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Nervous Tissue
Fundamentals
of the Nervous
P A R and
T A
System
Nervous
Tissue
1
Nervous System
The master controlling and
communicating system of the body
Functions
Sensory input –the stimuli goes to
CNS
Integration – interpretation of
sensory input
Motor output – response to
stimuli coming from CNS
2
Nervous System
3
Figure 11.1
Organization of the Nervous
System
Central nervous system (CNS)
Brain and spinal cord
Integration and command center
Peripheral nervous system
(PNS)
Paired spinal and cranial nerves
Carries messages to and from the
spinal cord and brain
4
Peripheral Nervous System (PNS):
Two Functional Divisions
Sensory (afferent) division
Somatic sensory fibers – carry
impulses from skin, skeletal muscles,
and joints to the brain
Visceral sensory fibers – transmit
impulses from visceral organs to the
brain
5
Motor Division: Two Main Parts
Motor (efferent) division
Somatic motor nervous
system (voluntary)- carries
impulses from the CNS to skeletal
muscles. Conscious control
6
Motor Division: Two Main Parts
Visceral motor nervous system
or Autonomic nervous system
(ANS) (involuntary) - carry
impulses from the CNS to smooth
muscle, cardiac muscle, and glands
Sympathetic and
Parasympathetic
Enteric Nervous System (ENS)
7
Histology of Nerve Tissue
The two principal cell types of the
nervous system are:
Neurons – excitable cells that
transmit electrical signals
Neuroglias (glial) – cells that
surround and wrap neurons
8
Supporting Cells: Neuroglia
Neuroglia
Provide a supportive scaffolding
for neurons
Segregate and insulate neurons
Guide young neurons to the
proper connections
Promote health and growth
9
Neuroglia of the CNS
Astrocytes
Most abundant, versatile, and highly
branched glial cells
Maintain blood-brain barrier
They cling to neurons and their
synaptic endings,
They wrap around capillaries
Regulate their permeability
10
Neuroglia of the CNS
Provide structural framework for the
neuron
Guide migration of young neurons
Control the chemical environment
Repair damaged neural tissue
11
Astrocytes
12
Figure 11.3a
Neuroglia of the CNS
Microglia – small, ovoid cells with
spiny processes
Phagocytes that monitor the
health of neurons
Ependymal cells – range in shape
from squamous to columnar
They line the central cavities of
the brain and spinal column
13
Microglia and Ependymal Cells
14
Figure 11.3b, c
Neuroglia of the CNS
Oligodendrocytes – branched cells
Myelin
Wraps of oligodendrocytes
processes around nerve fibers
Insulates the nerve fibers
15
Neuroglia of the PNS
Schwann cells (neurolemmocytes) –
Myelin
Wraps itself around nerve fibers
Insulates the nerve fibers
Satellite cells surround neuron cell
bodies located within the ganglia
Regulate the environment around the
neurons
16
Oligodendrocytes, Schwann
Cells, and Satellite Cells
17
Figure 11.3d, e
Neurons (Nerve Cells)
Structural units of the nervous
system
Composed of a body, axon, and
dendrites
Long-lived, amitotic, and have a
high metabolic rate
Their plasma membrane function in:
Electrical signaling
Cell-to-cell signaling during
18
development
Neurons (Nerve Cells)
19
Figure 11.4b
Nerve Cell Body (Perikaryon or
Soma)
Contains the nucleus and a
nucleolus
Is the major biosynthetic center
Is the focal point for the outgrowth
of neuronal processes
Has no centrioles (hence its amitotic
nature)
Has well-developed Nissl bodies
(rough ER)
Contains an axon hillock – coneshaped area from which axons arise20
Processes
Arm like extensions from the soma
There are two types of processes:
axons and dendrites
Myelinated axons are called tracts
in the CNS and nerves in the PNS
21
Dendrites: Structure
Short, tapering processes
They are the receptive, or input,
regions of the neuron
Electrical signals are conveyed as
graded potentials (not action
potentials)
22
Neurons (Nerve Cells)
23
Figure 11.4b
Axons: Structure
Slender processes of uniform
diameter arising from the hillock
Long axons are called nerve fibers
Usually there is only one
unbranched axon per neuron
Axon collaterals
Telodendria
Axonal terminal or synaptic
knobs
24
Axons: Function
Generate and transmit action
potentials
Secrete neurotransmitters from the
axonal terminals
Movement along axons occurs in
