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The master controlling and communicating system of the body
Functionso Sensory input – monitoring stimuli o Integration – interpretation of sensory inputo Motor output – response to stimuli
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Figure 11.1
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Central nervous system (CNS) o Brain and spinal cordo Integration and command center
Peripheral nervous system (PNS)o Paired spinal and cranial nerveso Carries messages to and from the spinal cord and brain
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Sensory (afferent) divisiono Sensory afferent fibers – carry impulses from skin,
skeletal muscles, and joints to the braino Visceral afferent fibers – transmit impulses from visceral
organs to the brain Motor (efferent) division
o Transmits impulses from the CNS to effector organs
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Somatic nervous systemo Conscious control of skeletal muscles
Autonomic nervous system (ANS)o Regulates smooth muscle, cardiac muscle, and glandso Divisions – sympathetic and parasympathetic
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The two principal cell types of the nervous system are:o Neurons – excitable cells that transmit electrical signalso Supporting cells – cells that surround and wrap neurons
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The supporting cells (neuroglia or glial cells):o Provide a supportive scaffolding for neuronso Segregate and insulate neuronso Guide young neurons to the proper connections o Promote health and growth
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Most abundant, versatile, and highly branched glial cells
They cling to neurons and their synaptic endings, and cover capillaries
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Functionally, they:o Support and brace neuronso Anchor neurons to their nutrient supplieso Guide migration of young neuronso Control the chemical environment
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Microglia – small, oval cells with spiny processeso Phagocytes that monitor the health of neurons
Ependymal cells – range in shape from squamous to columnaro They line the central cavities of the brain and spinal
column
Microglia Ependymal Cells
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Oligodendrocytes – branched cells that wrap CNS nerve fibers with myelin
Schwann cells (neurolemmocytes) – surround fibers of the PNS with myelin
Satellite cells surround neuron cell bodies with ganglia
Oligodendrocytes Schwann Cells Satellite Cells
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Structural units of the nervous systemo Composed of a body, axon, and dendriteso Long-lived, amitotic, and have a high metabolic rate
Their plasma membrane function in:o Electrical signaling o Cell-to-cell signaling during development
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Contains the nucleus and a nucleolus Is the major biosynthetic center Is the focal point for the outgrowth of neuronal
processes Has no centrioles (hence its amitotic nature) Has well-developed Nissl bodies (rough ER) Contains an axon hillock – cone-shaped area from
which axons arise
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Armlike extensions from the soma Called tracts in the CNS and nerves in the PNS There are two types: axons and dendrites
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Short, tapering, and diffusely branched processes They are the receptive, or input, regions of the
neuron Electrical signals are conveyed as graded
potentials (not action potentials)
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Slender processes of uniform diameter arising from the hillock
Long axons are called nerve fibers Usually there is only one unbranched axon per
neuron Rare branches, if present, are called axon
collaterals Axonal terminal – branched terminus of an axon
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Generate and transmit action potentials Secrete neurotransmitters from the axonal
terminals Movement along axons occurs in two ways
o Anterograde — toward axonal terminalo Retrograde — away from axonal terminal
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Whitish, fatty (protein-lipoid), segmented sheath around most long axons
It functions to:o Protect the axono Electrically insulate fibers from one anothero Increase the speed of nerve impulse transmission
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Formed by Schwann cells in the PNS A Schwann cell:
o Envelopes an axon in a trougho Encloses the axon with its plasma membraneo Has concentric layers of membrane that make up the
myelin sheath Neurilemma – remaining visible nucleus and
cytoplasm of a Schwann cell
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Figure 11.5a–c
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Gaps in the myelin sheath between adjacent Schwann cells
They are the sites where axon collaterals can emerge
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A Schwann cell surrounds nerve fibers but coiling does not take place
Schwann cells partially enclose 15 or more axons
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Both myelinated and unmyelinated fibers are present
Myelin sheaths are formed by oligodendrocytes Nodes of Ranvier are widely spaced There is no neurilemma
