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Transcript
Chapter 12
Neural Tissue
An Introduction to the Nervous System
•
Learning Outcomes
o 12-1 Describe the anatomical and functional divisions of the nervous
system.
o 12-2 Sketch and label the structure of a typical neuron, describe the
functions of each component, and classify neurons on the basis of
their structure and function.
o 12-3 Describe the locations and functions of the various types of
neuroglia.
An Introduction to the Nervous System
•
Learning Outcomes
o 12-4 Explain how the resting membrane potential is created and
maintained.
o 12-5 Describe the events involved in the generation and propagation of
an action potential.
o 12-6 Discuss the factors that affect the speed with which action
potentials are propagated.
An Introduction to the Nervous System
•
Learning Outcomes
o 12-7 Describe the structure of a synapse, and explain the mechanism
involved in synaptic activity.
o 12-8 Describe the major types of neurotransmitters and
neuromodulators, and discuss their effects on postsynaptic
membranes.
o 12-9 Discuss the interactions that enable information processing to occur
in neural tissue.
An Introduction to the Nervous System
•
The Nervous System
o Includes all neural tissue in the body
o Neural tissue contains two kinds of cells
1. Neurons
o Cells that send and receive signals
2. Neuroglia (glial cells)
o Cells that support and protect neurons
© 2015 Pearson Education, Inc.
An Introduction to the Nervous System
•
Organs of the Nervous System
o Brain and spinal cord
o Sensory receptors of sense organs (eyes, ears, etc.)
o Nerves connect nervous system with other systems
12-1 Divisions of the Nervous System
•
Anatomical Divisions of the Nervous System
o Central nervous system (CNS)
o Peripheral nervous system (PNS)
12-1 Divisions of the Nervous System
•
The Central Nervous System (CNS)
o Consists of the spinal cord and brain
o Contains neural tissue, connective tissues, and blood vessels
o Functions of the CNS are to process and coordinate:
 Sensory data from inside and outside body
 Motor commands control activities of peripheral organs (e.g., skeletal
muscles)
 Higher functions of brain: intelligence, memory, learning, emotion
12-1 Divisions of the Nervous System
•
The Peripheral Nervous System (PNS)
o Includes all neural tissue outside the CNS
o Functions of the PNS
 Deliver sensory information to the CNS
 Carry motor commands to peripheral tissues and systems
12-1 Divisions of the Nervous System
•
The Peripheral Nervous System (PNS)
o Nerves (also called peripheral nerves)
 Bundles of axons with connective tissues and blood vessels
 Carry sensory information and motor commands in PNS
o Cranial nerves – connect to brain
o Spinal nerves – attach to spinal cord
12-1 Divisions of the Nervous System
•
Functional Divisions of the PNS
o Afferent division
 Carries sensory information
 From PNS sensory receptors to CNS
o Efferent division
© 2015 Pearson Education, Inc.


Carries motor commands
From CNS to PNS muscles and glands
12-1 Divisions of the Nervous System
•
Functional Divisions of the PNS
o Receptors and effectors of afferent division
 Receptors
o Detect changes or respond to stimuli
o Neurons and specialized cells
o Complex sensory organs (e.g., eyes, ears)
 Effectors
o Respond to efferent signals
o Cells and organs
12-1 Divisions of the Nervous System
•
Functional Divisions of the PNS
o The efferent division
 Somatic nervous system (SNS)
o Controls voluntary and involuntary (reflexes) skeletal muscle
contractions
12-1 Divisions of the Nervous System
•
Functional Divisions of the PNS
o The efferent division
 Autonomic nervous system (ANS)
o Controls subconscious actions, contractions of smooth muscle and
cardiac muscle, and glandular secretions
o Sympathetic division has a stimulating effect
o Parasympathetic division has a relaxing effect
12-2 Neurons
•
Neurons
o The basic functional units of the nervous system
o The structure of neurons
 The multipolar neuron
o Common in the CNS
 Cell body (soma)
 Short, branched dendrites
 Long, single axon
12-2 Neurons
•
The Cell Body
© 2015 Pearson Education, Inc.
