16 Neural Integration II: The Autonomic Nervous System and Higher-Order Functions
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Transcript of 16 Neural Integration II: The Autonomic Nervous System and Higher-Order Functions
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PowerPoint® Lecture Presentations prepared byJason LaPresLone Star College—North Harris
16Neural Integration II: The Autonomic Nervous System and Higher-Order Functions
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An Introduction to the ANS and Higher-Order Functions
• Learning Outcomes
• 16-1 Compare the organization of the autonomic nervous system with that of the somatic
nervous system.
• 16-2 Describe the structures and functions of the sympathetic division of the autonomic
nervous system.
• 16-3 Describe the mechanisms of sympathetic neurotransmitter release and their
effects on target organs and tissues.
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An Introduction to the ANS and Higher-Order Functions • Learning Outcomes
• 16-4 Describe the structures and functions of the parasympathetic division of the autonomic
nervous system.
• 16-5 Describe the mechanisms of parasympathetic neurotransmitter release and their effects on
target organs and tissues.
• 16-6 Discuss the functional significance of dual innervation and autonomic tone.
• 16-7 Describe the hierarchy of interacting levels of control in the autonomic nervous system,
including the significance of visceral reflexes.
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An Introduction to the ANS and Higher-Order Functions
• Learning Outcomes
• 16-8 Explain how memories are created, stored, and recalled, and distinguish among the levels
of consciousness and unconsciousness.
• 16-9 Describe some of the ways in which the interactions of neurotransmitters influence
brain function.
• 16-10 Summarize the effects of aging on the nervous system and give examples of
interactions between the nervous system and other organ systems.
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An Introduction to the ANS and Higher-Order Functions
• Somatic Nervous System (SNS)
• Operates under conscious control
• Seldom affects long-term survival
• SNS controls skeletal muscles
• Autonomic Nervous System (ANS)
• Operates without conscious instruction
• ANS controls visceral effectors
• Coordinates system functions
• Cardiovascular, respiratory, digestive, urinary, reproductive
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Figure 16-1 An Overview of Neural Integration
CHAPTER 16CHAPTER 15
Higher-Order FunctionsConscious andsubconsciousmotor centers
in brain
Motorpathways
Memory, learning, andintelligence may
influence interpretationof sensory information
and nature of motoractivities
AutonomicNervous
System (ANS)
Visceral effectors(examples: smooth
muscle, glands,cardiac muscle,
adipocytes)
Sensoryprocessingcenters in
brain
Sensorypathways
Generalsensory
receptors
SomaticNervous
System (SNS)
Skeletalmuscles
OVERVIEW OF NEURAL INTEGRATION
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16-1 Autonomic Nervous System
• Organization of the ANS
• Integrative centers
• For autonomic activity in hypothalamus
• Neurons comparable to upper motor neurons in SNS
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16-1 Autonomic Nervous System
• Organization of the ANS
• Visceral motor neurons
• In brain stem and spinal cord, are known as
preganglionic neurons
• Preganglionic fibers
• Axons of preganglionic neurons
• Leave CNS and synapse on ganglionic neurons
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16-1 Autonomic Nervous System
• Visceral Motor Neurons
• Autonomic ganglia
• Contain many ganglionic neurons
• Ganglionic neurons innervate visceral effectors
• Such as cardiac muscle, smooth muscle, glands,
and adipose tissue
• Postganglionic fibers
• Axons of ganglionic neurons
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Figure 16-2a The Organization of the Somatic and Autonomic Nervous Systems
Somatic nervous system
Somaticmotor nucleiof spinal cord
SPINALCORD
Lowermotor
neurons
Skeletalmuscle
Skeletalmuscle
Somatic motornuclei of brain
stem
BRAIN
Upper motorneurons in
primary motorcortex
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Figure 16-2b The Organization of the Somatic and Autonomic Nervous Systems
Autonomic nervous system
Smoothmuscle
Cardiacmuscle
Adipocytes
Glands
Visceral Effectors
Preganglionicneuron
BRAIN
Autonomicganglia
Ganglionicneurons
Preganglionicneurons
Autonomicnuclei inbrain stem
SPINALCORD
Autonomicnuclei inspinal cord
Visceral motornuclei in
hypothalamus
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16-1 Divisions of the ANS
• The Autonomic Nervous System
• Operates largely outside our awareness
• Has two divisions
1. Sympathetic division
• Increases alertness, metabolic rate, and muscular
abilities
2. Parasympathetic division
• Reduces metabolic rate and promotes digestion
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16-1 Divisions of the ANS
• Sympathetic Division
• “Kicks in” only during exertion, stress, or
emergency
• “Fight or flight”
• Parasympathetic Division
• Controls during resting conditions
• “Rest and digest”
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16-1 Divisions of the ANS
• Sympathetic and Parasympathetic Division
1. Most often, these two divisions have opposing effects
• If the sympathetic division causes excitation, the
parasympathetic causes inhibition
2. The two divisions may also work independently
• Only one division innervates some structures
3. The two divisions may work together, with each
controlling one stage of a complex process
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16-1 Divisions of the ANS
• Sympathetic Division
• Preganglionic fibers (thoracic and superior lumbar;
thoracolumbar) synapse in ganglia near spinal cord
• Preganglionic fibers are short
• Postganglionic fibers are long
• Prepares body for crisis, producing a “fight or flight”
response
• Stimulates tissue metabolism
• Increases alertness
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16-1 Divisions of the ANS
• Seven Responses to Increased Sympathetic Activity
1. Heightened mental alertness
2. Increased metabolic rate
3. Reduced digestive and urinary functions
4. Energy reserves activated
5. Increased respiratory rate and respiratory passageways dilate
6. Increased heart rate and blood pressure
7. Sweat glands activated
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16-1 Divisions of the ANS
• Parasympathetic Division
• Preganglionic fibers originate in brain stem and sacral
segments of spinal cord; craniosacral
• Synapse in ganglia close to (or within) target organs
• Preganglionic fibers are long
• Postganglionic fibers are short
• Parasympathetic division stimulates visceral activity