two ways
Anterograde — toward the axon
terminal
Retrograde — toward the cell
body
25
Myelin Sheath
Whitish, fatty (protein-lipoid),
segmented sheath around most
long axons
It functions to:
Protect the axon
Electrically insulate fibers from
one another
Increase the speed of nerve
impulse transmission
26
Myelin Sheath and Neurilemma:
Formation
Formed by Schwann cells in the PNS
A Schwann cell:
Envelopes an axon
Encloses the axon with its plasma
membrane
Has concentric layers of
membrane that make up the
myelin sheath
27
Myelin Sheath and Neurilemma:
Formation
28
Figure 11.5a–c
Nodes of Ranvier (Neurofibral
Nodes)
Gaps in the myelin sheath between
adjacent Schwann cells
They are the sites where axon
collaterals can emerge
29
Unmyelinated Axons
A Schwann cell surrounds nerve
fibers but coiling does not take
place
Schwann cells partially enclose 15
or more axons
Conduct nerve impulse slowly
30
Axons of the CNS
Both myelinated and unmyelinated
fibers are present
Myelin sheaths are formed by
oligodendrocytes
Nodes of Ranvier are widely spaced
31
Regions of the Brain and Spinal
Cord
White matter – dense collections
of myelinated fibers
Gray matter – mostly soma,
dendrites, glial cells and
unmyelinated fibers
32
Neuron Classification
Structural:
Multipolar — three or more
processes
Bipolar — two processes (axon
and dendrite)
Unipolar — single, short process
33
A Structural Classification of
Neurons
34
Figure 12.4
Neuron Classification
Functional:
Sensory (afferent) — transmit
impulses toward the CNS
Motor (efferent) — carry impulses
toward the body surface
Interneurons (association
neurons) — any neurons between
a sensory and a motor neuron
35
Comparison of Structural
Classes of Neurons
36
Table 11.1.1
Comparison of Structural
Classes of Neurons
37
Table 11.1.2
Neurophysiology
Neurons are highly irritable
Action potentials, or nerve
impulses, are:
Electrical impulses carried along
the length of axons
Always the same regardless of
stimulus
38
Electricity Definitions
Voltage (V) – measure of potential
energy generated by separated
charge
Potential difference – voltage
measured between two points
Current (I) – the flow of electrical
charge between two points
Resistance (R) – hindrance to charge
flow
39
Electricity Definitions
Insulator – substance with high
electrical resistance
Conductor – substance with low
electrical resistance
40
Electrical Current and the Body
Reflects the flow of ions rather than
electrons
There is a potential on either side of
membranes when:
The number of ions is different
across the membrane
The membrane provides a
resistance to ion flow
41
Role of Ion Channels
Types of plasma membrane ion
channels:
Nongated, or leakage channels
– always open
Chemically gated channels –
open with binding of a specific
neurotransmitter
42
Role of Ion Channels
Voltage-gated channels – open
and close in response to
membrane potential.
2 gates
Mechanically gated channels –
open and close in response to
physical deformation of receptors
43
Gated Channels
44
Figure 12.13
Operation of a Chemically Gated
Channel
Example: Na+-K+ gated channel
Closed when a neurotransmitter is
not bound to the extracellular
receptor
Na+ cannot enter the cell and K+
cannot exit the cell
Open when a neurotransmitter is
attached to the receptor
Na+ enters the cell and K+ exits
the cell
45
Operation of a Voltage-Gated
Channel
Example: Na+ channel
Closed when the intracellular
environment is negative
Na+ cannot enter the cell
Open when the intracellular
environment is positive
Na+ can enter the cell
46
Gated Channels
When gated channels are open:
Ions move quickly across the
membrane
Movement is along their
electrochemical gradients
An electrical current is created
Voltage changes across the
membrane
47
Electrochemical Gradient
Ions flow along their chemical
gradient when they move from an
area of high concentration to an
area of low concentration
Ions flow along their electrical
gradient when they move toward an
area of opposite charge
Electrochemical gradient – the
electrical and chemical gradients
taken together
48
Resting Membrane Potential (Vr)
The potential difference (–70 mV)
across the membrane of a resting
neuron
It is generated by different
concentrations of Na+, K+, Cl, and
