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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 bodyo Neural tissue contains two kinds of cells
An Introduction to the Nervous System• Organs of the Nervous System
o Brain and spinal cordo 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 vesselso 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 CNSo 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 braino Spinal nerves – attach to spinal cord
12-1 Divisions of the Nervous System• Functional Divisions of the PNS
o Are chemical messengerso Are released at presynaptic membraneo Affect receptors of postsynaptic membrane o Are broken down by enzymeso 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
12-4 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,
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 3:2o 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 potentialo They are either passive or active
o Open in presence of specific chemicals (e.g., ACh) at a binding siteo Found on neuron cell body and dendrites
12-4 Membrane Potential• Voltage-gated Channels
o Respond to changes in membrane potentialo Have activation gates (open) and inactivation gates (close)o Characteristic of excitable membraneo Found in neural axons, skeletal muscle sarcolemma, cardiac muscle
The plasma membrane is selectively permeable • Membrane Potential
o Changes with plasma membrane permeabilityo In response to chemical or physical stimuli
12-4 Membrane Potential• Graded Potentials
o Also called local potentialso Changes in membrane potential
That cannot spread far from site of stimulationo Any stimulus that opens a gated channel
Produces a graded potential
12-4 Membrane Potential• Graded Potentials
o The resting state Opening sodium channel produces graded potential
o Resting membrane exposed to chemicalo Sodium channel openso Sodium ions enter the cello Membrane potential riseso 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 characteristics1. The membrane potential is most changed at the site of stimulation, and
the effect decreases with distance2. The effect spreads passively, due to local currents
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 changeo 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 characteristics4. The stronger the stimulus, the greater the change in the membrane
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 functionso For example, exocytosis of glandular secretions
At motor end plateo Release ACh into synaptic cleft
12-5 Action Potential• Action Potentials
o Propagated changes in membrane potentialo Affect an entire excitable membraneo Link graded potentials at cell body with motor end plate actions
o All-or-none principle If a stimulus exceeds threshold amount
o The action potential is the sameo No matter how large the stimulus
Action potential is either triggered, or not
12-5 Action Potential• Four Steps in the Generation of Action Potentials
o Step 1: Depolarization to thresholdo Step 2: Activation of Na channelso Step 3: Inactivation of Na channels and activation of K channelso Step 4: Return to normal permeability
12-5 Action Potential• Step 1: Depolarization to Threshold• Step 2: Activation of Na Channels
o Rapid depolarizationo Na+ ions rush into cytoplasmo 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 mVo Inactivation gates close (Na channel inactivation)o K channels openo 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)
Step 1: Action potential in segment 1 o Depolarizes membrane to 30 mVo 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 axono Faster and uses less energy than continuous propagationo Myelin insulates axon, prevents continuous propagationo Local current “jumps” from node to nodeo Depolarization occurs only at nodes
12-6 Axon Diameter and Speed• Axon Diameter and Propagation Speed
o Ion movement is related to cytoplasm concentrationo Axon diameter affects action potential speedo The larger the diameter, the lower the resistance
12-6 Axon Diameter and Speed• Three Groups of Axons
1. Type A fibers2. Type B fibers3. Type C fibers
These groups are classified by:o Diametero Myelinationo Speed of action potentials
o Are locked together at gap junctions (connexons)o Allow ions to pass between cellso Produce continuous local current and action potential propagationo 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 contacto Action potential may or may not be propagated to postsynaptic cell,
depending on: Amount of neurotransmitter released Sensitivity of postsynaptic cell
12-7 Synapses• 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
12-7 Synapses• 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
12-7 Synapses• Cholinergic Synapses
o Any synapse that releases ACh at:1. All neuromuscular junctions with skeletal muscle fibers2. Many synapses in CNS3. All neuron-to-neuron synapses in PNS4. All neuromuscular and neuroglandular junctions of ANS
1. Action potential arrives, depolarizes synaptic terminal2. Calcium ions enter synaptic terminal, trigger exocytosis of ACh3. ACh binds to receptors, depolarizes postsynaptic membrane4. ACh removed by AChE
AChE breaks ACh into acetate and choline
12-7 Synapses• 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 responseo Reflexes may involve only one synapse
12-7 Synapses• 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• Other Neurotransmitters
o At least 50 neurotransmitters other than ACh, including: Biogenic amines Amino acids Neuropeptides Dissolved gases
12-8 Neurotransmitters and Neuromodulators• Important Neurotransmitters
o Other than acetylcholine Norepinephrine (NE) Dopamine Serotonin Gamma aminobutyric acid (GABA)
Responses involve multiple steps, intermediary compounds Affect presynaptic membrane, postsynaptic membrane, or both Released alone or with a neurotransmitter
12-8 Neurotransmitters and Neuromodulators• Neuropeptides
o Neuromodulators that bind to receptors and activate enzymes• Opioids
o Neuromodulators in the CNSo Bind to the same receptors as opium or morphineo Relieve pain
12-8 Neurotransmitters and Neuromodulators• Four Classes of Opioids
12-8 Neurotransmitters and Neuromodulators• How Neurotransmitters and Neuromodulators Work
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)
12-8 Neurotransmitters and Neuromodulators• Direct Effects
o Ionotropic effectso Open/close gated ion channels
12-8 Neurotransmitters and Neuromodulators• Indirect Effects – G Proteins
o Work through second messengerso Enzyme complex that binds GTPo Link between neurotransmitter (first messenger) and second messengero Activate enzyme adenylyl cyclase
12-8 Neurotransmitters and Neuromodulators• Indirect Effects – Intracellular Receptors
o Lipid-soluble gases (NO, CO)o Bind to enzymes in brain cells
12-9 Information Processing• 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
12-9 Information Processing• Postsynaptic Potentials
o Graded potentials developed in a postsynaptic cell In response to neurotransmitters
• Two Types of Postsynaptic Potentials1. Excitatory postsynaptic potential (EPSP)
Graded depolarization of postsynaptic membrane2. Inhibitory postsynaptic potential (IPSP)
Graded hyperpolarization of postsynaptic membrane
12-9 Information Processing• Inhibition
o A neuron that receives many IPSPs: Is inhibited from producing an action potential Because the stimulation needed to reach threshold is increased
• Summationo To trigger an action potential:
One EPSP is not enough EPSPs (and IPSPs) combine through summation
1. Temporal summation2. Spatial summation
12-9 Information Processing• Temporal Summation
o Multiple timeso Rapid, repeated stimuli at one synapse
• Spatial Summationo Multiple locationso Many stimuli, arrive at multiple synapses
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 potentialso 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
12-9 Information Processing• Summary
o Neuromodulators Can alter either the rate of neurotransmitter release or the response of
a postsynaptic neuron to specific neurotransmitterso Neurons
May be facilitated or inhibited by extracellular chemicals other than neurotransmitters or neuromodulators
12-9 Information Processing• 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 synapse2. Activity under way at other synapses affecting the postsynaptic cell3. Modification of the rate of neurotransmitter release through presynaptic