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Functions of the Nervous System 1. Sensory input 2. Integration 3. Motor output Divisions of the Nervous System • Central nervous system (CNS) – Brain and spinal cord – Integration and command center • Peripheral nervous system (PNS) – Paired spinal and cranial nerves carry messages to and from the CNS Peripheral Nervous System (PNS) 1. Sensory (afferent) division • • Somatic afferent fibers—convey impulses from skin, skeletal muscles, and joints Visceral afferent fibers—convey impulses from visceral organs 2. Motor (efferent) division • Transmits impulses from the CNS to effector organs Motor Division of PNS 1. Somatic (voluntary) nervous system – Conscious control of skeletal muscles 2. Autonomic (involuntary) nervous system (ANS) – Visceral motor nerve fibers – Regulates smooth muscle, cardiac muscle, and glands – Two functional subdivisions • • Sympathetic Parasympathetic Peripheral nervous system (PNS) Central nervous system (CNS) Cranial nerves and spinal nerves Communication lines between the CNS and the rest of the body Brain and spinal cord Integrative and control centers Sensory (afferent) division Somatic and visceral sensory nerve fibers Conducts impulses from receptors to the CNS Somatic sensory fiber Motor (efferent) division Motor nerve fibers Conducts impulses from the CNS to effectors (muscles and glands) Somatic nervous system Somatic motor (voluntary) Conducts impulses from the CNS to skeletal muscles Skin Visceral sensory fiber Stomach Skeletal muscle Motor fiber of somatic nervous system Sympathetic division Mobilizes body systems during activity Sympathetic motor fiber of ANS Structure Function Sensory (afferent) division of PNS Motor (efferent) division of PNS Parasympathetic motor fiber of ANS Autonomic nervous system (ANS) Visceral motor (involuntary) Conducts impulses from the CNS to cardiac muscles, smooth muscles, and glands Parasympathetic division Conserves energy Promotes housekeeping functions during rest Heart Bladder Figure 11.2 Histology of Nervous Tissue 1. Neurons—excitable cells that transmit electrical signals 2. Neruoglia – glial cells • • • • • • Astrocytes (CNS) Microglia (CNS) Ependymal cells (CNS) Oligodendrocytes (CNS) Satellite cells (PNS) Schwann cells (PNS) Astrocytes • Most abundant, versatile, and highly branched glial cells • Cling to neurons, synaptic endings, and capillaries • Support and brace neurons • Help determine capillary permeability • Guide migration of young neurons • Control the chemical environment • Participate in information processing in the brain Microglia • Small, ovoid cells with thorny processes • Migrate toward injured neurons • Phagocytize microorganisms and neuronal debris Ependymal Cells • Range in shape from squamous to columnar • May be ciliated – Line the central cavities of the brain and spinal column • Separate the CNS interstitial fluid from the cerebrospinal fluid in the cavities Oligodendrocytes • Branched cells • Processes wrap CNS nerve fibers, forming insulating myelin sheaths Satellite Cells and Schwann Cells • Satellite cells – Surround neuron cell bodies in the PNS • Schwann cells (neurolemmocytes) – Surround peripheral nerve fibers and form myelin sheaths – Vital to regeneration of damaged peripheral nerve fibers Neurons (Nerve Cells) • Special characteristics: – Long-lived ( 100 years or more) – Amitotic—with few exceptions – High metabolic rate—depends on continuous supply of oxygen and glucose – Plasma membrane functions in: • Electrical signaling • Cell-to-cell interactions during development Cell Body (Perikaryon or Soma) • • • • Biosynthetic center of a neuron Spherical nucleus with nucleolus Well-developed Golgi apparatus Rough ER called Nissl bodies (chromatophilic substance) • Network of neurofibrils (neurofilaments) • Axon hillock—cone-shaped area from which axon arises • Clusters of cell bodies are called nuclei in the CNS, ganglia in the PNS Processes • Dendrites and axons • Bundles of processes are called – Tracts in the CNS – Nerves in the PNS Dendrites • Short, tapering, and diffusely branched • Receptive (input) region of a neuron • Convey electrical signals toward the cell body as graded potentials The Axon • • • • • One axon per cell arising from the axon hillock Long axon = nerve fiber Occasional branches = axon collaterals Numerous terminal branches Knoblike axon terminals (synaptic knobs or boutons) – Secretory region of neuron – Release neurotransmitters to excite or inhibit other cells Axons: Function • Conducting region of a neuron • Generates and transmits nerve impulses (action potentials) away from the cell body • Molecules and organelles are moved along axons by motor molecules in two directions: – Anterograde—toward axonal terminal • Examples: mitochondria, membrane components, enzymes – Retrograde—toward the cell body • Examples: organelles to be degraded, signal molecules, viruses, and bacterial toxins Dendrites (receptive regions) Cell body (biosynthetic center and receptive region) Nucleolus Axon (impulse generating and conducting region) Nucleus Nissl bodies Axon hillock (b) Impulse direction Node of Ranvier Schwann cell Neurilemma (one interTerminal node) branches Axon terminals (secretory region) Figure 11.4b Myelin Sheath • Segmented protein-lipoid sheath around most long or large-diameter axons • It functions to: – Protect and electrically