protein anions (A)
49
Resting Membrane Potential (Vr)
Ionic differences are the
consequence of:
Differential permeability of the
neurilemma to Na+ and K+
Operation of the sodiumpotassium pump
50
Measuring Membrane Potential
51
Figure 11.7
Resting Membrane Potential
(Vr)
52
Figure 11.8
Nervous Tissue
Fundamentals
P ANervous
RT B
of the
System and
Nervous Tissue
53
Membrane Potentials: Signals
Used to integrate, send, and receive
information
Membrane potential changes are
produced by:
Changes in membrane
permeability to ions
Alterations of ion concentrations
across the membrane
Types of signals – graded
potentials and action potentials
54
Changes in Membrane Potential
Changes are caused by three events
Depolarization – the inside of
the membrane becomes less
negative
Repolarization – the membrane
returns to its resting membrane
potential
Hyperpolarization – the inside
of the membrane becomes more
negative than the resting
55
potential
Depolarization, Repolarization and
Hyperpolarization
56
Figure 12.15
Graded Potentials
Short-lived, local changes in
membrane potential
Decrease in intensity with distance
Magnitude varies directly with the
strength of the stimulus
Depolarization or hyperpolarization
Sufficiently strong graded potentials
can initiate action potentials
57
Graded Potentials
58
Figure 11.10
Graded Potentials
Voltage changes are decremental
Current is quickly dissipated due to
the leaky plasma membrane
Only travel over short distances
59
Action Potentials (APs)
A brief reversal of membrane
potential with a total amplitude of
100 mV
Action potentials are only generated
by muscle cells and neurons
They do not decrease in strength
over distance
They are the principal means of
neural communication
An action potential in the axon of a
neuron is a nerve impulse
60
Action Potential: Resting State
Na+ and K+ voltage gated channels
are closed
Leakage accounts for small
movements of Na+ and K+
Each Na+ channel has two voltageregulated gates
Activation gates
Inactivation gates
61
Figure 11.12.1
Action Potential: Resting State
Na channel is closed, but
capable of opening
62
Action Potential: Depolarization
Phase
Na+ permeability increases;
membrane potential reverses
Na+ gates are opened; K+ gates
are closed
Threshold – a critical level of
depolarization (-55 to -50 mV)
At threshold, depolarization
becomes self-generating
63
Figure 11.12.2
Action Potential: Depolarization
Phase
Both Na channels are opened
64
Action Potential: Repolarization
Phase
Sodium inactivation gates close
Membrane permeability to Na+
declines to resting levels
As sodium gates close, voltagesensitive K+ gates open
K+ exits the cell and internal
negativity of the resting neuron
is restored
65
Figure 11.12.3
Action Potential: Repolarization
Phase
Na activation channel is opened and
inactivation is closed.
66
Action Potential: Hyperpolarization
Potassium gates remain open,
causing an excessive efflux of K+
This efflux causes hyperpolarization
of the membrane (undershoot)
67
Figure 11.12.4
Action Potential:
Hyperpolarization
68
Action Potential:
Role of the Sodium-Potassium Pump
Repolarization
Restores the resting electrical
conditions of the neuron
Does not restore the resting ionic
conditions
Ionic redistribution back to resting
conditions is restored by the sodiumpotassium pump
69
Phases of the Action Potential
• 1 – resting state
• 2 – depolarization
phase
• 3 – repolarization
phase
• 4 –hyperpolarization
70
Figure 11.12
Action Potential
71
Propagation of an Action Potential
along an Unmyelinated Axon
72
Propagation of an action potential
Continuous propagation
Unmyelinated axon
Saltatory propagation
Myelinated axon
73
Propagation of an Action Potential
(Time = 0ms)- continuous propagation
Na gates open
Na+ influx causes a patch of the axonal
membrane to depolarize
Positive Na ions in the axoplasm move
toward the polarized (negative) portion
of the membrane
74
Propagation of an Action Potential
(Time = 2ms)- continuous propagation
A local current is created that depolarizes
the adjacent membrane in a forward
direction
Local voltage-gated Na channels are
opened
The action potential moves away from
the stimulus
75
Propagation of an Action Potential
(Time = 4ms)- continuous propagation
Sodium gates close