• Conserves energy and promotes sedentary activities
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16-1 Divisions of the ANS
• Five Responses to Increased Parasympathetic
Activity
1. Decreased metabolic rate
2. Decreased heart rate and blood pressure
3. Increased secretion by salivary and digestive glands
4. Increased motility and blood flow in digestive tract
5. Urination and defecation stimulation
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16-1 Divisions of the ANS
• Enteric Nervous System (ENS)
• Third division of ANS
• Extensive network in digestive tract walls
• Complex visceral reflexes coordinated locally
• Roughly 100 million neurons
• All neurotransmitters are found in the brain
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16-2 The Sympathetic Division
• The Sympathetic Division
• Preganglionic neurons located between segments T1
and L2 of spinal cord
• Ganglionic neurons in ganglia near vertebral column
• Cell bodies of preganglionic neurons in lateral gray
horns
• Axons enter ventral roots of segments
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Figure 16-3 The Organization of the Sympathetic Division of the ANS
Sympathetic Division of ANS
Ganglionic NeuronsPreganglionic
Neurons
Target Organs
KEYPreganglionic fibers
Postganglionic fibers
Hormones releasedinto circulation
Lateral grayhorns of spinal
segmentsT1–L2
Sympatheticchain ganglia
(paired)
Collateralganglia
(unpaired)
Adrenalmedullae(paired)
Visceral effectorsin thoracic cavity,head, body wall,
and limbs
Visceral effectorsin abdominopelvic
cavity
Organs and systemsthroughout body
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16-2 The Sympathetic Division
• Ganglionic Neurons
• Occur in three locations
1. Sympathetic chain ganglia
2. Collateral ganglia
3. Suprarenal medullae
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16-2 The Sympathetic Division
• Sympathetic Chain Ganglia
• Are on both sides of vertebral column
• Control effectors:
• In body wall
• Inside thoracic cavity
• In head
• In limbs
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Figure 16-4a Sites of Ganglia in Sympathetic Pathways
SYMPATHETIC CHAIN GANGLIA
Preganglionic neuronsGanglionic neurons
KEY
Gray ramus
White ramusGanglionicneuron
Preganglionicneuron
Autonomic ganglion ofright sympathetic chain
Innervatesvisceral
effectors viaspinal nerves
Innervates visceralorgans in thoracic
cavity viasympathetic nerves
Autonomic ganglionof left sympathetic chain
Spinal nerve
Sympathetic nerve(postganglionic
fibers)
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16-2 The Sympathetic Division
• Collateral Ganglia
• Are anterior to vertebral bodies
• Contain ganglionic neurons that innervate tissues and
organs in abdominopelvic cavity
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Figure 16-4b Sites of Ganglia in Sympathetic Pathways
Preganglionic neuronsGanglionic neurons
KEY
COLLATERAL GANGLIA
Splanchnicnerve
(preganglionicfibers)
Postganglionicfibers
Collateralganglion
Innervatesvisceral organs inabdominopelvic
cavity
Whiteramus
Lateralgrayhorn
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16-2 The Sympathetic Division
• Adrenal Medullae (Suprarenal Medullae)
• Very short axons
• When stimulated, release neurotransmitters into
bloodstream (not at synapse)
• Function as hormones to affect target cells throughout
body
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Figure 16-4c Sites of Ganglia in Sympathetic Pathways
Preganglionic neuronsGanglionic neurons
KEY
THE ADRENAL MEDULLAE
Preganglionic fibers Secretesneurotransmitters
into generalcirculation
Endocrine cells(specialized ganglionic
neurons)
Adrenalmedullae
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16-2 The Sympathetic Division
• Fibers in Sympathetic Division
• Preganglionic fibers
• Are relatively short
• Ganglia located near spinal cord
• Postganglionic fibers
• Are relatively long, except at adrenal medullae
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• Organization and Anatomy of the Sympathetic Division
• Ventral roots of spinal segments T1–L2 contain
sympathetic preganglionic fibers
• Give rise to myelinated white ramus
• Carry myelinated preganglionic fibers into sympathetic
chain ganglion
• May synapse at collateral ganglia or in adrenal
medullae
16-2 The Sympathetic Division
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16-2 The Sympathetic Division
• Sympathetic Chain Ganglia
• Preganglionic fibers
• One preganglionic fiber synapses on many
ganglionic neurons
• Fibers interconnect sympathetic chain ganglia
• Each ganglion innervates particular body
segment(s)
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16-2 The Sympathetic Division
• Sympathetic Chain Ganglia
• Postganglionic Fibers
• Paths of unmyelinated postganglionic fibers
depend on targets
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16-2 The Sympathetic Division
• Sympathetic Chain Ganglia
• Postganglionic fibers control visceral effectors
• In body wall, head, neck, or limbs
• Enter gray ramus
• Return to spinal nerve for distribution
• Postganglionic fibers innervate effectors
• Sweat glands of skin
• Smooth muscles in superficial blood vessels
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16-2 The Sympathetic Division
• Sympathetic Chain Ganglia
• Postganglionic fibers innervating structures in thoracic
cavity form bundles
• Sympathetic nerves
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16-2 The Sympathetic Division
• Sympathetic Chain Ganglia
• Each sympathetic chain ganglia contains:
• 3 cervical ganglia
• 10–12 thoracic ganglia
• 4–5 lumbar ganglia
• 4–5 sacral ganglia
• 1 coccygeal ganglion
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16-2 The Sympathetic Division
• Sympathetic Chain Ganglia
• Preganglionic neurons
• Limited to spinal cord segments T1–L2
• White rami (myelinated preganglionic fibers)
• Innervate neurons in:
• Cervical, inferior lumbar, and sacral sympathetic chain
ganglia
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16-2 The Sympathetic Division
• Sympathetic Chain Ganglia
• Chain ganglia provide postganglionic fibers
• Through gray rami (unmyelinated postganglionic
fibers)
• To cervical, lumbar, and sacral spinal nerves
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16-2 The Sympathetic Division
• Sympathetic Chain Ganglia
• Only spinal nerves T1–L2 have white rami
• Every spinal nerve has gray ramus
• That carries sympathetic postganglionic fibers for
distribution in body wall
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16-2 The Sympathetic Division
• Sympathetic Chain Ganglia
• Postganglionic sympathetic fibers
• In head and neck leave superior cervical
sympathetic ganglia
• Supply the regions and structures innervated by
cranial nerves III, VII, IX, X
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Figure 16-5 The Distribution of Sympathetic Innervation