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White matter – dense collections of myelinated fibers
Gray matter – mostly soma and unmyelinated fibers
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Structural: o Multipolar — three or more processeso Bipolar — two processes (axon and dendrite)o Unipolar — single, short process
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Functional: o Sensory (afferent) — transmit impulses toward the CNSo Motor (efferent) — carry impulses away from the CNSo Interneurons (association neurons) — shuttle signals
through CNS pathways
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Table 11.1.1
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Table 11.1.2
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Table 11.1.3
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Neurons are highly irritable Action potentials, or nerve impulses, are:
o Electrical impulses carried along the length of axonso Always the same regardless of stimuluso The underlying functional feature of the nervous system
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Voltage (V) – measure of potential energy generated by separated charge
Potential difference – voltage measured between two points
Current (I) – the flow of electrical charge between two points
Resistance (R) – hindrance to charge flow Insulator – substance with high electrical
resistance Conductor – substance with low electrical
resistance
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Reflects the flow of ions rather than electrons There is a potential on either side of membranes
when:o The number of ions is different across the membraneo The membrane provides a resistance to ion flow
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Types of plasma membrane ion channels:o Passive, or leakage, channels – always openo Chemically gated channels – open with binding of a
specific neurotransmittero Voltage-gated channels – open and close in response to
membrane potentialo Mechanically gated channels – open and close in
response to physical deformation of receptors
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Example: Na+-K+ gated channel Closed when a neurotransmitter is not bound to
the extracellular receptoro Na+ cannot enter the cell and K+ cannot exit the cell
Open when a neurotransmitter is attached to the receptoro Na+ enters the cell and K+ exits the cell
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Example: Na+ channel Closed when the intracellular environment is
negative o Na+ cannot enter the cell
Open when the intracellular environment is positive o Na+ can enter the cell
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When gated channels are open: o Ions move quickly across the membrane o Movement is along their electrochemical gradientso An electrical current is createdo Voltage changes across the membrane
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Ions flow along their chemical gradient when they move from an area of high concentration to an area of low concentration
Ions flow along their electrical gradient when they move toward an area of opposite charge
Electrochemical gradient – the electrical and chemical gradients taken together
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The potential difference (–70 mV) across the membrane of a resting neuron
It is generated by different concentrations of Na+, K+, Cl, and protein anions (A)
Ionic differences are the consequence of:o Differential permeability of the neurilemma to Na+ and
K+
o Operation of the sodium-potassium pump
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Figure 11.8
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Used to integrate, send, and receive information Membrane potential changes are produced by:
o Changes in membrane permeability to ionso Alterations of ion concentrations across the membrane
Types of signals – graded potentials and action potentials
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Changes are caused by three eventso Depolarization – the inside of the membrane becomes
less negative o Repolarization – the membrane returns to its resting
membrane potentialo Hyperpolarization – the inside of the membrane
becomes more negative than the resting potential
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Short-lived, local changes in membrane potential Decrease in intensity with distance Magnitude varies directly with the strength of the
stimulus Sufficiently strong graded potentials can initiate
action potentials
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Figure 11.10
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Voltage changes are decremental Current is quickly dissipated due to the leaky
plasma membrane Only travel over short distances
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A brief reversal of membrane potential with a total amplitude of 100 mV
Action potentials are only generated by muscle cells and neurons
They do not decrease in strength over distance They are the principal means of neural
communication An action potential in the axon of a neuron is a
nerve impulse
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Na+ and K+ channels are closed Leakage accounts for small movements of
Na+ and K+
Each Na+ channel has two voltage-regulated gates o Activation gates –
closed in the resting state
o Inactivation gates – open in the resting state