o
o
o
o
Large nucleus and nucleolus
Perikaryon (cytoplasm)
Mitochondria (produce energy)
RER and ribosomes (produce neurotransmitters)
12-2 Neurons
•
The Cell Body
o Cytoskeleton
 Neurofilaments and neurotubules in place of microfilaments and
microtubules
 Neurofibrils: bundles of neurofilaments that provide support for
dendrites and axon
o Nissl bodies
 Dense areas of RER and ribosomes
 Make neural tissue appear gray (gray matter)
12-2 Neurons
•
Dendrites
o Highly branched
o Dendritic spines
 Many fine processes
 Receive information from other neurons
 80–90 percent of neuron surface area
12-2 Neurons
•
The axon
o Is long
o Carries electrical signal (action potential) to target
o Axon structure is critical to function
o
12-2 Neurons
•
Structures of the Axon
o Axoplasm
 Cytoplasm of axon
 Contains neurofibrils, neurotubules, enzymes, organelles
o Axolemma
 Specialized cell membrane
 Covers the axoplasm
12-2 Neurons
•
Structures of the Axon
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o Axon hillock
 Thick section of cell body
 Attaches to initial segment
o Initial segment
 Attaches to axon hillock
12-2 Neurons
•
Structures of the Axon
o Collaterals
 Branches of a single axon
o Telodendria
 Fine extensions of distal axon
o Axon terminals
 Tips of telodendria
12-2 Neurons
•
The Structure of Neurons
o The synapse
 Area where a neuron communicates with another cell
12-2 Neurons
•
The Structure of Neurons
o The synapse
 Presynaptic cell
o Neuron that sends message
 Postsynaptic cell
o Cell that receives message
 The synaptic cleft
o The small gap that separates the presynaptic membrane and the
postsynaptic membrane
12-2 Neurons
•
The Synapse
o The synaptic terminal
 Is expanded area of axon of presynaptic neuron
 Contains synaptic vesicles of neurotransmitters
12-2 Neurons
•
Neurotransmitters
o Are chemical messengers
o Are released at presynaptic membrane
o Affect receptors of postsynaptic membrane
© 2015 Pearson Education, Inc.
o Are broken down by enzymes
o Are reassembled at axon terminal
12-2 Neurons
•
Recycling Neurotransmitters
o Axoplasmic transport
 Neurotubules within the axon
 Transport raw materials
 Between cell body and axon terminal
 Powered by mitochondria, kinesin, and dynein
12-2 Neurons
•
Types of Synapses
o Neuromuscular junction
 Synapse between neuron and muscle
o Neuroglandular junction
 Synapse between neuron and gland
12-2 Neurons
•
Structural Classification of Neurons
o Anaxonic neurons
 Found in brain and sense organs
o Bipolar neurons
 Found in special sensory organs (sight, smell, hearing)
o Unipolar neurons
 Found in sensory neurons of PNS
o Multipolar neurons
 Common in the CNS
 Include all skeletal muscle motor neurons
12-2 Neurons
•
•
Anaxonic Neurons
o Small
o All cell processes look alike
Bipolar Neurons
o Are small
o One dendrite, one axon
12-2 Neurons
•
Unipolar Neurons
o Also called pseudounipolar neurons
o Have very long axons
© 2015 Pearson Education, Inc.
