CH11(12) Functions of the Nervous System 1._________ input • Information gathered by sensory receptors about internal and _________ changes 2.Integration • _________ of sensory input 3.Motor output • Activation of _________ organs (_________ and _________) produces a response Divisions of the Nervous System • _________ nervous system (CNS) • Brain and spinal cord • Integration and command center • _________ nervous system (PNS) • Paired _________ and _________ nerves carry messages to and from the CNS Peripheral Nervous System (PNS) • Two functional divisions 1. Sensory (_________) division • Somatic afferent fibers—convey impulses from skin, skeletal muscles, and joints • _________ afferent fibers—convey impulses from visceral organs 2. Motor (_________) division • Transmits impulses from the CNS to effector organs Motor Division of PNS 1.Somatic (_________) nervous system • _________ control of skeletal muscles Motor Division of PNS 2.Autonomic (_________) nervous system (_________) • _________ motor nerve fibers • Regulates _________ muscle, cardiac muscle, and glands • _________ functional subdivisions • _________ • Parasympathetic Histology of Nervous Tissue • Two principal cell types 1. _________ —excitable cells that transmit electrical signals Histology of Nervous Tissue 2.Neuroglia (glial cells)—supporting cells: • • • • • • _________ (CNS) Microglia (CNS) _________ cells (CNS) Oligodendrocytes (CNS) _________ cells (PNS) Schwann cells (PNS) Astrocytes • Most abundant, _________, and highly branched glial cells • Cling to _________, synaptic endings, and _________ • Support and _________ neurons Astrocytes • Help determine capillary _________ • Guide _________ of young neurons • Control the _________ environment • Participate in information processing in the _________ Microglia • Small, _________ cells with thorny processes • _________ toward injured neurons • Phagocytize _________ and neuronal _________ Ependymal Cells • Range in shape from _________ to _________ • May be ciliated • Line the central cavities of the brain and spinal column • Separate the CNS _________ fluid from the cerebrospinal fluid in the cavities Oligodendrocytes • _________ cells • Processes wrap _________ nerve fibers, forming insulating myelin sheaths Satellite Cells and Schwann Cells • _________ cells • Surround neuron cell bodies in the PNS • _________ cells (_________) • Surround peripheral nerve fibers and form myelin sheaths • Vital to _________ of damaged _________ nerve fibers Neurons (Nerve Cells) • Special characteristics: • Long-lived (_________ years or more) • _________ —with few exceptions • _________ metabolic rate—depends on continuous supply of oxygen and glucose • Plasma membrane functions in: • _________ signaling • Cell-to-cell interactions during development Cell Body (Perikaryon or Soma) • _________ center of a neuron • Spherical nucleus with nucleolus • Well-developed _________ _________ • Rough ER called _________ bodies (chromatophilic substance) Cell Body (Perikaryon or Soma) • Network of _________ (neurofilaments) • _________ hillock—cone-shaped area from which axon arises • Clusters of cell bodies are called nuclei in the CNS, ganglia in the _________ Processes • _________ and _________ • Bundles of processes are called • Tracts in the _________ • Nerves in the _________ Dendrites • _________, tapering, and diffusely branched • Receptive (input) region of a _________ • Convey _________ signals toward the cell body as graded potentials The Axon • _________ axon per cell arising from the axon hillock • Long axons (nerve fibers) • Occasional branches (_________ collaterals) The Axon • _________ terminal branches (_________) • Knoblike axon terminals (synaptic knobs or boutons) • _________ region of neuron • Release neurotransmitters to _________ or _________ other cells Axons: Function • _________ region of a neuron • Generates and transmits nerve impulses (_________ potentials) away from the cell body Axons: Function • Molecules and organelles are moved along _________ by _________ molecules in two directions: • _________ —toward axonal terminal • Examples: mitochondria, membrane components, enzymes • _________ —toward the cell body • Examples: organelles to be degraded, signal molecules, viruses, and bacterial toxins Myelin Sheath • Segmented _________ -lipoid sheath around most long or large-diameter axons • It functions to: • Protect and _________ insulate the axon • _________ speed of nerve impulse transmission Myelin Sheaths in the PNS • _________ cells wraps many times around the axon • Myelin sheath—concentric layers of Schwann cell membrane • _________ —peripheral bulge of Schwann cell cytoplasm Myelin Sheaths in the PNS • Nodes of _________ • Myelin sheath gaps between adjacent _________ cells • Sites where _________ collaterals can emerge Unmyelinated Axons • Thin nerve fibers are _________ • One Schwann cell may incompletely enclose 15 or more _________ axons Myelin Sheaths in the CNS • Formed by processes of oligodendrocytes, not the whole cells • Nodes of _________ are present • No _________ • _________ fibers are unmyelinated White Matter and Gray Matter • _________ matter • Dense collections of myelinated fibers • _________ matter • Mostly neuron cell bodies and unmyelinated fibers Structural Classification of Neurons • Three types: 1. _________ —1 axon and several dendrites • Most abundant • Motor neurons and _________ 2. _________ —1 axon and 1 dendrite • Rare, e.g., retinal neurons Structural Classification of Neurons 3. Unipolar (_________)—single, short process that has two branches: • _________ process—more distal branch, often associated with a sensory receptor • _________ process—branch entering the _________ Functional Classification of Neurons • Three types: 1. _________ (afferent) • Transmit impulses from sensory receptors toward the CNS 2. _________ (efferent) • Carry impulses from the CNS to effectors Functional Classification of Neurons 3. _________ (association neurons) • Shuttle signals through CNS pathways; most are entirely within the CNS Neuron Function • Neurons are _________ irritable • Respond to adequate _________ by generating an action potential (nerve impulse) • Impulse is always the _________ regardless of stimulus Principles of Electricity • _________ charges attract each other • Energy is required to separate _________ charges across a membrane • Energy is _________ when the charges move _________ one another • If opposite charges are _________, the system has potential energy Definitions • _________ (V): measure of potential energy generated by separated charge • _________ difference: voltage measured between two points • _________ (I): the flow of electrical charge (ions) between two points Definitions • _________ (R): hindrance to charge flow (provided by the plasma membrane) • _________: substance with high electrical resistance • _________: substance with low electrical resistance Role of Membrane Ion Channels • _________ serve as membrane ion channels • Two main types of ion channels 1. Leakage (_________) channels—always _________ Role of Membrane Ion Channels 2. _________ channels (three types): • Chemically gated (_________ -gated) channels—open with binding of a specific neurotransmitter • _________ -gated channels—open and close in response to changes in membrane potential • _________ gated channels—open and close in response to physical deformation of receptors Gated Channels • When gated channels are _________: • Ions diffuse quickly across the membrane along their _________ gradients • Along _________ concentration gradients from higher concentration to _________ concentration • Along electrical gradients toward _________ electrical charge • Ion flow creates an electrical current and _________ changes across the membrane Resting Membrane Potential (Vr) • Potential difference across the membrane of a _________ cell • Approximately –70 mV in neurons (_________ side of membrane is negatively charged relative to outside) • Generated by: • Differences in ionic makeup of _________ and _________ • Differential permeability of the plasma membrane Resting Membrane Potential • Differences in ionic makeup • _________ has lower concentration of Na+ and Cl– than ECF • _________ has higher concentration of K+ and negatively charged proteins (A–) than ECF Resting Membrane Potential • Differential permeability of membrane • • • • _________ to A– Slightly permeable to Na+ (through _________ channels) 75 times _________ permeable to K+ (more leakage channels) _________ permeable to Cl– Resting Membrane Potential • _________ interior of the cell is due to much _________ diffusion of K+ out of the cell than Na+ diffusion into the cell • _________ -_________ pump stabilizes the resting membrane potential by maintaining the concentration gradients for Na + and K+ Membrane Potentials That Act as Signals • Membrane _________ changes when: • Concentrations of ions across the membrane change • _________ of membrane to ions changes • Changes in membrane potential are _________ used to _________, integrate and _________ information Membrane Potentials That Act as Signals • Two types of signals • _________ potentials • Incoming short-distance signals • _________ potentials • Long-distance signals of axons Changes in Membrane Potential • Depolarization • A _________ in membrane potential (toward zero) • Inside of the membrane becomes less _________ than the resting potential • _________ the probability of producing a nerve impulse Changes in Membrane Potential • Hyperpolarization • An increase in membrane potential (away from _________) • Inside of the membrane becomes more _________ than the resting potential • _________ the probability of producing a nerve impulse Graded Potentials • _________ -lived, localized changes in membrane potential • _________ or hyperpolarizations • Graded potential spreads as local currents change the membrane potential of adjacent regions Graded Potentials • Occur when a stimulus causes gated ion channels to open • E.g., _________ potentials, _________ potentials, _________ potentials • _________ varies directly (graded) with stimulus strength • _________ in magnitude with distance as ions flow and diffuse through leakage channels • Short-distance signals Action Potential (AP) • Brief _________ of membrane potential with a total amplitude of ~100 mV • Occurs in _________ cells and _________ of neurons • Does not decrease in magnitude over distance • Principal means of _________ -distance _________ communication Generation of an Action Potential • _________ state • Only leakage channels for Na+ and K+ are open • All gated Na+ and K+ channels are closed Properties of Gated Channels • Properties of _________ channels • Each _________ + channel has two voltage-sensitive gates • _________ gates • Closed at rest; open with depolarization • _________ gates • Open at rest; block channel once it is open Properties of Gated Channels • Each _________ + channel has one voltage-sensitive gate • _________ at rest • Opens _________ with depolarization Depolarizing Phase • _________ local currents open voltage-gated Na+ channels • Na+ _________ causes more depolarization • At _________ (–55 to –50 mV) _________ feedback leads to opening of all Na+ channels, and a reversal of membrane polarity to +30mV (spike of action potential) Repolarizing Phase • _________ phase • • • • _________ + channel slow inactivation gates close Membrane permeability to Na+ _________ to resting levels Slow _________ -sensitive K+ gates open K+ exits the cell and internal _________ is restored Hyperpolarization • Hyperpolarization • Some K+ channels remain _________, allowing _________ K+ efflux • This causes after-_________ of the membrane (undershoot) Role of the Sodium-Potassium Pump • Repolarization • _________ the resting electrical conditions of the neuron • Does not restore the _________ ionic conditions • Ionic redistribution back to resting conditions is restored by the thousands of _________ -_________ pumps Propagation of an Action Potential • Na+ influx causes a patch of the axonal membrane to depolarize • _________ currents occur • Na+ channels toward the point of _________ are inactivated and not affected by the local currents Propagation of an Action Potential • _________ currents affect adjacent areas in the forward direction • _________ opens voltage-gated channels and triggers an AP • Repolarization wave follows the depolarization wave • (Fig. 11.12 shows the propagation process in unmyelinated axons.) Threshold • At threshold: • • • • Membrane is _________ by 15 to 20 mV Na+ permeability increases Na _________ exceeds K+ efflux The _________ feedback cycle begins Threshold • _________ stimulus—weak local depolarization that does not reach threshold • _________ stimulus—strong enough to push the membrane potential toward and beyond threshold • AP is an _________ phenomenon—action potentials either happen completely, or not at all Coding for Stimulus Intensity • All _________ potentials are alike and are independent of stimulus intensity • How does the _________ tell the difference between a weak stimulus and a strong one? • Strong stimuli can generate action potentials _________ often than weaker stimuli • The CNS determines stimulus intensity by the _________ of impulses Absolute Refractory Period • Time from the opening of the Na+ channels until the _________ of the channels • Ensures that each AP is an all-or-none event • Enforces _________ transmission of nerve impulses Relative Refractory Period • Follows the _________ refractory period • Most Na+ channels have returned to their resting state • Some K+ _________ are still open • _________ is occurring • _________ for AP generation is elevated • Exceptionally _________ stimulus may generate an AP Conduction Velocity • _________ velocities of neurons vary widely • Effect of axon _________ • Larger diameter fibers have less _________ to local current flow and have faster impulse conduction • Effect of _________ • Continuous conduction in _________ axons is slower than saltatory conduction in myelinated axons Conduction Velocity • Effects of _________ • Myelin sheaths insulate and prevent leakage of charge • _________ conduction in _________ axons is about 30 times faster • Voltage-gated Na+ channels are located at the nodes • APs appear to jump rapidly from node to node Multiple Sclerosis (MS) • An _________ disease that mainly affects young adults • Symptoms: visual disturbances, _________, loss of muscular control, speech disturbances, and urinary incontinence • Myelin sheaths in the _________ become nonfunctional scleroses • Shunting and short-circuiting of nerve impulses occurs • Impulse conduction _________ and eventually _________ Multiple Sclerosis: Treatment • Some immune system–modifying drugs, including interferons and Copazone: • Hold symptoms at bay • Reduce complications • Reduce disability Nerve Fiber Classification • Nerve fibers are classified according to: • Diameter • Degree of _________ • Speed of _________ Nerve Fiber Classification • Group _________ fibers • Large diameter, _________ somatic sensory and motor fibers • Group B fibers • _________ diameter, lightly myelinated ANS fibers • Group C fibers • _________ diameter, unmyelinated ANS fibers The Synapse • A junction that mediates information transfer from one neuron: • To another _________, or • To an _________ cell The Synapse • _________ neuron—conducts impulses toward the synapse • _________ neuron—transmits impulses away from the synapse Types of Synapses • _________ —between the axon of one neuron and the dendrite of another • _________ —between the axon of one neuron and the soma of another • Less common types: • _________ (axon to axon) • _________ (dendrite to dendrite) • _________ (dendrite to soma) Electrical Synapses • _________ common than chemical synapses • Neurons are electrically coupled (joined by _________ junctions) • Communication is very _________, and may be unidirectional or bidirectional • Are important in: • _________ nervous tissue • Some brain regions Chemical Synapses • Specialized for the release and reception of _________ • Typically composed of two parts • Axon terminal of the _________ neuron, which contains synaptic vesicles • _________ region on the postsynaptic neuron Synaptic Cleft • Fluid-filled space separating the _________ and _________ neurons • Prevents nerve impulses from directly passing from one neuron to the next Synaptic Cleft • Transmission across the _________ cleft: • Is a chemical event (as opposed to an electrical one) • Involves _________, _________, and binding of neurotransmitters • Ensures unidirectional communication _________ neurons Information Transfer • AP arrives at _________ terminal of the _________ neuron and opens voltage-gated Ca2+ channels • _________ protein binds Ca2+ and promotes fusion of synaptic vesicles with axon membrane • _________ of neurotransmitter occurs Information Transfer • _________ diffuses and binds to receptors (often chemically gated ion channels) on the postsynaptic neuron • _________ channels are opened, causing an excitatory or inhibitory event (graded potential) Termination of Neurotransmitter Effects • Within a few milliseconds, the neurotransmitter effect is terminated • Degradation by _________ • Reuptake by _________ or axon terminal • _________ away from the synaptic cleft Synaptic Delay • _________ must be released, diffuse across the synapse, and bind to receptors • _________ delay—time needed to do this (0.3–5.0 ms) • Synaptic delay is the rate-limiting step of _________ transmission Postsynaptic Potentials • Graded potentials • _________ determined by: • Amount of _________ released • Time the neurotransmitter is in the area • Types of _________ potentials 1. _________ —excitatory postsynaptic potentials 2. _________ —inhibitory postsynaptic potentials Excitatory Synapses and EPSPs • _________ binds to and opens chemically gated channels that allow simultaneous flow of Na+ and K+ in opposite directions • Na+ influx is greater that K+ _________, causing a net depolarization • EPSP helps trigger AP at axon _________ if EPSP is of threshold strength and opens the voltage-gated channels Inhibitory Synapses and IPSPs • Neurotransmitter binds to and opens channels for K+ or Cl– • Causes a _________ (the inner surface of membrane becomes more negative) • Reduces the _________ neuron’s ability to produce an action potential Integration: Summation • A single _________ cannot induce an action potential • _________ can summate to reach threshold • _________ can also summate with _________, canceling each other out Integration: Summation • _________ summation • One or more presynaptic neurons transmit impulses in rapid-fire order • _________ summation • Postsynaptic neuron is stimulated by a large number of terminals at the same time Integration: Synaptic Potentiation • Repeated use _________ the efficiency of _________ • Ca2+ concentration increases in _________ terminal and ostsynaptic neuron • Brief _________ -frequency stimulation partially depolarizes the postsynaptic neuron • _________ gated channels (NMDA receptors) allow Ca2+ entry • Ca2+ activates kinase enzymes that promote more effective responses to subsequent stimuli Integration: Presynaptic Inhibition • Release of excitatory _________ by one neuron may be inhibited by the activity of another neuron via an axoaxonic synapse • _________ neurotransmitter is released and _________ EPSPs are formed Neurotransmitters • Most neurons make _________ or more neurotransmitters, which are released at different stimulation frequencies • _________ or more neurotransmitters have been identified • Classified by chemical structure and by function Chemical Classes of Neurotransmitters • _________ (Ach) • Released at neuromuscular junctions and some ANS neurons • Synthesized by enzyme choline acetyltransferase • _________ by the enzyme _________ (AChE) Chemical Classes of Neurotransmitters • _________ amines include: • Catecholamines • _________, norepinephrine (NE), and _________ • Indolamines • Serotonin and histamine • _________ distributed in the brain • Play roles in _________ behaviors and the biological clock Chemical Classes of Neurotransmitters • Amino acids include: • _________ —Gamma ()-aminobutyric acid • Glycine • _________ • Glutamate Chemical Classes of Neurotransmitters • Peptides (_________) include: • _________ P • Mediator of pain signals • Endorphins • Act as natural opiates; reduce pain perception • _________ -brain peptides • Somatostatin and _________ Chemical Classes of Neurotransmitters • _________ such as ATP: • Act in both the _________ and PNS • Produce fast or slow responses • Induce Ca2+ influx in _________ • Provoke pain sensation Chemical Classes of Neurotransmitters • _________ and lipids • _________ oxide (NO) • Synthesized on demand • Activates the intracellular receptor guanylyl cyclase to cyclic _________ • Involved in learning and memory • _________ _________ (CO) is a regulator of _________ in the brain Chemical Classes of Neurotransmitters • _________ and lipids • Endocannabinoids • _________ soluble; synthesized on demand from membrane lipids • Bind with _________ protein–coupled receptors in the brain • Involved in learning and memory Functional Classification of Neurotransmitters • Neurotransmitter effects may be excitatory (depolarizing) and/or inhibitory (hyperpolarizing) • • • • _________ by the receptor type of the postsynaptic neuron GABA and glycine are usually inhibitory _________ is usually excitatory Acetylcholine • _________ at neuromuscular junctions in skeletal muscle • Inhibitory in cardiac muscle Neurotransmitter Actions • Direct action • _________ binds to channel-linked receptor and opens ion channels • Promotes _________ responses • Examples: _________ and _________ _________ Neurotransmitter Actions • _________ action • Neurotransmitter binds to a _________ protein-linked receptor and acts through an _________ second messenger • Promotes long-lasting effects • Examples: biogenic amines, neuropeptides, and dissolved gases Neurotransmitter Receptors • Types 1. Channel-linked receptors 2. G protein-linked receptors Channel-Linked (Ionotropic) Receptors • _________ -gated ion channels • Action is immediate and brief • _________ receptors are channels for small cations • Na+ influx contributes most to depolarization • Inhibitory receptors allow Cl– _________ or K+ efflux that causes _________ G Protein-Linked (Metabotropic) Receptors • _________ protein complexes • Responses are indirect, _________, _________, and often prolonged and widespread • Examples: muscarinic ACh receptors and those that bind biogenic amines and neuropeptides G Protein-Linked Receptors: Mechanism • Neurotransmitter binds to _________ protein–linked receptor • G protein is activated • _________ G protein controls production of second messengers, e.g., _________ AMP, cyclic _________, diacylglycerol or Ca2+ G Protein-Linked Receptors: Mechanism • Second messengers • • • • _________ or close ion channels Activate kinase enzymes _________ channel proteins Activate genes and induce protein synthesis Neural Integration: Neuronal Pools • Functional groups of _________ that: • Integrate _________ information • Forward the processed information to other destinations Neural Integration: Neuronal Pools • Simple neuronal pool • Single _________ fiber branches and synapses with several neurons in the pool • _________ zone—neurons most closely associated with the incoming fiber • _________ zone—neurons farther away from incoming fiber Types of Circuits in Neuronal Pools • _________ circuit • One incoming fiber stimulates an ever-increasing number of fibers, often amplifying circuits • May affect a _________ pathway or several • Common in both sensory and motor systems Types of Circuits in Neuronal Pools • _________ circuit • Opposite of diverging circuits, resulting in either strong stimulation or inhibition • Also common in sensory and motor systems Types of Circuits in Neuronal Pools • Reverberating (_________) circuit • Chain of neurons containing collateral synapses with previous neurons in the chain Types of Circuits in Neuronal Pools • _________ after-discharge circuit • Incoming fiber stimulates _________ neurons in parallel arrays to stimulate a common output cell Patterns of Neural Processing • _________ processing • Input travels along _________ pathway to a specific destination • Works in an all-or-none manner to produce a specific response Patterns of Neural Processing • _________ processing • Example: _________ —rapid, automatic responses to stimuli that always cause the same response • Reflex arcs (_________) have _________ essential components: receptor, sensory neuron, CNS integration center, motor neuron, and effector Patterns of Neural Processing • Parallel processing • Input travels along _________ pathways • _________ stimulus promotes numerous responses • Important for _________ -level mental functioning • Example: a smell may remind one of the odor and associated experiences Developmental Aspects of Neurons • The nervous system _________ from the neural tube and neural crest formed from ectoderm • The _________ tube becomes the CNS • _________ cells of the neural tube undergo differentiation to form cells needed for development • Cells (neuroblasts) become amitotic and migrate • Neuroblasts sprout axons to connect with targets and become neurons Axonal Growth • Growth cone at tip of axon interacts with its environment via: • Cell surface adhesion proteins (_________, integrin, and nerve cell adhesion molecules or N-CAMs) • _________ that attract or repel the growth cone • Nerve growth factor (NGF), which keeps the neuroblast alive • _________ provide physical support and cholesterol essential for construction of synapses Cell Death • About 2/3 of neurons _________ before birth • Death results in cells that fail to make functional synaptic contacts • Many cells also die due to _________ (programmed cell death) during development