Potassium gates open
Repolarization
Hyperpolarization
Sluggish K channels are kept opened
Resting membrane potential is restored
in this region
76
Saltatory Conduction
Current passes through a
myelinated axon only at the nodes
of Ranvier
Voltage-gated Na+ channels are
concentrated at these nodes
Action potentials are triggered only
at the nodes and jump from one
node to the next
Much faster than conduction along
unmyelinated axons
77
Saltatory Conduction
78
Figure 11.16
Types of stimuli
Threshold stimulus
Stimulus strong enough to bring
the membrane potential to a
threshold voltage causing an action
potential
Subthreshold stimulus
weak stimuli that cause
depolarization (graded potentials)
but not action potentials
79
Coding for Stimulus Intensity
Action potential are All-or-none
phenomenon
Strong stimuli can generate an
action potential more often than
weaker stimuli
The CNS determines stimulus
intensity by the frequency of
impulse transmission
80
Stimulus Strength and AP
Frequency
81
Figure 11.14
Absolute Refractory Period
Time from the opening of the Na+
activation gates until the closing of
inactivation gates
The absolute refractory period:
Prevents the neuron from
generating an action potential
Ensures that each action potential
is separate
Enforces one-way transmission of
nerve impulses
82
Absolute and Relative
Refractory Periods
83
Figure 11.15
Relative Refractory Period
The interval following the absolute
refractory period when:
Sodium gates are closed
Potassium gates are open
Repolarization is occurring
Stimulus stronger than the original
one cause a new action potential
84
Conduction Velocities of Axons
Conduction velocities vary widely
among neurons
Rate of impulse propagation is
determined by:
Axon diameter – the larger the
diameter, the faster the impulse
Presence of a myelin sheath –
myelination dramatically
increases impulse speed
85
Nerve Fiber Classification
Nerve fibers are classified according
to:
Diameter
Degree of myelination
Speed of conduction
86
Axon classification
Type A fibers
Diameter of 4 to 20 µm and myelinated
Type B fibers
Diameter of 2 to 4 µm and myelinated
Type C fibers
Diameter less than 2 µm and
unmyelinated
87
Multiple Sclerosis (MS)
An autoimmune disease that mainly
affects young adults
Symptoms: visual disturbances,
weakness, loss of muscular control,
and urinary incontinence
Nerve fibers are severed and myelin
sheaths in the CNS become
nonfunctional scleroses
Shunting and short-circuiting of
nerve impulses occurs
88
Synapses
A junction that mediates
information transfer from one
neuron:
To another neuron
To an effector cell
Presynaptic neuron – conducts
impulses toward the synapse
Postsynaptic neuron – transmits
impulses away from the synapse
89
Synapses
90
Figure 11.17
Types of Synapses
Axodendritic – synapses between
the axon of one neuron and the
dendrite of another
Axosomatic – synapses between the
axon of one neuron and the soma of
another
Others: Axoaxonic, etc
91
Electrical Synapses
Electrical synapses:
Are less common than chemical
synapses
Correspond to gap junctions found in
other cell types
Very fast propagation of action
potentials
92
Electrical Synapses
Are important in the CNS in:
Arousal from sleep
Mental attention
Emotions and memory
Ion and water homeostasis
Light transmitting (eye)
93
Chemical Synapses
Specialized for the release and
reception of neurotransmitters
Typically composed of:
Presynaptic neuron
Contains synaptic vesicles
Postsynaptic neuron
The receptors are located
typically on dendrites and soma
94
Chemical Synapses
Synaptic Cleft
Fluid-filled space separating the
presynaptic and postsynaptic
neurons
Transmission across the synaptic
cleft:
Is a chemical event (as opposed
to an electrical one)
Ensures unidirectional
communication between neurons
95
Synaptic Cleft: Information
Transfer
Nerve impulses reach the axonal
terminal of the presynaptic neuron
and open Ca2+ channels
Neurotransmitter is released into
the synaptic cleft via exocytosis
96
Synaptic Cleft: Information Transfer
Neurotransmitter crosses the
synaptic cleft and binds to receptors
on the postsynaptic neuron
Postsynaptic membrane
permeability changes, causing an
excitatory or inhibitory effect
97
Synaptic Cleft: Information