•
o Fused dendrites and axon
o Cell body to one side
Multipolar Neurons
o Have very long axons
o Multiple dendrites, one axon
12-2 Neurons
•
Three Functional Classifications of Neurons
1. Sensory neurons
 Afferent neurons of PNS
2. Motor neurons
 Efferent neurons of PNS
3. Interneurons
 Association neurons
12-2 Neurons
•
•
Functions of Sensory Neurons
o Monitor internal environment (visceral sensory neurons)
o Monitor effects of external environment (somatic sensory neurons)
Structures of Sensory Neurons
o Unipolar
o Cell bodies grouped in sensory ganglia
o Processes (afferent fibers) extend from sensory receptors to CNS
12-2 Neurons
•
Three Types of Sensory Receptors
1. Interoceptors
 Monitor internal systems (digestive, respiratory, cardiovascular, urinary,
reproductive)
 Internal senses (taste, deep pressure, pain)
2. Exteroceptors
 External senses (touch, temperature, pressure)
 Distance senses (sight, smell, hearing)
3. Proprioceptors
 Monitor position and movement (skeletal muscles and joints)
12-2 Neurons
•
Motor Neurons
o Carry instructions from CNS to peripheral effectors
o Via efferent fibers (axons)
12-2 Neurons
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•
Motor Neurons
o Two major efferent systems
1. Somatic nervous system (SNS)
o Includes all somatic motor neurons that innervate skeletal muscles
2. Autonomic (visceral) nervous system (ANS)
o Visceral motor neurons innervate all other peripheral effectors
 Smooth muscle, cardiac muscle, glands, adipose tissue
12-2 Neurons
•
Motor Neurons
o Two groups of efferent axons
 Signals from CNS motor neurons to visceral effectors pass synapses
at autonomic ganglia dividing axons into:
o Preganglionic fibers
o Postganglionic fibers
12-2 Neurons
•
Interneurons
o Most are located in brain, spinal cord, and autonomic ganglia
 Between sensory and motor neurons
o Are responsible for:
 Distribution of sensory information
 Coordination of motor activity
o Are involved in higher functions
 Memory, planning, learning
12-3 Neuroglia
•
Neuroglia
o Half the volume of the nervous system
o Many types of neuroglia in CNS and PNS
12-3 Neuroglia
•
Four Types of Neuroglia in the CNS
1. Ependymal cells
 Cells with highly branched processes; contact neuroglia directly
2. Astrocytes
 Large cell bodies with many processes
3. Oligodendrocytes
 Smaller cell bodies with fewer processes
4. Microglia
 Smallest and least numerous neuroglia with many fine-branched
processes
© 2015 Pearson Education, Inc.
12-3 Neuroglia
•
Ependymal Cells
o Form epithelium called ependyma
o Line central canal of spinal cord and ventricles of brain
 Secrete cerebrospinal fluid (CSF)
 Have cilia or microvilli that circulate CSF
 Monitor CSF
 Contain stem cells for repair
12-3 Neuroglia
•
Astrocytes
o Maintain blood–brain barrier (isolates CNS)
o Create three-dimensional framework for CNS
o Repair damaged neural tissue
o Guide neuron development
o Control interstitial environment
12-3 Neuroglia
•
Oligodendrocytes
o Myelination
 Increases speed of action potentials
 Myelin insulates myelinated axons
 Makes nerves appear white
12-3 Neuroglia
•
Oligodendrocytes
o Nodes and internodes
 Internodes – myelinated segments of axon
 Nodes (also called nodes of Ranvier)
o Gaps between internodes
o Where axons may branch
12-3 Neuroglia
•
Myelination
o White matter
 Regions of CNS with many myelinated nerves
o Gray matter
 Unmyelinated areas of CNS
12-3 Neuroglia
•
Microglia
o Migrate through neural tissue
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o Clean up cellular debris, waste products, and pathogens
12-3 Neuroglia
•
Neuroglia of the Peripheral Nervous System
o Ganglia
 Masses of neuron cell bodies
 Surrounded by neuroglia
 Found in the PNS
12-3 Neuroglia
•
Neuroglia of the Peripheral Nervous System
o Satellite cells
 Also called amphicytes
 Surround ganglia
 Regulate environment around neuron
12-3 Neuroglia
•
Neuroglia of the Peripheral Nervous System
o Schwann cells
 Also called neurilemma cells
 Form myelin sheath (neurilemma) around peripheral axons
 One Schwann cell sheaths one segment of axon
o Many Schwann cells sheath entire axon
12-3 Neuroglia
•
Neurons and Neuroglia
o Neurons perform:
 All communication, information processing, and control functions of the
nervous system
o Neuroglia preserve:
 Physical and biochemical structure of neural tissue
o Neuroglia are essential to:
