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Copyright © 2010 Pearson Education, Inc.
Chapter 13
The Peripheral Nervous System
Part D
Shilla Chakrabarty, Ph.D.
Copyright © 2010 Pearson Education, Inc.
Motor Endings
• Are PNS elements that activate effectors by releasing neurotransmitters
• Found at neuromuscular junctions
Terminals of somatic fibers innervating voluntary muscles forms elaborate neuromuscular junctions with their effector cells
As each axon branch reaches its target, a single muscle cell, the ending splits into a cluster of axon terminals
Axon terminals branch like a tree over the junctional folds of sarcolemma of muscle fiber
Axon terminals contain mitochondria and synaptic vesicles filled with the neurotransmitter acetylcholine (ACh)
When a nerve impulse reaches an axon terminal, Ach is released by exocytosis and a series of events is initiated.
Copyright © 2010 Pearson Education, Inc.
Nucleus
Actionpotential (AP)
Myelinated axonof motor neuron
Axon terminal of neuromuscular junction
Sarcolemma ofthe muscle fiber
Ca2+Ca2+
Axon terminalof motor neuron
Synaptic vesiclecontaining ACh
Mitochondrion
Synaptic cleft
Junctionalfolds of sarcolemma
Fusing synaptic vesicles
ACh
Sarcoplasm ofmuscle fiber
Postsynaptic membraneion channel opens;ions pass.
Na+ K+
AChNa+
K+
Degraded ACh
Acetylcholinesterase
Postsynaptic membraneion channel closed;ions cannot pass.
Action potential arrives at axon terminal of motor neuron.
Voltage-gated Ca2+
channels open and Ca2+
enters the axon terminal.
Ca2+ entry causes some synaptic vesicles to release their contents (acetylcholine)by exocytosis.
Acetylcholine, a neurotransmitter, diffuses across the synaptic cleft and binds to receptors in the sarcolemma.
ACh binding opens ion channels that allow simultaneous passage of Na+ into the muscle fiber and K+ out of the muscle fiber.
ACh effects are terminated by its enzymatic breakdown in the synaptic cleft by acetylcholinesterase.
1
2
3
4
5
6
Events At A Neuromuscular Junction When A Nerve Impulse Arrives
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Review of Innervation of Visceral Muscle and Glands
• Autonomic motor endings and visceral effectors (such as smooth and cardiac muscles and glands) are simpler than somatic junctions
• Branches form synapses en passant via varicosities
Varicosities are knoblike swellings containing mitochondria and synaptic vesicles, and appear like a string of beads
• Autonomic synaptic vesicles typically contain acetylcholine or norepinephrine, both of which act indirectly via second messengers
• Visceral motor responses are slower than somatic responses which directly open ion channels
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Smoothmusclecell
Varicosities releasetheir neurotransmittersinto a wide synaptic cleft (a diffuse junction).
Synapticvesicles
Mitochondrion
Autonomicnerve fibersinnervatemost smoothmuscle fibers.
Varicosities
Innervations Of Smooth Muscles
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Feedback
Reflex activity Motoroutput
Sensoryinput
Precommand Level(highest)• Cerebellum and basal nuclei• Programs and instructions (modified by feedback)
Projection Level (middle)
• Motor cortex (pyramidal system) and brain stem nuclei (vestibular, red, reticular formation, etc.)• Convey instructions to spinal cord motor neurons and send a copy of that information to higher levels
Segmental Level (lowest)• Spinal cord• Contains central pattern generators (CPGs)
Internalfeedback
Levels of Motor Control And Their Interactions
Copyright © 2010 Pearson Education, Inc.
Levels of Motor Control: Segmental Level
• The lowest level of the motor hierarchy
• Central pattern generators (CPGs): segmental circuits that activate networks of ventral horn neurons to stimulate specific groups of muscles
• Controls locomotion and specific, oft-repeated motor activity
Copyright © 2010 Pearson Education, Inc.
Levels of Motor Control: Projection Level
• Consists of:
Upper motor neurons that direct the pyramidal system to directly produce voluntary skeletal muscle movements
Brain stem motor areas that oversee the indirect (extrapyramidal) system to control reflex and CPG-controlled motor actions
• Projection motor pathways keep higher command levels informed of what is happening
Copyright © 2010 Pearson Education, Inc.
Levels of Motor Control: Precommand Level
• Neurons in the cerebellum and basal nuclei
Regulate motor activity
Precisely start or stop movements
Coordinate movements with posture
Block unwanted movements
Monitor muscle tone
Perform unconscious planning and discharge in advance of willed movements
Copyright © 2010 Pearson Education, Inc.
