and K+ channels

Velocity of a Nerve Signal: Propagation


Saltatory conduction
Occurs in myelinated axons
Action potentials propagated only at neurofibril nodes
Myelinated regions
with limited numbers of voltage gated Na+ and K+ channels
well insulated, preventing ion movement
Neurofibril nodes
with large number of voltage-gated Na+ and K+ channels
lack myelin insulation

Velocity of a Nerve Signal: Propagation


Nerve signal in myelinated axon
Neurofibril node
initiation of action potential
diffusion of Na+ into axon
Myelinated regions
diffusion through axoplasm of axon
relatively fast
becomes weaker with distance as experiences resistance

Velocity of a Nerve Signal: Propagation


Nerve signal in myelinated axon (continued)
Next neurofibril node
arrival of weak Na+ current
sufficient to cause opening of voltage-gated Na + channels
new action potential initiated
Repetition of process
continuation of nerve signal to synaptic knob

Velocity of a Nerve Signal: Propagation


Nerve signal in myelinated axon (continued)
Increased speed in myelinated axons
action potential generated only at neurofibril nodes
More efficient
less energy required for Na+/K+ pump to maintain RMP

Saltatory Conduction (Figure 12.23)

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Neurofibril node

Na+

K+

Myelin sheath

Diffusion of Na+
through axoplasm

+ + + + +

+ + + + +

+ + +
+ + +

+ + +


+ + + + +


+ + + + +

+ + +
+ + +


+ + +

Action
potential
Repolarization

Depolarization

Velocity of a Nerve Signal: Propagation


How does conduction of an action potential
in an unmyelinated axon and myelinated
axon differ?
In an unmyelinated axon action potentials occur
down the whole length of the axon. In an
unmyelinated axon, action potentials only occur at
neurofibril nodes. In myelinated regions Na+ quickly
diffuses through axoplasm, initiating action
potentials at the next neurofibril node.

Velocity of a Nerve Signal:


Nerve Fiber Classification
Nerve fiber groups
Nerve fiber: an axon and its myelin sheath
Classified into three major groups
Group A
conduction velocity as fast as 150 m/sec
large diameter myelinated fibers
e.g., most somatic sensory neurons, somatic motor neurons

Velocity of a Nerve Signal:


Nerve Fiber Classification
Nerve fiber groups (continued)
Group B and Group C
small in diameter, unmyelinated, or both
e.g., sensory and motor visceral neurons
group B: 15 m/sec
group C: 1 m/sec

Velocity of a Nerve Signal:


Nerve Fiber Classification
What are the general characteristics of
group A nerve fibers, and what functions do
they normally serve?
Group A nerve fibers have the fastest conduction
velocity. This is due to their large diameter and their
myelination.

Neurotransmitters and Neuromodulation


Learning Objectives:
1) Identify the four classes of neurotransmitters, and give
examples of their actions.
2) Explain the two ways in which neurotransmitters are removed
from the synaptic cleft.
3) Define neuromodulation, including facilitation and inhibition.

Neurotransmitters and Neuromodulation


Classes of neurotransmitters

Neurotransmitters, various small organic compounds


Released at synaptic cleft
Approximately 100 known
Classified into major groups

Neurotransmitters and Neuromodulation


Classes of neurotransmitters (continued)
Acetylcholine
excitatory or inhibitory neurotransmitter
released in both CNS and PNS
molecule released from motor neuron at neuromuscular junction
Amino acids
building blocks of proteins
some also neurotransmitters
e.g., glutamate, glycine, aspartate

Neurotransmitters and Neuromodulation


Classes of neurotransmitters (continued)
Monoamines
derived from certain amino acids
carboxyl group removed and functional group added
subgroup added determines type
includes subgroup, catecholamines (norepinephrine, epinephrine,
dopamine)
Neuropeptides
chains of amino acids
include enkephalins and somatostatin
See Table 12.4: Neurotransmitters

Neurotransmitters and Neuromodulation


Removal of neurotransmitters from the synaptic
cleft
Temporary association between neurotransmitter and receptor
Necessary to eliminate molecule after stimulation
Can occur by degradation
neurotransmitter chemically inactivated in synaptic cleft
e.g., breakdown of ACh by acetylcholinesterase

Neurotransmitters and Neuromodulation


Removal of neurotransmitters from the synaptic
cleft (continued)
Can occur by reuptake
neurotransmitter reabsorbed by transport protein in presynaptic
neuron
recycled into another synaptic vesicle for reuse
e.g., drugs, selective serotonin reuptake inhibitors
block reuptake of serotonin and used in treatment of depression

Neurotransmitters and Neuromodulation


What are the four primary classes of
neurotransmitters?
Acetylcholine, amino acids, monoamines,
neuropeptides

Neurotransmitters and Neuromodulation:


Neuromodulation
Neuromodulators

Chemical released from cells


Locally regulate or alter response of neurons to neurotransmitters
Release termed neuromodulation
Facilitation
occurs when greater response in postsynaptic neuron
may increase amount of neurotransmitter in synaptic cleft
may increase number of receptors on postsynaptic neurons

Neurotransmitters and Neuromodulation:


Neuromodulation
Neuromodulators (continued)
Inhibition
occurs when less response from postsynaptic neuron
may decrease amount of neurotransmitter
may decrease number of receptors on postsynaptic neuron

Neurotransmitters and Neuromodulation:


Neuromodulation
What is the term for the type of
neuromodulation that results in a greater
response from a postsynaptic neuron? A
lesser response?
Facilitation
Inhibition

Neural Integration and


Neuronal Pools of the CNS
Learning Objective:
1) Identify the four different types of neuronal pools, and explain
how they function.

Neural Integration and


Neuronal Pools of the CNS
Neuronal pools (neuronal circuits)
Complex patterns of grouped interneurons
Four types of circuits:
converging, diverging, reverberating, parallel-after-discharge
Pool may be localized or distributed in several regions of CNS
All restricted in number of input sources and output destinations

Copyright The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

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)

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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

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Types of circuits (continued)


Reverberating circuits
utilize feedback to produce
repeated, cyclical stimulation
once activated, may continue to
function until cycle is broken
broken by inhibitory stimuli or
synaptic fatigue
due to exhaustion of
neurotransmitter in
presynaptic cell
e.g., circuits that keep us
breathing during sleep
(Figure 12.24c)

Input

Output

(c) Reverberating
circuit

Types of circuits (continued)


Parallel-after-discharge circuits
input transmitted
simultaneously along several
pathways to common
postsynaptic cell
vary in number of neurons in
pathway
due to synaptic delay, signal
arriving at postsynaptic cell at
various times
believed to be involved in
higher-order thinking
(Figure 12.24d)

Copyright The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

Input

Output
(d) Parallel-after-discharge
circuit

Neural Integration and


Neuronal Pools of the CNS
How are neurons arranged in a converging
circuit?
Inputs converge (come together) at a single
postsynaptic neuron.

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