 Survival and function of neurons
12-3 Neuroglia
•
Neural Responses to Injuries
o Wallerian degeneration
 Axon distal to injury degenerates
o Schwann cells
 Form path for new growth
 Wrap new axon in myelin
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12-3 Neuroglia
•
Nerve Regeneration in CNS
o Limited by chemicals released by astrocytes that:
 Block growth
 Produce scar tissue
12-4 Membrane Potential
•
Ion Movements and Electrical Signals
o All plasma (cell) membranes produce electrical signals by ion movements
o Membrane potential is particularly important to neurons
12-4 Membrane Potential
•
Five Main Membrane Processes in Neural Activities
1. Resting potential
 The membrane potential of resting cell
2. Graded potential
 Temporary, localized change in resting potential
 Caused by stimulus
12-4 Membrane Potential
•
Five Main Membrane Processes in Neural Activities
3. Action potential
 Is an electrical impulse
 Produced by graded potential
 Propagates along surface of axon to synapse
12-4 Membrane Potential
•
Five Main Membrane Processes in Neural Activities
4. Synaptic activity
 Releases neurotransmitters at presynaptic membrane
 Produces graded potentials in postsynaptic membrane
5. Information processing
 Response (integration of stimuli) of postsynaptic cell
12-4 Membrane Potential
•
The Membrane Potential
o Three important concepts
1. The extracellular fluid (ECF) and intracellular fluid (cytosol) differ
greatly in ionic composition
o Concentration gradient of ions (Na+, K+)
2. Cells have selectively permeable membranes
3. Membrane permeability varies by ion
© 2015 Pearson Education, Inc.
12-4 Membrane Potential
•
Passive Forces Acting across the Plasma Membrane
o Chemical gradients
 Concentration gradients (chemical gradient) of ions (Na+, K+)
o Electrical gradients
 Separate charges of positive and negative ions
 Result in potential difference
12-4 Membrane Potential
•
Electrical Currents and Resistance
o Electrical current
 Movement of charges to eliminate potential difference
o Resistance
 The amount of current a membrane restricts
12-4 Membrane Potential
•
The Electrochemical Gradient
o For a particular ion (Na+, K+) is:
 The sum of chemical and electrical forces
o Acting on the ion across a plasma membrane
 A form of potential energy
12-4 Membrane Potential
•
Equilibrium Potential
o The membrane potential at which there is no net movement of a particular
ion across the cell membrane
o Examples:
 K+ 
 Na+  66 mV
12-4 Membrane Potential
•
Active Forces across the Membrane
o Sodium–potassium ATPase (exchange pump)
 Is powered by ATP
 Carries 3 Na+ out and 2 K+ in
 Balances passive forces of diffusion
 Maintains resting potential (70 mV)
12-4 Membrane Potential
•
The Resting Potential
o Because the plasma membrane is highly permeable to potassium ions:
 The resting potential of approximately 70 mV is fairly close to 90 mV,
© 2015 Pearson Education, Inc.
the equilibrium potential for K+
o The electrochemical gradient for sodium ions is very large, but the
membrane’s permeability to these ions is very low
 Na+ has only a small effect on the normal resting potential, making it
just slightly less negative than the equilibrium potential for K +
12-4 Membrane Potential
•
The Resting Potential
o The sodium–potassium exchange pump ejects 3 Na+ ions for every 2 K+
ions that it brings into the cell
 It serves to stabilize the resting potential when the ratio of Na + entry to
K+ loss through passive channels is 3:2
o At the normal resting potential, these passive and active mechanisms are
in balance
 The resting potential varies widely with the type of cell
 A typical neuron has a resting potential of approximately 70 mV
12-4 Membrane Potential
•
Changes in the Membrane Potential
o Membrane potential rises or falls
 In response to temporary changes in membrane permeability
 Resulting from opening or closing specific membrane channels
12-4 Membrane Potential
•
Sodium and Potassium Channels
o Membrane permeability to Na+ and K+ determines membrane potential
o They are either passive or active
12-4 Membrane Potential
•
•
Passive Channels (Leak Channels)
o Are always open
o Permeability changes with conditions
Active Channels (Gated Channels)
o Open and close in response to stimuli
o At resting potential, most gated channels are closed
12-4 Membrane Potential
•