Preganglionic neuronsGanglionic neurons
KEY
T1 T1
Gray rami tospinal nerves
Cervicalsympathetic
ganglia
Superior
Middle
Inferior
PONS
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Figure 16-5 The Distribution of Sympathetic Innervation
Preganglionic neuronsGanglionic neurons
KEY
T1
PONS
Sympathetic nerves
Cardiac andpulmonary plexuses(see Figure 16-10)
Eye
Salivaryglands
Heart
Lung
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Figure 16-5 The Distribution of Sympathetic Innervation
Preganglionic neuronsGanglionic neurons
KEY
Coccygealganglia (Co1)
fused together
L2
Superiormesenteric
ganglion
Splanchnicnerves
Uterus Ovary
Greatersplanchnicnerve
T1
Celiac ganglion
Inferiormesentericganglion
Penis Scrotum Urinary bladder
Liver and gallbladder
Stomach
SpleenPancreas
Largeintestine
Smallintestine
AdrenalmedullaKidney
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Figure 16-5 The Distribution of Sympathetic Innervation
Preganglionic neuronsGanglionic neurons
KEY
Spinal cord
Sympatheticchain ganglia
L2
Postganglionic fibersto spinal nerves
(innervating skin,blood vessels,sweat glands,
arrector pili muscles,adipose tissue)
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16-2 The Sympathetic Division
• Collateral Ganglia
• Receive sympathetic innervation via sympathetic preganglionic fibers
• Splanchnic nerves
• Formed by preganglionic fibers that innervate collateral ganglia
• In dorsal wall of abdominal cavity
• Originate as paired ganglia (left and right)
• Usually fuse together in adults
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16-2 The Sympathetic Division
• Collateral Ganglia
• Postganglionic fibers
• Leave collateral ganglia
• Extend throughout abdominopelvic cavity
• Innervate variety of visceral tissues and organs
• Reduction of blood flow and energy by organs not
vital to short-term survival
• Release of stored energy reserves
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16-2 The Sympathetic Division
• Collateral Ganglia
• Preganglionic fibers from seven inferior thoracic
segments
• End at celiac ganglion or superior mesenteric
ganglion
• Ganglia embedded in network of autonomic nerves
• Preganglionic fibers from lumbar segments
• Form splanchnic nerves
• End at inferior mesenteric ganglion
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16-2 The Sympathetic Division
• Collateral Ganglia
• Celiac ganglion
• Pair of interconnected masses of gray matter
• May form single mass or many interwoven masses
• Postganglionic fibers innervate stomach, liver,
gallbladder, pancreas, and spleen
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16-2 The Sympathetic Division
• Collateral Ganglia
• Superior mesenteric ganglion
• Near base of superior mesenteric artery
• Postganglionic fibers innervate small intestine and
proximal 2/3 of large intestine
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16-2 The Sympathetic Division
• Collateral Ganglia
• Inferior mesenteric ganglion
• Near base of inferior mesenteric artery
• Postganglionic fibers provide sympathetic innervation to
portions of:
• Large intestine
• Kidney
• Urinary bladder
• Sex organs
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16-2 The Sympathetic Division
• Adrenal Medullae
• Preganglionic fibers entering adrenal gland proceed
to center (adrenal medulla)
• Modified sympathetic ganglion
• Preganglionic fibers synapse on neuroendocrine cells
• Specialized neurons secrete hormones into
bloodstream
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16-2 The Sympathetic Division
• Adrenal Medullae
• Neuroendocrine cells
• Secrete neurotransmitters epinephrine (E) and
norepinephrine (NE)
• Epinephrine
• Also called adrenaline
• Is 75–80% of secretory output
• Remaining is norepinephrine (NE)
• Noradrenaline
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16-2 The Sympathetic Division
• Adrenal Medullae
• Bloodstream carries neurotransmitters through body
• Causing changes in metabolic activities of different
cells including cells not innervated by sympathetic
postganglionic fibers
• Effects last longer
• Hormones continue to diffuse out of bloodstream
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16-2 The Sympathetic Division
• Sympathetic Activation
• Change activities of tissues and organs by:
• Releasing NE at peripheral synapses
• Target specific effectors, smooth muscle fibers in
blood vessels of skin
• Are activated in reflexes
• Do not involve other visceral effectors
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16-2 The Sympathetic Division
• Sympathetic Activation
• Changes activities of tissues and organs by:
• Distributing E and NE throughout body in bloodstream
• Entire division responds (sympathetic activation)
• Are controlled by sympathetic centers in
hypothalamus
• Effects are not limited to peripheral tissues
• Alters CNS activity
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16-2 The Sympathetic Division
• Changes Caused by Sympathetic Activation
• Increased alertness
• Feelings of energy and euphoria
• Change in breathing
• Elevation in muscle tone
• Mobilization of energy reserves
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16-3 Various Sympathetic Neurotransmitters
• Stimulation of Sympathetic Preganglionic
Neurons
• Releases ACh at synapses with ganglionic neurons
• Excitatory effect on ganglionic neurons
• Ganglionic Neurons
• Release neurotransmitters at specific target organs
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16-3 Various Sympathetic Neurotransmitters
• Ganglionic Neurons
• Axon terminals
• Form branching networks of telodendria instead of synaptic
terminals
• Telodendria form sympathetic varicosities
• Resemble string of pearls
• Swollen segment packed with neurotransmitter vesicles
• Pass along or near surface of effector cells
• No specialized postsynaptic membranes
• Membrane receptors on surfaces of target cells
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Figure 16-6 Sympathetic Varicosities
Preganglionic fiber(myelinated)
Ganglionicneuron
Ganglion
Vesicles containingnorepinephrine
Postganglionic fiber(unmyelinated)
Varicosities
Mitochondrion
5 m
Smooth muscle cells Varicosities
Schwann cellcytoplasm
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16-3 Various Sympathetic Neurotransmitters
• Ganglionic Neurons
• Axon terminals
• Release NE at most varicosities
• Called adrenergic neuron
• Some ganglionic neurons release ACh instead
• Are located in body wall, skin, brain, and skeletal
muscles
• Called cholinergic neurons
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16-3 Various Sympathetic Neurotransmitters
• Sympathetic Stimulation and the Release of NE
and E
• Primarily from interactions of NE and E with two
types of adrenergic membrane receptors
1. Alpha receptors (NE more potent)