Precommand Level
• Cerebellum
• Lacks direct connections to the spinal cord
• Acts on motor pathways through projection areas of the brain stem
• Acts on the motor cortex via the thalamus
• Basal nuclei
• Receive inputs from all cortical areas and send output to premotor and prefrontal cortical areas via thalamus
• Inhibit various motor centers under resting conditions
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(b) Structures involved
Precommand level • Cerebellum• Basal nuclei
Projection level • Primary motor cortex• Brain stem nuclei
Segmental level • Spinal cord
Levels of Motor Control: Segmental, Projection and Precommand
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Reflexes
• Inborn (intrinsic) reflex: a rapid, involuntary, predictable motor response to a stimulus
• Learned (acquired) reflexes result from practice or repetition,
Example: driving skills
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Reflex Arc• Reflexes occur over highly specific neural paths
called reflex arcs
• Components of a reflex arc (neural path)
1. Receptor—site of stimulus action
2. Sensory neuron—transmits afferent impulses to the CNS
3. Integration center—either monosynaptic or polysynaptic region within the CNS
4. Motor neuron—conducts efferent impulses from the integration center to an effector organ
5. Effector—muscle fiber or gland cell that responds to the efferent impulses by contracting or secreting
Copyright © 2010 Pearson Education, Inc. Figure 13.14
Receptor
Sensory neuron
Integration center
Motor neuron
Effector
Spinal cord(in cross section)
Interneuron
Stimulus
Skin
1
2
3
4
5
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Spinal Reflexes
• Spinal somatic reflexes
• Integration center is in the spinal cord
• Effectors are skeletal muscle
• Testing of somatic reflexes is important clinically to assess the condition of the nervous system
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Stretch and Golgi Tendon Reflexes
• For skeletal muscle activity to be smoothly coordinated, proprioceptor input is necessary
Muscle spindles inform the nervous system of the length of the muscle
Golgi tendon organs inform the brain as to the amount of tension in the muscle and tendons
Copyright © 2010 Pearson Education, Inc.
Secondary sensoryendings (type II fiber)
Efferent (motor)fiber to muscle spindle
Primary sensoryendings (type Iafiber)
Connectivetissue capsule
Muscle spindle
Tendon
Sensory fiber
Golgi tendonorgan
Efferent (motor)fiber to extrafusalmuscle fibers
Extrafusal musclefiber
Intrafusal musclefibers
Muscle Spindles
• Composed of 3–10 short intrafusal muscle fibers in a connective tissue capsule
• Intrafusal fibers
Noncontractile in their central regions (lack myofilaments)
Wrapped with two types of afferent endings: primary sensory endings of type Ia fibers and secondary sensory endings of type II fibers
• Contractile end regions are innervated by gamma () efferent fibers that maintain spindle sensitivity
Note: Extrafusal fibers (contractile muscle fibers) are innervated by alpha () efferent fibers
Copyright © 2010 Pearson Education, Inc.
Operation Of The Muscle Spindle
(a) Unstretched muscle. Action potentials (APs) are generated at a constant rate in the associated sensory (la) fiber.
Musclespindle
Intrafusalmuscle fiber
Primarysensory (la)nerve fiberExtrafusalmuscle fiber
Time
(b) Stretched muscle. Stretching activates the muscle spindle, increasing the rate of APs.
Time
(d) Coactivation. Both extrafusal and intrafusal muscle fibers contract. Muscle spindle tension is main- tained and it can still signal changes in length.
Time
(c) Only motor neurons activated. Only the extrafusal muscle fibers contract. The muscle spindle becomes slack and no APs are fired. It is unable to signal further length changes.
Time
Action potentials generated in sensory fibers are shown as black lines in yellow bars
Copyright © 2010 Pearson Education, Inc.
Stretch Reflexes• Maintain muscle tone in large postural muscles
• Cause muscle contraction in response to increased muscle length (stretch)
How a stretch reflex works:
• Stretch activates the muscle spindle
• IIa sensory neurons synapse directly with motor neurons in the spinal cord
• motor neurons cause the stretched muscle to contract
• All stretch reflexes are monosynaptic and ipsilateral
• Reciprocal inhibition also occurs—IIa fibers synapse with interneurons that inhibit the motor neurons of antagonistic muscles
Example: In the patellar reflex, the stretched muscle (quadriceps) contracts and the antagonists (hamstrings) relax
Copyright © 2010 Pearson Education, Inc. Figure 13.17 (1 of 2)
Stretched muscle spindles initiate a stretch reflex,causing contraction of the stretched muscle andinhibition of its antagonist.
When muscle spindles are activatedby stretch, the associated sensoryneurons (blue) transmit afferent impulsesat higher frequency to the spinal cord.
The sensory neurons synapse directly with alphamotor neurons (red), which excite extrafusal fibersof the stretched muscle. Afferent fibers alsosynapse with interneurons (green) that inhibit motorneurons (purple) controlling antagonistic muscles.
The events by which muscle stretch is damped
Efferent impulses of alpha motor neuronscause the stretched muscle to contract,which resists or reverses the stretch.
Efferent impulses of alpha motorneurons to antagonist muscles arereduced (reciprocal inhibition).
Initial stimulus(muscle stretch)
Cell body ofsensory neuron
Sensoryneuron
Muscle spindleAntagonist muscle
Spinal cord
12
3a 3b
Copyright © 2010 Pearson Education, Inc. Figure 13.17 (2 of 2)
The patellar (knee-jerk) reflex—a specific example of a stretch reflex
Musclespindle
Quadriceps(extensors)
Hamstrings(flexors)
Patella
Patellarligament
Spinal cord(L2–L4)
Tapping the patellar ligament excitesmuscle spindles in the quadriceps.