Transfer
Ca2+
1
Neurotransmitter
Axon terminal of
presynaptic neuron
Postsynaptic
membrane
Mitochondrion
Axon of
presynaptic
neuron
Na+
Receptor
Postsynaptic
membrane
Ion channel open
Synaptic vesicles
containing
neurotransmitter
molecules
5
Degraded
neurotransmitter
2
Synaptic
cleft
Ion channel
(closed)
3
4
Ion channel closed
Ion channel (open)
98
Figure 11.18
Nervous Tissue
Fundamentals
ART C
of thePNervous
System and
Nervous
Tissue
99
Termination of Neurotransmitter
Effects
Neurotransmitter bound to a
postsynaptic neuron:
Produces a continuous postsynaptic
effect
Blocks reception of additional
“messages”
Must be removed from its receptor
100
Termination of Neurotransmitter
Effects
Removal of neurotransmitters
occurs when they:
Are degraded by enzymes
Are reabsorbed by astrocytes or
the presynaptic terminals
Diffuse from the synaptic cleft
101
Synaptic Delay
Neurotransmitter must be released,
diffuse across the synapse, and bind
to receptors
Synaptic delay – time needed to do
this
Synaptic delay is the rate-limiting
step of neural transmission
102
Postsynaptic Potentials
Neurotransmitter receptors mediate
changes in membrane potential
according to:
The amount of neurotransmitter
released
The amount of time the
neurotransmitter is bound to
receptors
103
Postsynaptic Potentials
The two types of postsynaptic
potentials are:
EPSP – excitatory postsynaptic
potentials
IPSP – inhibitory postsynaptic
potentials
104
Excitatory Postsynaptic Potentials
EPSPs are graded potentials that
can initiate an action potential in an
axon
Use only chemically gated
channels
Na+ and K+ flow in opposite
directions at the same time
Postsynaptic membranes do not
generate action potentials
105
Excitatory Postsynaptic
Potential (EPSP)
106
Figure 11.19a
Inhibitory Synapses and IPSPs
Neurotransmitter binding to a
receptor at inhibitory synapses:
Causes the membrane to become
more permeable to potassium and
chloride ions
Leaves the charge on the inner
surface negative
Reduces the postsynaptic
neuron’s ability to produce an
action potential
107
Inhibitory Postsynaptic (IPSP)
108
Figure 11.19b
Summation
A single EPSP cannot induce an
action potential
EPSPs must summate temporally or
spatially to induce an action
potential
Temporal summation –
presynaptic neurons transmit
impulses in rapid-fire order
109
Temporal Summation
110
Summation
Spatial summation – postsynaptic
neuron is stimulated by a large
number of terminals at the same
time
IPSPs can also summate with
EPSPs, canceling each other out
111
Spatial Summation
112
EPSP – IPSP Interactions
113
Neurotransmitters
Chemicals used for neuronal
communication with the body and
the brain
50 different neurotransmitters have
been identified
Classified chemically and
functionally
114
Chemical Neurotransmitters
Acetylcholine (ACh)
Biogenic amines
Amino acids
Peptides
Novel messengers: ATP and
dissolved gases NO and CO
115
Neurotransmitters: Acetylcholine
First neurotransmitter identified,
and best understood
Released at the neuromuscular
junction
Synthesized and enclosed in
synaptic vesicles
116
Neurotransmitters: Acetylcholine
Degraded by the enzyme
acetylcholinesterase (AChE)
Released by:
All neurons that stimulate skeletal
muscle
Some neurons in the autonomic
nervous system
117
Neurotransmitters: Biogenic
Amines
Include:
Catecholamines – dopamine,
norepinephrine (NE), and
epinephrine
Indolamines – serotonin and
histamine
Broadly distributed in the brain
Play roles in emotional behaviors
and our biological clock
118
Synthesis of Catecholamines
Enzymes present in the cell
determine length of biosynthetic
pathway
Norepinephrine and dopamine are
synthesized in axonal terminals
Epinephrine is released by the
adrenal medulla
119
Figure
11.21
Synthesis of Catecholamines
120
Neurotransmitters: Amino Acids
Include:
GABA – Gamma ()-aminobutyric
acid
Glycine
Aspartate
Glutamate
Found only in the CNS
121
Neurotransmitters: Peptides
Include:
Substance P – mediator of pain
signals
Beta endorphin, dynorphin, and
enkephalins
Act as natural opiates; reduce pain
perception
Bind to the same receptors as
opiates and morphine
Gut-brain peptides – somatostatin,
122
and cholecystokinin
Neurotransmitters: Novel
Messengers
ATP
Is found in both the CNS and PNS