2. Beta receptors
• Activates enzymes on inside of cell membrane via G
proteins
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16-3 Various Sympathetic Neurotransmitters
• Sympathetic Stimulation and the Release of NE
and E
• Alpha-1 (1)
• More common type of alpha receptor
• Releases intracellular calcium ions from reserves
in endoplasmic reticulum
• Has excitatory effect on target cell
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16-3 Various Sympathetic Neurotransmitters
• Sympathetic Stimulation and the Release of NE and E
• Alpha-2 (2)
• Lowers cAMP levels in cytoplasm
• Has inhibitory effect on the cell
• Helps coordinate sympathetic and parasympathetic activities
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16-3 Various Sympathetic Neurotransmitters
• Sympathetic Stimulation and the Release of NE
and E
• Beta () receptors
• Affect membranes in many organs (skeletal muscles,
lungs, heart, and liver)
• Trigger metabolic changes in target cell
• Stimulation increases intracellular cAMP levels
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16-3 Various Sympathetic Neurotransmitters
• Three Main Types of Beta Receptors
1. Beta-1 (1)
• Increases metabolic activity
2. Beta-2 (2)
• Triggers relaxation of smooth muscles along respiratory tract
3. Beta-3 (3)
• Leads to lipolysis, the breakdown of triglycerides in
adipocytes
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16-3 Various Sympathetic Neurotransmitters
• Sympathetic Stimulation and the Release of ACh
and NO
• Cholinergic (ACh) sympathetic terminals
• Innervate sweat glands of skin and blood vessels of
skeletal muscles and brain
• Stimulate sweat gland secretion and dilate blood
vessels
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16-3 Various Sympathetic Neurotransmitters
• Sympathetic Stimulation and the Release of ACh
and NO
• Nitroxidergic synapses
• Release nitric oxide (NO) as neurotransmitter
• Neurons innervate smooth muscles in walls of blood
vessels in skeletal muscles and the brain
• Produce vasodilation and increased blood flow
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16-4 The Parasympathetic Division
• Autonomic Nuclei
• Are contained in the mesencephalon, pons, and
medulla oblongata
• Associated with cranial nerves III, VII, IX, X
• In lateral gray horns of spinal segments S2–S4
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16-4 The Parasympathetic Division
• Ganglionic Neurons in Peripheral Ganglia
• Terminal ganglion
• Near target organ
• Usually paired
• Intramural ganglion
• Embedded in tissues of target organ
• Interconnected masses
• Clusters of ganglion cells
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16-4 The Parasympathetic Division
• Organization and Anatomy of the Parasympathetic
Division
• Parasympathetic preganglionic fibers leave brain as
components of cranial nerves
• III (oculomotor)
• VII (facial)
• IX (glossopharyngeal)
• X (vagus)
• Parasympathetic preganglionic fibers leave spinal cord at
sacral level
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Figure 16-7 The Organization of the Parasympathetic Division of the ANS
Preganglionic fibersPostganglionic fibers
KEYNuclei in
spinal cordsegments
S2–S4
Pelvicnerves
Nuclei inbrain stem
Preganglionic Neurons Ganglionic NeuronsN III
N VII
N IX
N X
Intramuralganglia
Intramuralganglia
Otic ganglion
Pterygopalatineand submandibular
ganglia
Ciliary ganglion
Target Organs
Intrinsic eye muscles(pupil and lens shape)
Nasal glands, tearglands, and salivary
glands
Parotid salivary gland
Visceral organsof neck,
thoracic cavity,and most of
abdominal cavity
Visceral organs ininferior portion of
abdominopelvic cavity
Parasympathetic Division of ANS
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16-4 The Parasympathetic Division
• Oculomotor, Facial, and Glossopharyngeal
Nerves
• Control visceral structures in head
• Synapse in ciliary, pterygopalatine, submandibular,
and otic ganglia
• Short postganglionic fibers continue to their peripheral
targets
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16-4 The Parasympathetic Division
• Vagus Nerve
• Provides preganglionic parasympathetic innervation to
structures in:
• Neck
• Thoracic and abdominopelvic cavity as distant as a distal
portion of large intestine
• Provides 75% of all parasympathetic outflow
• Branches intermingle with fibers of sympathetic division
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16-4 The Parasympathetic Division
• Sacral Segments of Spinal Cord
• Preganglionic fibers carry sacral parasympathetic
output
• Do not join ventral roots of spinal nerves, instead form
pelvic nerves
• Pelvic nerves innervate intramural ganglia in walls of
kidneys, urinary bladder, portions of large intestine, and
the sex organs
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Figure 16-8 The Distribution of Parasympathetic Innervation
Preganglionic neuronsGanglionic neurons
KEY
N X (Vagus)
N IX
N VII
N III
PONS
Otic ganglion
Ciliary ganglion
Pterygopalatine ganglion
Submandibular ganglion
Lacrimal gland
Eye
Salivary glands
Heart
Lungs
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Figure 16-8 The Distribution of Parasympathetic Innervation
Preganglionic neuronsGanglionic neurons
KEY
Liver andgallbladder
Stomach
Spleen
Pancreas
LargeintestineSmallintestineRectum
Kidney
Uterus Ovary Penis Scrotum
Lungs
Urinary bladder
S2
S3
S4
Spinalcord
Pelvicnerves
Autonomic plexuses(see Figure 16-10)
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16-4 The Parasympathetic Division
• Parasympathetic Activation
• Centers on relaxation, food processing, and
energy absorption
• Localized effects, last a few seconds at most
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16-4 The Parasympathetic Division
• Major Effects of Parasympathetic Division
• Constriction of the pupils
• (To restrict the amount of light that enters the eyes)
• And focusing of the lenses of the eyes on nearby objects
• Secretion by digestive glands
• Including salivary glands, gastric glands, duodenal
glands, intestinal glands, the pancreas (exocrine and
endocrine), and the liver
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16-4 The Parasympathetic Division
• Major Effects of Parasympathetic Division
• Secretion of hormones
• That promote the absorption and utilization of nutrients
by peripheral cells
• Changes in blood flow and glandular activity
• Associated with sexual arousal
• Increase in smooth muscle activity
• Along the digestive tract
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16-4 The Parasympathetic Division
• Major Effects of Parasympathetic Division
• Stimulation and coordination of defecation
• Contraction of the urinary bladder during urination
• Constriction of the respiratory passageways
• Reduction in heart rate and in the force of contraction