Three States of Gated Channels
1. Closed, but capable of opening
2. Open (activated)
3. Closed, not capable of opening (inactivated)
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12-4 Membrane Potential
•
Three Classes of Gated Channels
1. Chemically gated channels
2. Voltage-gated channels
3. Mechanically gated channels
12-4 Membrane Potential
•
Chemically Gated Channels
o Open in presence of specific chemicals (e.g., ACh) at a binding site
o Found on neuron cell body and dendrites
12-4 Membrane Potential
•
Voltage-gated Channels
o Respond to changes in membrane potential
o Have activation gates (open) and inactivation gates (close)
o Characteristic of excitable membrane
o Found in neural axons, skeletal muscle sarcolemma, cardiac muscle
12-4 Membrane Potential
•
Mechanically Gated Channels
o Respond to membrane distortion
o Found in sensory receptors (touch, pressure, vibration)
12-4 Membrane Potential
•
•
Membrane Potential Exists across Plasma Membrane
o Because:
 Cytosol and extracellular fluid have different chemical/ionic balance
 The plasma membrane is selectively permeable
Membrane Potential
o Changes with plasma membrane permeability
o In response to chemical or physical stimuli
12-4 Membrane Potential
•
Graded Potentials
o Also called local potentials
o Changes in membrane potential
 That cannot spread far from site of stimulation
o Any stimulus that opens a gated channel
 Produces a graded potential
12-4 Membrane Potential
© 2015 Pearson Education, Inc.
•
Graded Potentials
o The resting state
 Opening sodium channel produces graded potential
o Resting membrane exposed to chemical
o Sodium channel opens
o Sodium ions enter the cell
o Membrane potential rises
o Depolarization occurs
12-4 Membrane Potential
•
Graded Potentials
o Depolarization
 A shift in membrane potential toward 0 mV
o Movement of Na+ through channel
o Produces local current
o Depolarizes nearby plasma membrane (graded potential)
o Change in potential is proportional to stimulus
12-4 Membrane Potential
•
Graded Potentials
o Whether depolarizing or hyperpolarizing, share four basic characteristics
1. The membrane potential is most changed at the site of stimulation, and
the effect decreases with distance
2. The effect spreads passively, due to local currents
12-4 Membrane Potential
•
Graded Potentials
o Whether depolarizing or hyperpolarizing, share
four basic characteristics
3. The graded change in membrane potential may involve either
depolarization or hyperpolarization
 The properties and distribution of the membrane channels involved
determine the nature of the change
o For example, in a resting membrane, the opening of sodium
channels causes depolarization, whereas the opening of potassium
channels causes hyperpolarization
 The change in membrane potential reflects
whether positive charges enter or leave the cell
12-4 Membrane Potential
•
Graded Potentials
o Whether depolarizing or hyperpolarizing, share four basic characteristics
4. The stronger the stimulus, the greater the change in the membrane
© 2015 Pearson Education, Inc.
potential and the larger the area affected
12-4 Membrane Potential
•
Graded Potentials
o Repolarization
 When the stimulus is removed, membrane potential returns to normal
o Hyperpolarization
 Increasing the negativity of the resting potential
 Result of opening a potassium channel
 Opposite effect of opening a sodium channel
 Positive ions move out, not into cell
12-4 Membrane Potential
•
Graded Potentials
o Effects of graded potentials
 At cell dendrites or cell bodies
o Trigger specific cell functions
o For example, exocytosis of glandular secretions
 At motor end plate
o Release ACh into synaptic cleft
12-5 Action Potential
•
Action Potentials
o Propagated changes in membrane potential
o Affect an entire excitable membrane
o Link graded potentials at cell body with motor end plate actions
12-5 Action Potential
•
Initiating Action Potential
o Initial stimulus
 A graded depolarization of axon hillock large enough (10 to 15 mV) to
change resting potential (70 mV) to threshold level of voltage-gated
sodium channels (60 to 55 mV)
12-5 Action Potential
•
Initiating Action Potential
o All-or-none principle
 If a stimulus exceeds threshold amount
o The action potential is the same
o No matter how large the stimulus
 Action potential is either triggered, or not
© 2015 Pearson Education, Inc.