Figure 11.12.1
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Na+ permeability increases; membrane potential reverses
Na+ gates are opened; K+ gates are closed Threshold – a critical level of depolarization
(-55 to -50 mV) At threshold,
depolarization becomes self-generating
Figure 11.12.2
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Sodium inactivation gates close Membrane permeability to Na+ declines to resting
levels As sodium gates close, voltage-sensitive K+ gates
open K+ exits the cell and
internal negativity of the resting neuron is restored
Figure 11.12.3
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Potassium gates remain open, causing an excessive efflux of K+
This efflux causes hyperpolarization of the membrane (undershoot)
The neuron is insensitive to stimulus and depolarization during this time
Figure 11.12.4
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Repolarization o Restores the resting electrical conditions of the neurono Does not restore the resting ionic conditions
Ionic redistribution back to resting conditions is restored by the sodium-potassium pump
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1 – resting state 2 – depolarization
phase 3 – repolarization
phase 4 –
hyperpolarization
Figure 11.12
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Na+ influx causes a patch of the axonal membrane to depolarize
Positive ions in the axoplasm move toward the polarized (negative) portion of the membrane
Sodium gates are shown as closing, open, or closed
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Figure 11.13a
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Ions of the extracellular fluid move toward the area of greatest negative charge
A current is created that depolarizes the adjacent membrane in a forward direction
The impulse propagates away from its point of origin
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Figure 11.13b
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The action potential moves away from the stimulus
Where sodium gates are closing, potassium gates are open and create a current flow
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Figure 11.13c
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Threshold – membrane is depolarized by 15 to 20 mV
Established by the total amount of current flowing through the membrane
Weak (subthreshold) stimuli are not relayed into action potentials
Strong (threshold) stimuli are relayed into action potentials
All-or-none phenomenon – action potentials either happen completely, or not at all
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All action potentials are alike and are independent of stimulus intensity
Strong stimuli can generate an action potential more often than weaker stimuli
The CNS determines stimulus intensity by the frequency of impulse transmission
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Figure 11.14
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Time from the opening of the Na+ activation gates until the closing of inactivation gates
The absolute refractory period:o Prevents the neuron from generating an action potentialo Ensures that each action potential is separateo Enforces one-way transmission of nerve impulses
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Figure 11.15
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The interval following the absolute refractory period when:o Sodium gates are closedo Potassium gates are openo Repolarization is occurring
The threshold level is elevated, allowing strong stimuli to increase the frequency of action potential events
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Conduction velocities vary widely among neurons Rate of impulse propagation is determined by:
o Axon diameter – the larger the diameter, the faster the impulse
o Presence of a myelin sheath – myelination dramatically increases impulse speed
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Current passes through a myelinated axon only at the nodes of Ranvier
Voltage-gated Na+ channels are concentrated at these nodes
Action potentials are triggered only at the nodes and jump from one node to the next
Much faster than conduction along unmyelinated axons
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Figure 11.16
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An autoimmune disease that mainly affects young adults
Symptoms: visual disturbances, weakness, loss of muscular control, and urinary incontinence
Nerve fibers are severed and myelin sheaths in the CNS become nonfunctional scleroses
Shunting and short-circuiting of nerve impulses occurs
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The advent of disease-modifying drugs including interferon beta-1a and -1b, Avonex, Betaseran, and Copazone:o Hold symptoms at bayo Reduce complicationso Reduce disability
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Nerve fibers are classified according to:o Diametero Degree of myelinationo Speed of conduction
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A junction that mediates information transfer from one neuron:o To another neurono To an effector cell
Presynaptic neuron – conducts impulses toward the synapse
Postsynaptic neuron – transmits impulses away from the synapse
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Axodendritic – synapses between the axon of one neuron and the dendrite of another