Produces excitatory or inhibitory
responses depending on receptor
type
Induces Ca2+ wave propagation in
astrocytes
Provokes pain sensation
123
Neurotransmitters: Novel
Messengers
Nitric oxide (NO)
Activates the intracellular
receptor guanylyl cyclase
Is involved in learning and
memory
Carbon monoxide (CO) is a main
regulator of cGMP in the brain
124
Functional Classification of
Neurotransmitters
Two classifications: excitatory and
inhibitory
Excitatory neurotransmitters
cause depolarizations
(e.g., glutamate)
Inhibitory neurotransmitters
cause hyperpolarizations (e.g.,
GABA and glycine)
125
Functional Classification of
Neurotransmitters
Some neurotransmitters have both
excitatory and inhibitory effects
Determined by the receptor type
of the postsynaptic neuron
Example: acetylcholine
Excitatory at neuromuscular
junctions with skeletal muscle
Inhibitory in cardiac muscle
126
Neurotransmitter Receptor
Mechanisms
Direct: neurotransmitters that open
ion channels
Promote rapid responses
Examples: ACh and amino acids
Indirect: neurotransmitters that
act through second messengers
Promote long-lasting effects
Examples: biogenic amines,
peptides, and dissolved gases
127
Channel-Linked Receptors
Composed of integral membrane
protein
Mediate direct neurotransmitter
action
Action is immediate, brief, simple,
and highly localized
128
Channel-Linked Receptors
Ligand binds the receptor, and ions
enter the cells
Excitatory receptors depolarize
membranes
Inhibitory receptors hyperpolarize
membranes
129
Channel-Linked Receptors
130
Figure 11.22a
G Protein-Linked Receptors
Responses are indirect, slow,
complex, prolonged, and often
diffuse
These receptors are transmembrane
protein complexes
First and Second messengers.
131
Neurotransmitter Receptor
Mechanism
Ions flow
Blocked ion flow
(a)
Channel closed
Adenylate
cyclase
Channel open
Neurotransmitter (ligand)
released from axon terminal
of presynaptic neuron
3
1
PPi
4
GTP
5
cAMP
ATP
5
3
Changes in
membrane
permeability
and potential
GTP
2
GDP
Protein
synthesis
Enzyme
activation
GTP
Receptor
G protein
(b)
Nucleus
Activation of
specific genes
132
Figure 11.22b
Neural Integration: Neuronal Pools
Functional groups of neurons that:
Integrate incoming information
Forward the processed
information to its appropriate
destination
133
Neural Integration: Neuronal Pools
Simple neuronal pool
Input fiber – presynaptic fiber
Discharge zone – neurons most
closely associated with the
incoming fiber
Facilitated zone – neurons farther
away from incoming fiber
134
Simple Neuronal Pool
135
Figure 11.23
Types of Neuronal Pools
136
Types of Circuits in Neuronal
Pools
Divergent – information spreads
from one neuron to several neurons
or from one pool to several pools.
One motor neuron of the brain
stimulates many muscle fibers
137 b
Figure 11.24a,
Types of Circuits in Neuronal
Pools
Convergent – input from many
neurons is funneled to one neuron or
neuronal pools
Motor neurons of the diaphragm is
controlled by different areas of the
brain for subconscious and
conscious functioning
138 d
Figure 11.24c,
Types of Circuits in Neuronal
Pools
Reverberating –collateral
branches of axons extend back
toward the source of the impulse
Involved in rhythmic activities
Breathing, sleep-wake cycle, etc
Figure139
11.24e
Types of Circuits in Neuronal
Pools
Parallel - incoming neurons
stimulate several neurons in parallel
arrays
Divergence must take place before
the parallel processing
Response to a painfull stimuli
causes
Withdrawal of the limb
Shift of the weight
Fell pain
Shout “ouch!”
Figure140
11.24f
Patterns of Neural Processing
Serial
Transmitting of the impulse from
one neuron to another or from
one pool to another
Spinal reflexes
141
Development of Neurons
The nervous system originates from
the neural tube and neural crest
The neural tube becomes the CNS
There is a three-phase process of
differentiation:
Proliferation of cells needed for
development
Migration – neuroblasts become
amitotic and move externally
142
Differentiation into neurons
Axonal Growth
Guided by:
Scaffold laid down by older neurons
Orienting glial fibers
Release of nerve growth factor by
astrocytes
Neurotropins released by other
neurons
Repulsion guiding molecules
Attractants released by target cells
143