12-5 Action Potential
•
Four Steps in the Generation of Action Potentials
o Step 1: Depolarization to threshold
o Step 2: Activation of Na channels
o Step 3: Inactivation of Na channels and activation of K channels
o Step 4: Return to normal permeability
12-5 Action Potential
•
•
Step 1: Depolarization to Threshold
Step 2: Activation of Na Channels
o Rapid depolarization
o Na+ ions rush into cytoplasm
o Inner membrane changes from negative to positive
12-5 Action Potential
•
Step 3: Inactivation of Na Channels and Activation of K Channels
o At 30 mV
o Inactivation gates close (Na channel inactivation)
o K channels open
o Repolarization begins
12-5 Action Potential
•
Step 4: Return to Normal Permeability
o K+ channels begin to close
 When membrane reaches normal resting potential (70 mV)
o K+ channels finish closing
 Membrane is hyperpolarized to 90 mV
 Membrane potential returns to resting level
 Action potential is over
12-5 Action Potential
•
The Refractory Period
o The time period:
 From beginning of action potential
 To return to resting state
 During which membrane will not respond normally to additional stimuli
12-5 Action Potential
•
Absolute Refractory Period
o Sodium channels open or inactivated
o No action potential possible
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•
Relative Refractory Period
o Membrane potential almost normal
o Very large stimulus can initiate action potential
12-5 Action Potential
•
Powering the Sodium–Potassium Exchange Pump
o To maintain concentration gradients of Na+ and K+ over time
 Requires energy (1 ATP for each 2 K+/3 Na+ exchange)
o Without ATP
 Neurons stop functioning
12-5 Action Potential
•
Propagation of Action Potentials
o Propagation
 Moves action potentials generated in axon hillock
 Along entire length of axon
o Two methods of propagating action potentials
1. Continuous propagation (unmyelinated axons)
2. Saltatory propagation (myelinated axons)
12-5 Action Potential
•
Continuous Propagation
o Of action potentials along an unmyelinated axon
o Affects one segment of axon at a time
o Steps in propagation
 Step 1: Action potential in segment 1
o Depolarizes membrane to 30 mV
o Local current
 Step 2: Depolarizes second segment to threshold
o Second segment develops action potential
12-5 Action Potential
•
Continuous Propagation
o Steps in propagation
 Step 3: First segment enters refractory period
 Step 4: Local current depolarizes next segment
o Cycle repeats
 Action potential travels in one direction (1 m/sec)
12-5 Action Potential
•
Saltatory Propagation
o Action potential along myelinated axon
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o
o
o
o
Faster and uses less energy than continuous propagation
Myelin insulates axon, prevents continuous propagation
Local current “jumps” from node to node
Depolarization occurs only at nodes
12-6 Axon Diameter and Speed
•
Axon Diameter and Propagation Speed
o Ion movement is related to cytoplasm concentration
o Axon diameter affects action potential speed
o The larger the diameter, the lower the resistance
12-6 Axon Diameter and Speed
•
Three Groups of Axons
1. Type A fibers
2. Type B fibers
3. Type C fibers
 These groups are classified by:
o Diameter
o Myelination
o Speed of action potentials
12-6 Axon Diameter and Speed
•
Type A Fibers
o Myelinated
o Large diameter
o High speed (140 m/sec)
o Carry rapid information to/from CNS
o For example, position, balance, touch, and motor impulses
12-6 Axon Diameter and Speed
•
Type B Fibers
o Myelinated
o Medium diameter
o Medium speed (18 m/sec)
o Carry intermediate signals
o For example, sensory information, peripheral effectors
12-6 Axon Diameter and Speed
•
Type C Fibers
o Unmyelinated
o Small diameter
o Slow speed (1 m/sec)
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o Carry slower information
o For example, involuntary muscle, gland controls
12-6 Axon Diameter and Speed
•
Information
o “Information” travels within the nervous system
 As propagated electrical signals (action potentials)
o The most important information (vision, balance, motor commands)
 Is carried by large-diameter, myelinated axons
12-7 Synapses
•
Synaptic Activity
o Action potentials (nerve impulses)
 Are transmitted from presynaptic neuron
 To postsynaptic neuron (or other postsynaptic cell)
 Across a synapse
12-7 Synapses
•
Two Types of Synapses
1. Electrical synapses
 Direct physical contact between cells
2. Chemical synapses
 Signal transmitted across a gap by chemical neurotransmitters
12-7 Synapses
•
Electrical Synapses
o Are locked together at gap junctions (connexons)
o Allow ions to pass between cells
o Produce continuous local current and action potential propagation
o Are found in areas of brain, eye, ciliary ganglia
12-7 Synapses
•
Chemical Synapses
o Are found in most synapses between neurons and all synapses between