Axosomatic – synapses between the axon of one neuron and the soma of another
Other types of synapses include:o Axoaxonic (axon to axon)o Dendrodendritic (dendrite to dendrite)o Dendrosomatic (dendrites to soma)
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Electrical synapses:o Are less common than chemical synapseso Correspond to gap junctions found in other cell
typeso Are important in the CNS in:
• Arousal from sleep• Mental attention• Emotions and memory• Ion and water homeostasis
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Specialized for the release and reception of neurotransmitters
Typically composed of two parts: o Axonal terminal of the presynaptic neuron, which
contains synaptic vesicles o Receptor region on the dendrite(s) or soma of the
postsynaptic neuron
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Fluid-filled space separating the presynaptic and postsynaptic neurons
Prevents nerve impulses from directly passing from one neuron to the next
Transmission across the synaptic cleft: o Is a chemical event (as opposed to an electrical one)o Ensures unidirectional communication between neurons
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Nerve impulses reach the axonal terminal of the presynaptic neuron and open Ca2+ channels
Neurotransmitter is released into the synaptic cleft via exocytosis in response to synaptotagmin
Neurotransmitter crosses the synaptic cleft and binds to receptors on the postsynaptic neuron
Postsynaptic membrane permeability changes, causing an excitatory or inhibitory effect
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Synaptic vesiclescontaining neurotransmitter molecules
Axon of presynapticneuron
Synapticcleft
Ion channel(closed)
Ion channel (open)
Axon terminal of presynaptic neuron
PostsynapticmembraneMitochondrion
Ion channel closed
Ion channel open
Neurotransmitter
Receptor
Postsynapticmembrane
Degradedneurotransmitter
Na+Ca2+
1
2
34
5
Action
potential
Figure 11.18
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Neurotransmitter bound to a postsynaptic neuron: o Produces a continuous postsynaptic effecto Blocks reception of additional “messages” o Must be removed from its receptor
Removal of neurotransmitters occurs when they:o Are degraded by enzymeso Are reabsorbed by astrocytes or the presynaptic
terminals o Diffuse from the synaptic cleft
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Neurotransmitter must be released, diffuse across the synapse, and bind to receptors
Synaptic delay – time needed to do this (0.3-5.0 ms)
Synaptic delay is the rate-limiting step of neural transmission
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Neurotransmitter receptors mediate changes in membrane potential according to:o The amount of neurotransmitter releasedo The amount of time the neurotransmitter is bound to
receptors The two types of postsynaptic potentials are:
o EPSP – excitatory postsynaptic potentials o IPSP – inhibitory postsynaptic potentials
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EPSPs are graded potentials that can initiate an action potential in an axono Use only chemically gated channelso Na+ and K+ flow in opposite directions at the same time
Postsynaptic membranes do not generate action potentials
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Figure 11.19a
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Neurotransmitter binding to a receptor at inhibitory synapses: o Causes the membrane to become more permeable to
potassium and chloride ions o Leaves the charge on the inner surface negativeo Reduces the postsynaptic neuron’s ability to produce an
action potential
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Figure 11.19b
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A single EPSP cannot induce an action potential EPSPs must summate temporally or spatially to
induce an action potential Temporal summation – presynaptic neurons
transmit impulses in rapid-fire order
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Spatial summation – postsynaptic neuron is stimulated by a large number of terminals at the same time
IPSPs can also summate with EPSPs, canceling each other out
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Figure 11.20
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Chemicals used for neuronal communication with the body and the brain
50 different neurotransmitters have been identified
Classified chemically and functionally
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Acetylcholine (ACh) Biogenic amines Amino acids Peptides Novel messengers: ATP and dissolved gases NO
and CO
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First neurotransmitter identified, and best understood
Released at the neuromuscular junction Synthesized and enclosed in synaptic vesicles
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Degraded by the enzyme acetylcholinesterase (AChE)
Released by:o All neurons that stimulate skeletal muscleo Some neurons in the autonomic nervous system
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Include:o Catecholamines – dopamine, norepinephrine (NE), and
epinephrineo Indolamines – serotonin and histamine
Broadly distributed in the brain Play roles in emotional behaviors and our
biological clock
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Enzymes present in the cell determine length of biosynthetic pathway