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16-5 Parasympathetic Neurons Release ACh
• Neuromuscular and Neuroglandular Junctions
• All release ACh as neurotransmitter
• Small, with narrow synaptic clefts
• Effects of stimulation are short lived
• Inactivated by acetylcholinesterase (AChE) at synapse
• ACh is also inactivated by tissue cholinesterase in
surrounding tissues
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16-5 Parasympathetic Neurons Release ACh
• Membrane Receptors and Responses
• Nicotinic receptors
• On surfaces of ganglion cells (sympathetic and
parasympathetic)
• Exposure to ACh causes excitation of ganglionic
neuron or muscle fiber
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16-5 Parasympathetic Neurons Release ACh
• Membrane Receptors and Responses
• Muscarinic receptors
• At cholinergic neuromuscular or neuroglandular junctions (parasympathetic)
• At few cholinergic junctions (sympathetic)
• G proteins
• Effects are longer lasting than nicotinic receptors
• Response reflects activation or inactivation of specific enzymes
• Can be excitatory or inhibitory
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16-5 Parasympathetic Neurons Release ACh
• Dangerous Environmental Toxins
• Produce exaggerated, uncontrolled responses
• Nicotine
• Binds to nicotinic receptors
• Targets autonomic ganglia and skeletal neuromuscular junctions
• 50 mg ingested or absorbed through skin
• Signs and symptoms:
• Vomiting, diarrhea, high blood pressure, rapid heart rate, sweating, profuse salivation, convulsions
• May result in coma or death
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16-5 Parasympathetic Neurons Release ACh
• Dangerous Environmental Toxins
• Produce exaggerated, uncontrolled responses
• Muscarine
• Binds to muscarinic receptors
• Targets parasympathetic neuromuscular or neuroglandular
junctions
• Signs and symptoms:
• Salivation, nausea, vomiting, diarrhea, constriction of
respiratory passages, low blood pressure, slow heart
rate (bradycardia)
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Table 16-1 Adrenergic and Cholinergic Receptors of the ANS
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16-6 Dual Innervation
• Sympathetic Division
• Widespread impact
• Reaches organs and tissues throughout body
• Parasympathetic Division
• Innervates only specific visceral structures
• Sympathetic and Parasympathetic Division
• Most vital organs receive instructions from both
sympathetic and parasympathetic divisions
• Two divisions commonly have opposing effects
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16-6 Dual Innervation
• Anatomy of Dual Innervation
• Parasympathetic postganglionic fibers accompany
cranial nerves to peripheral destinations
• Sympathetic innervation reaches same structures
• By traveling directly from superior cervical ganglia of
sympathetic chain
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Figure 16-9 Summary: The Anatomical Differences between the Sympathetic and Parasympathetic Divisions
KEYNeurotransmitters
Epinephrine
Norepinephrine
Acetylcholine
Parasympatheticganglion
TARGET
Circulatorysystem
Sympatheticganglion
or
Postganglionicfiber
Ganglionicneurons
Preganglionicneuron
Preganglionicfiber
CNS
PNS
Sympathetic Parasympathetic
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Table 16-2 A Structural Comparison of the Sympathetic and Parasympathetic Divisions of the ANS
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Table 16-3 A Functional Comparison of the Sympathetic and Parasympathetic Divisions of the ANS
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Table 16-3 A Functional Comparison of the Sympathetic and Parasympathetic Divisions of the ANS
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Table 16-3 A Functional Comparison of the Sympathetic and Parasympathetic Divisions of the ANS
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Table 16-3 A Functional Comparison of the Sympathetic and Parasympathetic Divisions of the ANS
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16-6 Dual Innervation
• Anatomy of Dual Innervation
• Autonomic plexuses
• Nerve networks in the thoracic and abdominopelvic
cavities
• Are formed by mingled sympathetic postganglionic
fibers and parasympathetic preganglionic fibers
• Travel with blood and lymphatic vessels that supply
visceral organs
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16-6 Dual Innervation
• Anatomy of Dual Innervation
• Cardiac plexus
• Pulmonary plexus
• Esophageal plexus
• Celiac plexus
• Inferior mesenteric plexus
• Hypogastric plexus
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16-6 Dual Innervation
• Cardiac and Pulmonary Plexuses
• Autonomic fibers entering thoracic cavity intersect
• Contain:
• Sympathetic and parasympathetic fibers for heart and
lungs
• Parasympathetic ganglia whose output affects those
organs
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16-6 Dual Innervation
• Esophageal Plexus
• Contains:
• Descending branches of vagus nerve
• Splanchnic nerves leaving sympathetic chain
• Parasympathetic preganglionic fibers of vagus nerve
enter abdominopelvic cavity with esophagus
• Fibers enter celiac plexus (solar plexus)
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Figure 16-10 The Autonomic Plexuses and Ganglia
Autonomic Plexusesand Ganglia
Right vagus nerve
Aortic arch
Cardiac plexus
Pulmonary plexus
Thoracic sympatheticchain ganglia
Esophageal plexus
Celiac plexusand ganglion
Superior mesentericganglion
Inferior mesentericplexus and ganglia
Pelvic sympatheticchain
Hypogastric plexus
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16-6 Dual Innervation
• Celiac Plexus
• Associated with smaller plexuses, such as inferior
mesenteric plexus
• Innervates viscera within abdominal cavity
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16-6 Dual Innervation
• Hypogastric Plexus
• Contains:
• Parasympathetic outflow of pelvic nerves
• Sympathetic postganglionic fibers from inferior
mesenteric ganglion
• Splanchnic nerves from sacral sympathetic chain
• Innervates digestive, urinary, and reproductive organs
of pelvic cavity
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Figure 16-10 The Autonomic Plexuses and Ganglia
Inferior mesenteric artery
Superior mesenteric artery
Diaphragm
Esophagus
Splanchnic nerves
Thoracic spinal nerves
Left vagus nerve
Trachea
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16-6 Dual Innervation
• Autonomic Tone
• Is an important aspect of ANS function
• If nerve is inactive under normal conditions, can only
increase activity
• If nerve maintains background level of activity, can
increase or decrease activity
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16-6 Dual Innervation
• Autonomic Tone
• Autonomic motor neurons
• Maintain resting level of spontaneous activity