neurons and other cells
o Cells not in direct contact
o Action potential may or may not be propagated to postsynaptic cell,
depending on:
 Amount of neurotransmitter released
 Sensitivity of postsynaptic cell
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12-7 Synapses
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Two Classes of Neurotransmitters
1. Excitatory neurotransmitters
 Cause depolarization of postsynaptic membranes
 Promote action potentials
2. Inhibitory neurotransmitters
 Cause hyperpolarization of postsynaptic membranes
 Suppress action potentials
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The Effect of a Neurotransmitter
o On a postsynaptic membrane
 Depends on the receptor
 Not on the neurotransmitter
o For example, acetylcholine (ACh)
 Usually promotes action potentials
 But inhibits cardiac neuromuscular junctions
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Cholinergic Synapses
o Any synapse that releases ACh at:
1. All neuromuscular junctions with skeletal muscle fibers
2. Many synapses in CNS
3. All neuron-to-neuron synapses in PNS
4. All neuromuscular and neuroglandular junctions of ANS
parasympathetic division
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Events at a Cholinergic Synapse
1. Action potential arrives, depolarizes synaptic terminal
2. Calcium ions enter synaptic terminal, trigger exocytosis of ACh
3. ACh binds to receptors, depolarizes postsynaptic membrane
4. ACh removed by AChE
 AChE breaks ACh into acetate and choline
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Synaptic Delay
o A synaptic delay of 0.2–0.5 msec occurs between:
 Arrival of action potential at synaptic terminal
 And effect on postsynaptic membrane
o Fewer synapses means faster response
o Reflexes may involve only one synapse
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12-7 Synapses
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Synaptic Fatigue
o Occurs when neurotransmitter cannot recycle fast enough to meet
demands of intense stimuli
o Synapse inactive until ACh is replenished
12-8 Neurotransmitters and Neuromodulators
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Other Neurotransmitters
o At least 50 neurotransmitters other than ACh, including:
 Biogenic amines
 Amino acids
 Neuropeptides
 Dissolved gases
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Important Neurotransmitters
o Other than acetylcholine
 Norepinephrine (NE)
 Dopamine
 Serotonin
 Gamma aminobutyric acid (GABA)
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Norepinephrine (NE)
o Released by adrenergic synapses
o Excitatory and depolarizing effect
o Widely distributed in brain and portions of ANS
Dopamine
o A CNS neurotransmitter
o May be excitatory or inhibitory
o Involved in Parkinson’s disease and cocaine use
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Serotonin
o A CNS neurotransmitter
o Affects attention and emotional states
Gamma Aminobutyric Acid (GABA)
o Inhibitory effect
o Functions in CNS
 Not well understood
12-8 Neurotransmitters and Neuromodulators
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Chemical Synapse
o The synaptic terminal releases a neurotransmitter that binds to the
postsynaptic plasma membrane
o Produces temporary, localized change in permeability or function of
postsynaptic cell
o Changes affect cell, depending on nature and number of stimulated
receptors
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Many Drugs
o Affect nervous system by stimulating receptors that respond to
neurotransmitters
o Can have complex effects on perception, motor control, and emotional
states
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Neuromodulators
o Other chemicals released by synaptic terminals
o Similar in function to neurotransmitters
o Characteristics of neuromodulators
 Effects are long term, slow to appear
 Responses involve multiple steps, intermediary compounds
 Affect presynaptic membrane, postsynaptic membrane, or both
 Released alone or with a neurotransmitter
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Neuropeptides
o Neuromodulators that bind to receptors and activate enzymes
Opioids
o Neuromodulators in the CNS
o Bind to the same receptors as opium or morphine
o Relieve pain
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Four Classes of Opioids
1. Endorphins
2. Enkephalins
3. Endomorphins
4. Dynorphins
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How Neurotransmitters and Neuromodulators Work
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o Direct effects on membrane channels
 For example, ACh, glycine, aspartate
o Indirect effects via G proteins
 For example, E, NE, dopamine, histamine, GABA
o Indirect effects via intracellular enzymes
 For example, lipid-soluble gases (NO, CO)
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Direct Effects