The motor neurons (red) sendactivating impulses to the quadricepscausing it to contract, extending theknee.
Afferent impulses (blue) travel to thespinal cord, where synapses occur withmotor neurons and interneurons.
The interneurons (green) makeinhibitory synapses with ventral horn neurons (purple) that prevent theantagonist muscles (hamstrings) fromresisting the contraction of thequadriceps.
Excitatory synapseInhibitory synapse
+
–
1
2
3a
3b
1
2
3a3b 3b
Copyright © 2010 Pearson Education, Inc. Figure 13.17 (2 of 2), step 3b
The patellar (knee-jerk) reflex—a specific example of a stretch reflex
Musclespindle
Quadriceps(extensors)
Hamstrings(flexors)
Patella
Patellarligament
Spinal cord(L2–L4)
Tapping the patellar ligament excitesmuscle spindles in the quadriceps.
The motor neurons (red) sendactivating impulses to the quadricepscausing it to contract, extending theknee.
Afferent impulses (blue) travel to thespinal cord, where synapses occur withmotor neurons and interneurons.
The interneurons (green) makeinhibitory synapses with ventral horn neurons (purple) that prevent theantagonist muscles (hamstrings) fromresisting the contraction of thequadriceps.
Excitatory synapseInhibitory synapse
+
–
1
2
3a
3b
1
2
3a3b 3b
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Golgi Tendon Reflexes
• Polysynaptic reflexes
• Help to prevent damage due to excessive stretch
• Important for smooth onset and termination of muscle contraction
Copyright © 2010 Pearson Education, Inc.
Golgi Tendon Reflexes
• Produce muscle relaxation (lengthening) in response to tension
• Contraction or passive stretch activates Golgi tendon organs
• Afferent impulses are transmitted to spinal cord
• Contracting muscle relaxes and the antagonist contracts (reciprocal activation)
• Information transmitted simultaneously to the cerebellum is used to adjust muscle tension
Copyright © 2010 Pearson Education, Inc. Figure 13.18
+ Excitatory synapse– Inhibitory synapse
Quadriceps strongly contracts. Golgi tendon organs are activated.
Afferent fibers synapse with interneurons in the spinal cord.
Efferent impulses to muscle with stretched tendon are damped. Muscle relaxes, reducing tension.
Efferent impulses to antagonist muscle cause it to contract.
Interneurons
Spinal cord
Quadriceps(extensors)
Golgitendon
organHamstrings
(flexors)
1 2
3a 3b
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Flexor and Crossed-Extensor Reflexes
• Flexor (withdrawal) reflex
• Initiated by a painful stimulus
• Causes automatic withdrawal of the threatened body part
• Ipsilateral and polysynaptic
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Flexor and Crossed-Extensor Reflexes
• Crossed extensor reflex
• Occurs with flexor reflexes in weight-bearing limbs to maintain balance
• Consists of an ipsilateral flexor reflex and a contralateral extensor reflex
• The stimulated side is withdrawn (flexed)
• The contralateral side is extended
Copyright © 2010 Pearson Education, Inc. Figure 13.19
Afferentfiber
Efferentfibers
Extensorinhibited
Flexorstimulated
Site of stimulus: a noxiousstimulus causes a flexorreflex on the same side,withdrawing that limb.
Site of reciprocalactivation: At thesame time, theextensor muscleson the oppositeside are activated.
Armmovements
Interneurons
Efferentfibers
FlexorinhibitedExtensorstimulated
+ Excitatory synapse– Inhibitory synapse
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Superficial Reflexes
• Elicited by gentle cutaneous stimulation
• Depend on upper motor pathways and cord-level reflex arcs
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Superficial Reflexes: Plantar Reflex
• Plantar reflex
Stimulus: stroking lateral aspect of the sole of the foot
Response: downward flexion of the toes
Tests for function of corticospinal tracts
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Superficial Reflexes
• Babinski’s sign
• Stimulus: as above
• Response: dorsiflexion of hallux and fanning of toes
• Present in infants due to incomplete myelination
• In adults, indicates corticospinal or motor cortex damage
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Superficial Reflexes
• Abdominal reflexes
• Cause contraction of abdominal muscles and movement of the umbilicus in response to stroking of the skin
• Vary in intensity from one person to another
• Absent when corticospinal tract lesions are present
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Developmental Aspects of the PNS
• Spinal nerves branch from the developing spinal cord and neural crest cells
• Supply both motor and sensory fibers to developing muscles to help direct their maturation
• Cranial nerves innervate muscles of the head
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Developmental Aspects of the PNS
• Distribution and growth of spinal nerves correlate with the segmented body plan
• Sensory receptors atrophy with age and muscle tone lessens due to loss of neurons, decreased numbers of synapses per neuron, and slower central processing
• Peripheral nerves remain viable throughout life unless subjected to trauma