Norepinephrine and dopamine are synthesized in axonal terminals
Epinephrine is released by the adrenal medulla
Figure 11.21
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Include:o GABA – Gamma ()-aminobutyric acid o Glycineo Aspartateo Glutamate
Found only in the CNS
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Include:o Substance P – mediator of pain signalso Beta endorphin, dynorphin, and enkephalins
Act as natural opiates; reduce pain perception Bind to the same receptors as opiates and
morphine Gut-brain peptides – somatostatin, and
cholecystokinin
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ATPo Is found in both the CNS and PNSo Produces excitatory or inhibitory responses depending
on receptor typeo Induces Ca2+ wave propagation in astrocyteso Provokes pain sensation
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Nitric oxide (NO) o Activates the intracellular receptor guanylyl cyclaseo Is involved in learning and memory
Carbon monoxide (CO) is a main regulator of cGMP in the brain
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Two classifications: excitatory and inhibitoryo Excitatory neurotransmitters cause depolarizations
(e.g., glutamate)o Inhibitory neurotransmitters cause hyperpolarizations
(e.g., GABA and glycine)
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Some neurotransmitters have both excitatory and inhibitory effects o Determined by the receptor type of the postsynaptic
neuron o Example: acetylcholine
• Excitatory at neuromuscular junctions with skeletal muscle• Inhibitory in cardiac muscle
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Direct: neurotransmitters that open ion channelso Promote rapid responses o Examples: ACh and amino acids
Indirect: neurotransmitters that act through second messengerso Promote long-lasting effectso Examples: biogenic amines, peptides, and dissolved
gases
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Composed of integral membrane protein- a protein permanently attached to the cell membrane
Mediate direct neurotransmitter action Action is immediate, brief, simple, and highly
localized Ligand binds the receptor, and ions enter the
cells Excitatory receptors depolarize membranes Inhibitory receptors hyperpolarize membranes
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Figure 11.22a
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Responses are indirect, slow, complex, prolonged, and often diffuse
These receptors are transmembrane protein complexes
Examples: muscarinic ACh receptors, neuropeptides, and those that bind biogenic amines
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G protein-linked receptors activate intracellular second messengers including Ca2+, cGMP, diacylglycerol, as well as cAMP
Second messengers:o Open or close ion channelso Activate kinase enzymeso Phosphorylate channel proteins o Activate genes and induce protein synthesis
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Functional groups of neurons that:o Integrate incoming informationo Forward the processed information to its appropriate
destination
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Simple neuronal poolo Input fiber – presynaptic fibero Discharge zone – neurons most closely associated with
the incoming fibero Facilitated zone – neurons farther away from incoming
fiber
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Figure 11.23
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Divergent – one incoming fiber stimulates ever increasing number of fibers, often amplifying circuits
Figure 11.24a, b
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Convergent – opposite of divergent circuits, resulting in either strong stimulation or inhibition
Figure 11.24c, d
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Reverberating – chain of neurons containing collateral synapses with previous neurons in the chain
Figure 11.24e
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Parallel after-discharge – incoming neurons stimulate several neurons in parallel arrays
Figure 11.24f
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Serial Processingo Input travels along one pathway to a specific destinationo Works in an all-or-none mannero Example: spinal reflexes
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Parallel Processingo Input travels along several pathwayso Pathways are integrated in different CNS systemso One stimulus promotes numerous responses
Example: a smell may remind one of the odor and associated experiences
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The nervous system originates from the neural tube and neural crest
The neural tube becomes the CNS There is a three-phase process of differentiation:
o Proliferation of cells needed for developmento Migration – cells become amitotic and move externallyo Differentiation into neuroblasts
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Guided by:o Scaffold laid down by older neuronso Orienting glial fiberso Release of nerve growth factor by astrocyteso Neurotropins released by other neuronso Repulsion guiding moleculeso Attractants released by target cells
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N-CAM – nerve cell adhesion molecule Important in establishing neural pathways Without N-CAM, neural function is impaired Found in the membrane of the growth cone