o Ionotropic effects
o Open/close gated ion channels
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Indirect Effects – G Proteins
o Work through second messengers
o Enzyme complex that binds GTP
o Link between neurotransmitter (first messenger) and second messenger
o Activate enzyme adenylyl cyclase
 Which produces second messenger cyclic-AMP (cAMP)
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Indirect Effects – Intracellular Receptors
o Lipid-soluble gases (NO, CO)
o Bind to enzymes in brain cells
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Information Processing
o At the simplest level (individual neurons)
 Many dendrites receive neurotransmitter messages simultaneously
 Some excitatory, some inhibitory
 Net effect on axon hillock determines if action potential is produced
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Postsynaptic Potentials
o Graded potentials developed in a postsynaptic cell
 In response to neurotransmitters
Two Types of Postsynaptic Potentials
1. Excitatory postsynaptic potential (EPSP)
 Graded depolarization of postsynaptic membrane
2. Inhibitory postsynaptic potential (IPSP)
 Graded hyperpolarization of postsynaptic membrane
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Inhibition
o A neuron that receives many IPSPs:
 Is inhibited from producing an action potential
 Because the stimulation needed to reach threshold is increased
Summation
o To trigger an action potential:
 One EPSP is not enough
 EPSPs (and IPSPs) combine through summation
1. Temporal summation
2. Spatial summation
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Temporal Summation
o Multiple times
o Rapid, repeated stimuli at one synapse
Spatial Summation
o Multiple locations
o Many stimuli, arrive at multiple synapses
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Facilitation
o A neuron becomes facilitated
 As EPSPs accumulate
 Raising membrane potential closer to threshold
 Until a small stimulus can trigger action potential
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Summation of EPSPs and IPSPs
o Neuromodulators and hormones
 Can change membrane sensitivity to neurotransmitters
 Shifting balance between EPSPs and IPSPs
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Axoaxonic Synapses
o Synapses between the axons of two neurons
o Presynaptic inhibition
 Action of an axoaxonic synapse at a synaptic terminal that decreases
the neurotransmitter released by presynaptic membrane
o Presynaptic facilitation
 Action of an axoaxonic synapse at a synaptic terminal that increases
the neurotransmitter released by presynaptic membrane
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12-9 Information Processing
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Frequency of Action Potentials
o Information received by a postsynaptic cell may be simply the frequency of
action potentials received
Rate of Generation of Action Potentials
o Frequency of action potentials depends on degree of depolarization above
threshold
o Holding membrane above threshold level
 Has same effect as a second, larger stimulus
 Reduces relative refractory period
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In the Nervous System
o A change in membrane potential that determines whether or not action
potentials are generated is the simplest form of information processing
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Summary
o Information is relayed in the form of action potentials
 In general, the degree of sensory stimulation or the strength of the
motor response is proportional to the frequency of action potentials
o The neurotransmitters released at a synapse may have either excitatory or
inhibitory effects
 The effect on the axon’s initial segment reflects a summation of the
stimuli that arrive at any moment
 The frequency of generation of action potentials is an indication of the
degree of sustained depolarization at the axon hillock
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Summary
o Neuromodulators
 Can alter either the rate of neurotransmitter release or the response of
a postsynaptic neuron to specific neurotransmitters
o Neurons
 May be facilitated or inhibited by extracellular chemicals other than
neurotransmitters or neuromodulators
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Summary
o The response of a postsynaptic neuron to the activation of a presynaptic
neuron can be altered by:
1. The presence of neuromodulators or other chemicals that cause
facilitation or inhibition at the synapse
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2. Activity under way at other synapses affecting the postsynaptic cell
3. Modification of the rate of neurotransmitter release through presynaptic
facilitation or presynaptic inhibition
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