• Background level of activation determines autonomic
tone
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16-6 Dual Innervation
• Autonomic Tone
• Significant where dual innervation occurs
• Two divisions have opposing effects
• More important when dual innervation does not
occur
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16-6 Dual Innervation
• The Heart Receives Dual Innervation• Two divisions have opposing effects on heart function
1. Parasympathetic division
• Acetylcholine released by postganglionic fibers slows heart rate
2. Sympathetic division
• NE released by varicosities accelerates heart rate
• Balance between two divisions
• Autonomic tone is present
• Releases small amounts of both neurotransmitters continuously
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16-6 Dual Innervation
• The Heart Receives Dual Innervation
• Parasympathetic innervation dominates under resting
conditions
• Crisis accelerates heart rate by:
• Stimulation of sympathetic innervation
• Inhibition of parasympathetic innervation
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16-6 Dual Innervation
• Autonomic Tone
• Blood vessel dilates and blood flow increases
• Blood vessel constricts and blood flow is reduced
• Sympathetic postganglionic fibers release NE
• Innervate smooth muscle cells in walls of
peripheral vessels
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16-6 Dual Innervation
• Autonomic Tone
• Background sympathetic tone keeps muscles partially
contracted
• To increase blood flow:
• Rate of NE release decreases
• Sympathetic cholinergic fibers are stimulated
• Smooth muscle cells relax
• Vessels dilate and blood flow increases
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16-7 Visceral Reflexes Regulate the ANS
• Somatic Motor Control
• Centers in all portions of CNS
• Lowest level regulatory control
• Lower motor neurons of cranial and spinal visceral reflex
arcs
• Highest level
• Pyramidal motor neurons of primary motor cortex
• Operating with feedback from cerebellum and basal nuclei
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16-7 Visceral Reflexes Regulate the ANS
• Visceral Reflexes
• Provide automatic motor responses
• Can be modified, facilitated, or inhibited by higher centers, especially hypothalamus
• Visceral reflex arc
• Receptor
• Sensory neuron
• Processing center (one or more interneurons)
• All polysynaptic
• Two visceral motor neurons
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16-7 Visceral Reflexes Regulate the ANS
• Visceral Reflexes
• Long reflexes
• Autonomic equivalents of polysynaptic reflexes
• Visceral sensory neurons deliver information to CNS along dorsal roots of spinal nerves
• Within sensory branches of cranial nerves
• Within autonomic nerves that innervate visceral effectors
• ANS carries motor commands to visceral effectors
• Coordinate activities of entire organ
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16-7 Visceral Reflexes Regulate the ANS
• Visceral Reflexes
• Short reflexes
• Bypass CNS
• Involve sensory neurons and interneurons located within autonomic ganglia
• Interneurons synapse on ganglionic neurons
• Motor commands distributed by postganglionic fibers
• Control simple motor responses with localized effects
• One small part of target organ
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Figure 16-11 Visceral Reflexes
Stimulus
Response
Receptors inperipheral tissue
Afferent (sensory)fibers
Shortreflex
Longreflex
Peripheraleffector
Ganglionicneuron
Preganglionicneuron
Processing centerin spinal cord
(or brain)
Autonomic ganglion(sympathetic orparasympathetic)
CENTRAL NERVOUSSYSTEM
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16-7 Visceral Reflexes Regulate the ANS
• Visceral Reflexes
• Regulating visceral activity
• Most organs
• Long reflexes most important
• Digestive tract
• Short reflexes provide most control and
coordination
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16-7 Visceral Reflexes Regulate the ANS
• Visceral Reflexes
• Enteric nervous system
• Ganglia in the walls of digestive tract contain cell
bodies of:
• Visceral sensory neurons
• Interneurons
• Visceral motor neurons
• Axons form extensive nerve nets
• Control digestive functions independent of CNS
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Table 16-4 Representative Visceral Reflexes
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Table 16-4 Representative Visceral Reflexes
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16-7 Visceral Reflexes Regulate the ANS
• Higher Levels of Autonomic Control
• Simple reflexes from spinal cord provide rapid and automatic responses
• Complex reflexes coordinated in medulla oblongata
• Contains centers and nuclei involved in:
• Salivation
• Swallowing
• Digestive secretions
• Peristalsis
• Urinary function
• Regulated by hypothalamus
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16-7 Visceral Reflexes Regulate the ANS
• The Integration of SNS and ANS Activities
• Many parallels in organization and function
• Integration at brain stem
• Both systems under control of higher centers
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Figure 16-12 A Comparison of Somatic and Autonomic Function
CentralNervousSystem
PeripheralNervousSystem SNS ANS
Cerebral cortex
Thalamus
Somatic sensory
Hypothalamus
Limbicsystem
Visceralsensory
Relay and processing centers in brain stem
Somaticreflexes
Longreflexes
Lower motorneuron
PreganglionicneuronSensory
pathways
Shortreflexes
Ganglionicneuron
Skeletalmuscles
Sensoryreceptors
Visceraleffectors
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Table 16-5 A Comparison of the ANS and SNS
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16-8 Higher-Order Functions
• Higher-Order Functions Share Three Characteristics
1. Require the cerebral cortex
2. Involve conscious and unconscious information processing
3. Are not part of programmed “wiring” of brain
• Can adjust over time
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16-8 Higher-Order Functions
• Memory
• Fact memories
• Are specific bits of information
• Skill memories
• Learned motor behaviors
• Incorporated at unconscious level with repetition
• Programmed behaviors stored in appropriate area of brain stem
• Complex are stored and involve motor patterns in the basal nuclei, cerebral cortex, and cerebellum
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16-8 Higher-Order Functions
• Memory
• Short-term memories
• Information that can be recalled immediately
• Contain small bits of information
• Primary memories
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16-8 Higher-Order Functions
• Memory
• Long-term memories
• Memory consolidation – conversion from short-
term to long-term memory
• Two types of long-term memory
1. Secondary memories fade and require effort
to recall
2. Tertiary memories are with you for life
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Figure 16-13 Memory Storage
Sensoryinput
Repetitionpromotesretention
Consolidation
Permanent loss dueto neural fatigue,shock, interferenceby other stimuli
Temporary loss
Permanent loss
• Cerebral cortex (fact memory)• Cerebral cortex and cerebellar cortex (skill memory)
Short-termMemory
Long-term Memory
SecondaryMemory
TertiaryMemory
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16-8 Higher-Order Functions
• Brain Regions Involved in Memory Consolidation
and Access
• Amygdaloid body and hippocampus
• Nucleus basalis
• Cerebral cortex
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16-8 Higher-Order Functions
• Amygdaloid Body and Hippocampus
• Are essential to memory consolidation
• Damage may cause:
• Inability to convert short-term memories to new long-
term memories
• Existing long-term memories remain intact and
accessible
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16-8 Higher-Order Functions
• Nucleus Basalis
• Cerebral nucleus near diencephalon
• Plays uncertain role in memory storage and retrieval
• Tracts connect with hippocampus, amygdaloid body,
and cerebral cortex
• Damage changes emotional states, memory, and
intellectual functions
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16-8 Higher-Order Functions
• Cerebral Cortex
• Stores long-term memories
• Conscious motor and sensory memories referred to
association areas
• Occipital and temporal lobes
• Special portions crucial to memories of faces, voices, and
words
• A specific neuron may be activated by combination of
sensory stimuli associated with particular individual; called
“grandmother cells”
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16-8 Higher-Order Functions
• Cerebral Cortex
• Visual association area
• Auditory association area
• Speech center
• Frontal lobes
• Related information stored in other locations
• If storage area is damaged, memory will be incomplete
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16-8 Higher-Order Functions
• Cellular Mechanisms of Memory Formation and
Storage
• Involves anatomical and physiological changes in
neurons and synapses
• Increased neurotransmitter release
• Facilitation at synapses
• Formation of additional synaptic connections
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16-8 Higher-Order Functions
• Increased Neurotransmitter Release
• Frequently active synapse increases the amount of
neurotransmitter it stores
• Releases more on each stimulation
• The more neurotransmitter released, the greater
effect on postsynaptic neuron
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16-8 Higher-Order Functions
• Facilitation at Synapses
• Neural circuit repeatedly activated
• Synaptic terminals begin continuously releasing
neurotransmitter
• Neurotransmitter binds to receptors on postsynaptic
membrane
• Produces graded depolarization
• Brings membrane closer to threshold
• Facilitation results affect all neurons in circuit
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16-8 Higher-Order Functions
• Formation of Additional Synaptic Connections
• Neurons repeatedly communicating
• Axon tip branches and forms additional synapses on
postsynaptic neuron
• Presynaptic neuron has greater effect on
transmembrane potential of postsynaptic neuron
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16-8 Higher-Order Functions
• Cellular Mechanisms of Memory Formation and
Storage
• Basis of memory storage
• Processes create anatomical changes
• Facilitate communication along specific neural circuit
• Memory Engram
• Single circuit corresponds to single memory
• Forms as result of experience and repetition
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16-8 Higher-Order Functions
• Cellular Mechanisms of Memory Formation and
Storage
• Efficient conversion of short-term memory
• Takes at least 1 hour
• Repetition crucial
• Factors of conversion
• Nature, intensity, and frequency of original stimulus
• Strong, repeated, and exceedingly pleasant or unpleasant
events likely converted to long-term memories
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16-8 Higher-Order Functions
• Cellular Mechanisms of Memory Formation and
Storage
• Drugs stimulate CNS
• Caffeine and nicotine are examples
• Enhance memory consolidation through facilitation
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16-8 Higher-Order Functions
• Cellular Mechanisms of Memory Formation and Storage
• Drugs stimulate CNS
• NMDA (N-methyl D-aspartate) Receptors
• Linked to consolidation
• Chemically gated calcium channels
• Activated by neurotransmitter glutamate
• Gates open, calcium enters cell
• Blocking NMDA receptors in hippocampus prevents long-term memory formation
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16-8 Higher-Order Functions
• States of Consciousness
• Many gradations of states
• Degree of wakefulness indicates level of ongoing
CNS activity
• When abnormal or depressed, state of wakefulness is
affected
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16-8 Higher-Order Functions
• States of Consciousness
• Deep sleep
• Also called slow-wave or Non-REM (NREM) sleep
• Entire body relaxes
• Cerebral cortex activity minimal
• Heart rate, blood pressure, respiratory rate, and energy
utilization decline up to 30%
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16-8 Higher-Order Functions
• States of Consciousness
• Rapid eye movement (REM) sleep
• Active dreaming occurs
• Changes in blood pressure and respiratory rate
• Less receptive to outside stimuli than in deep sleep
• Muscle tone decreases markedly
• Intense inhibition of somatic motor neurons
• Eyes move rapidly as dream events unfold
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16-8 Higher-Order Functions
• States of Consciousness
• Nighttime sleep pattern
• Alternates between levels
• Begins in deep sleep
• REM periods average 5 minutes in length; increase to
20 minutes over 8 hours
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16-8 Higher-Order Functions
• Sleep
• Has important impact on CNS
• Produces only minor changes in physiological
activities of organs and systems
• Protein synthesis in neurons increases during sleep
• Extended periods without sleep lead to disturbances
in mental function
• 25% of the U.S. population experiences sleep
disorders
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Figure 16-14a Levels of Sleep
EEG from the awake, REM, and deep (slow wave)sleep states. The EEG pattern during REM sleepresembles the alpha waves typical of awake adults.
Awake
REM sleep
Deep (slowwave) sleep
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Figure 16-14b Levels of Sleep
Time
Awake
REM sleep
Transitionperiod
Deep sleep
Typical pattern of sleep stages in a healthy youngadult during a single night’s sleep.
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16-8 Higher-Order Functions
• States of Consciousness
• Arousal and the reticular activating system (RAS)
• Awakening from sleep
• Function of reticular formation
• Extensive interconnections with sensory, motor,
integrative nuclei, and pathways along brain stem
• Determined by complex interactions between reticular
formation and cerebral cortex
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16-8 Higher-Order Functions
• Reticular Activating System (RAS)
• Important brain stem component
• Diffuse network in reticular formation
• Extends from medulla oblongata to midbrain
• Output of RAS projects to thalamic nuclei that
influence large areas of cerebral cortex
• When RAS inactive, so is cerebral cortex
• Stimulation of RAS produces widespread activation of
cerebral cortex
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16-8 Higher-Order Functions
• Arousal and the Reticular Activating System
• Ending sleep
• Any stimulus activates reticular formation and RAS
• Arousal occurs rapidly
• Effects of single stimulation of RAS last less than a
minute
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16-8 Higher-Order Functions
• Arousal and the Reticular Activating System
• Maintaining consciousness
• Activity in cerebral cortex, basal nuclei, and sensory
and motor pathways continue to stimulate RAS
• After many hours, reticular formation becomes less
responsive to stimulation
• Individual becomes less alert and more lethargic
• Neural fatigue reduces RAS activity
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16-8 Higher-Order Functions
• Arousal and the Reticular Activating System
• Regulation of sleep–awake cycles
• Involves interplay between brain stem nuclei that use
different neurotransmitters
• Group of nuclei stimulates RAS with NE and maintains
awake, alert state
• Other group promotes deep sleep by depressing RAS
activity with serotonin
• “Dueling” nuclei located in brain stem
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Figure 16-15 The Reticular Activating System
Reticularformation
RAS
Special sensoryinput
General cranialor spinal nerve input
N VIII
N II
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16-9 Brain Chemistry
• Brain Chemistry
• Changes in normal balance between two or more
neurotransmitters can profoundly affect brain function
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16-9 Brain Chemistry
• Huntington’s Disease
• Destruction of ACh-secreting and GABA-secreting
neurons in basal nuclei
• Symptoms appear as basal nuclei and frontal lobes
slowly degenerate
• Difficulty controlling movements
• Intellectual abilities gradually decline
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16-9 Brain Chemistry
• Lysergic Acid Diethylamide (LSD)
• Powerful hallucinogenic drug
• Activates serotonin receptors in brain stem,
hypothalamus, and limbic system
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16-9 Brain Chemistry
• Serotonin
• Compounds that enhance effects also produce
hallucinations (LSD)
• Compounds that inhibit or block action cause severe
depression and anxiety
• Variations in levels affect sensory interpretation and
emotional states
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16-9 Brain Chemistry
• Serotonin
• Fluoxetine (Prozac)
• Slows removal of serotonin at synapses
• Increases serotonin concentrations at postsynaptic
membrane
• Classified as selective serotonin reuptake inhibitors
(SSRIs)
• Other SSRIs
• Celexa, Luvox, Paxil, and Zoloft
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16-9 Brain Chemistry
• Parkinson’s Disease
• Inadequate dopamine production causes motor
problems
• Dopamine
• Secretion stimulated by amphetamines, or “speed”
• Large doses can produce symptoms resembling
schizophrenia
• Important in nuclei that control intentional movements
• Important in other centers of diencephalon and cerebrum
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16-10 Effects of Aging on the Nervous System
• Effects of Aging
• Anatomical and physiological changes begin after
maturity (age 30)
• Accumulate over time
• 85% of people over age 65 have changes in
mental performance and CNS function
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16-10 Effects of Aging on the Nervous System
• Common Age-related Anatomical Changes in
the Nervous System
• Reduction in Brain Size and Weight
• Reduction in Number of Neurons
• Decrease in Blood Flow to Brain
• Changes in Synaptic Organization of Brain
• Intracellular and Extracellular Changes in CNS Neurons
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16-10 Effects of Aging on the Nervous System
• Reduction in Brain Size and Weight
• Decrease in volume of cerebral cortex
• Narrower gyri and wider sulci
• Larger subarachnoid space
• Reduction in Number of Neurons
• Brain shrinkage linked to loss of cortical neurons
• No neuronal loss in brain stem nuclei
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16-10 Effects of Aging on the Nervous System
• Decrease in Blood Flow to Brain
• Arteriosclerosis
• Fatty deposits in walls of blood vessels
• Reduces blood flow through arteries
• Increases chances of rupture
• Cerebrovascular accident (CVA), or stroke
• May damage surrounding neural tissue
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16-10 Effects of Aging on the Nervous System
• Changes in Synaptic Organization of Brain
• Number of dendritic branches, spines, and
interconnections decreases
• Synaptic connections lost
• Rate of neurotransmitter production declines
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16-10 Effects of Aging on the Nervous System
• Intracellular and Extracellular Changes in CNS
Neurons
• Neurons in brain accumulate abnormal intracellular
deposits
• Lipofuscin
• Granular pigment with no known function
• Neurofibrillary tangles
• Masses of neurofibrils form dense mats inside cell body
and axon
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16-10 Effects of Aging on the Nervous System
• Intracellular and Extracellular Changes in
CNS Neurons
• Plaques
• Extracellular accumulations of fibrillar proteins
• Surrounded by abnormal dendrites and axons
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16-10 Effects of Aging on the Nervous System
• Intracellular and Extracellular Changes in CNS
Neurons
• Plaques and tangles
• Contain deposits of several peptides
• Primarily two forms of amyloid ß (Aß) protein
• Appear in brain regions specifically associated with
memory processing
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16-10 Effects of Aging on the Nervous System
• Anatomical Changes
• Linked to functional changes
• Neural processing becomes less efficient with age
• Memory consolidation more difficult
• Secondary memories harder to access
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16-10 Effects of Aging on the Nervous System
• Sensory Systems
• Hearing, balance, vision, smell, and taste become
less acute
• Reaction times slowed
• Reflexes weaken or disappear
• Motor Control
• Precision decreases
• Takes longer to perform
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16-10 Effects of Aging on the Nervous System
• Incapacitation
• 85% of elderly population develops changes that
do not interfere with abilities
• Some individuals become incapacitated by
progressive CNS changes
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16-10 Effects of Aging on the Nervous System
• Senility
• Also called senile dementia
• Degenerative changes
• Memory loss
• Anterograde amnesia (lose ability to store new
memories)
• Emotional disturbances
• Alzheimer’s disease is most common
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16-10 Nervous System Integration
• The Nervous System
• Monitors all other systems
• Issues commands that adjust their activities
• Like conductor of orchestra
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16-10 Nervous System Integration
• Neural Tissue
• Extremely delicate
• Extracellular environment must maintain homeostatic
limits
• If regulatory mechanisms break down, neurological
disorders appear
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Figure 16-16 System Integrator: The Nervous System
Provides sensations of touch, pressure,pain, vibration, and temperature; hairprovides some protection and insulationfor skull and brain; protects peripheral nerves
Provides calcium for neural function;protects brain and spinal cord
Facial muscles express emotionalstate; intrinsic laryngeal musclespermit communication; musclespindles provide proprioceptivesensations
Body System Nervous System Nervous System Body SystemS Y S T E M I N T E G R A T O R
Controls contraction of arrector pilimuscles and secrection of sweat glands
Controls skeletal muscle contractionsthat results in bone thickening andmaintenance and determine boneposition
Controls skeletal muscle contractions;coordinates respiratory andcardiovascular activities
The Nervous System
The nervous system is closelyintegrated with other body systems.Every moment of your life, billions ofneurons in your nervous systemare exchanging informationacross trillions of synapsesand performing the mostcomplex integrativefunctions in the body. Aspart of this process, thenervous system monitors allother systems and issuescommands that adjust theiractivities. However, thesignificance and impact ofthese commands varies greatly from one system toanother. The normal functions of themuscular system, for example, simplycannot be performed withoutinstructions from the nervous system.By contrast, the cardiovascular systemis relatively independent—the nervoussystem merely coordinates and adjustscardiovascular activities to meet thecirculatory demands of other systems. Inthe final analysis, the nervous system islike the conductor of an orchestra,directing the rhythm and balancing the performances of each section toproduce a symphony, instead of simplya very loud noise.
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