insulate the axon – Increase speed of nerve impulse transmission Myelin Sheaths in the PNS • Schwann cells wraps many times around the axon – Myelin sheath—concentric layers of Schwann cell membrane • Outer collar of perinuclear cytoplasm— peripheral bulge of Schwann cell cytoplasm • Nodes of Ranvier – gaps between adjacent Schwann cells – Sites where axon collaterals can emerge Myelin Sheaths in the CNS • Formed by processes of oligodendrocytes, NOT the whole cells • Nodes of Ranvier are present • No OCPC • Thinnest fibers are unmyelinated White Matter and Gray Matter • White matter – Dense collections of myelinated fibers • Gray matter – Mostly neuron cell bodies and unmyelinated fibers Structural Classification of Neurons Table 11.1 (1 of 3) Functional Classification of Neurons 1. Sensory (afferent) • Transmit impulses from sensory receptors toward the CNS 2. Motor (efferent) • Carry impulses from the CNS to effectors 3. Interneurons (association neurons) • Shuttle signals through CNS pathways; most are entirely within the CNS Neuron Function • Neurons are highly irritable • Respond to adequate stimulus by generating an action potential (nerve impulse) • Impulse is always the same regardless of stimulus (Action potential) Role of Membrane Ion Channels 1. Leakage (nongated) channels—always open 2. Gated channels (three types): – – – Chemically gated (ligand-gated) channels— open with binding of a specific neurotransmitter Voltage-gated channels—open and close in response to changes in membrane potential Mechanically gated channels—open and close in response to physical deformation of receptors Resting Membrane Potential (Vr) • Potential difference across the membrane of a resting cell – Approximately –70 mV in neurons Membrane Potentials That Act as Signals • Membrane potential changes when: 1. Concentrations of ions across the membrane change 2. Permeability of membrane to ions changes • Changes in membrane potential are signals used to receive, integrate and send information Membrane Potentials That Act as Signals • Two types of signals – Graded potentials • Incoming short-distance signals – Action potentials • Long-distance signals of axons (outgoing) Changes in Membrane Potential • Depolarization – A reduction in membrane potential (toward zero) • Hyperpolarization – An increase in membrane potential (away from zero) Graded Potentials • Short-lived, localized changes in membrane potential • Depolarizations or hyperpolarizations • Graded potential spreads as local currents change the membrane potential of adjacent regions Membrane potential (mV) Active area (site of initial depolarization) –70 Resting potential Distance (a few mm) (c) Decay of membrane potential with distance: Because current is lost through the “leaky” plasma membrane, the voltage declines with distance from the stimulus (the voltage is decremental ). Consequently, graded potentials are short-distance signals. Figure 11.10c Action Potential (AP) • Brief reversal of membrane potential with a total amplitude of ~100 mV • Occurs in muscle cells and axons of neurons • Does not decrease in magnitude over distance • Principal means of long-distance neural communication The big picture 1 Resting state 3 Repolarization Membrane potential (mV) 2 Depolarization 3 4 Hyperpolarization 2 Action potential Threshold 1 4 1 Time (ms) Figure 11.11 (1 of 5) 3 2 Action potential Na+ permeability K+ permeability 1 4 1 Relative membrane permeability Membrane potential (mV) The AP is caused by permeability changes in the plasma membrane Time (ms) Figure 11.11 (2 of 5) Coding for Stimulus Intensity • All action potentials are alike and are independent of stimulus intensity • Strong stimuli can generate action potentials more often than weaker stimuli • The CNS determines stimulus intensity by the frequency of impulses Action potentials Threshold Stimulus Time (ms) Figure 11.13 Absolute refractory period Relative refractory period Depolarization (Na+ enters) Repolarization (K+ leaves) After-hyperpolarization Stimulus Time (ms) Figure 11.14 Conduction Velocity • Conduction velocities of neurons vary widely • Effect of axon diameter – Larger = faster • Effect of myelination – Myelination = faster Multiple Sclerosis (MS) • Autoimmune disease that mainly affects young adults • Symptoms: visual disturbances, weakness, loss of muscular control, speech disturbances, and urinary incontinence • Myelin sheaths in the CNS become nonfunctional scleroses • Shunting and short-circuiting of nerve impulses occurs • Impulse conduction slows and eventually ceases The Synapse • A junction that mediates information transfer from one neuron: – To another neuron – To an effector cell • Electrical or Chemical • Presynaptic neuron—conducts impulses toward the synapse • Postsynaptic neuron—transmits impulses away from the synapse Axodendritic synapses Dendrites Axosomatic synapses Cell body Axoaxonic synapses (a) Axon Axon Axosomatic synapses (b) Cell body (soma) of postsynaptic neuron Figure 11.16 Electrical Synapses • Less common than chemical synapses – Neurons are electrically coupled (joined by gap junctions) – Communication = very rapid • may be unidirectional or bidirectional – Important in: • Embryonic nervous tissue • Some brain regions Chemical Synapses • Specialized for the release and reception of neurotransmitters • Typically composed of two parts – Axon terminal of the presynaptic neuron, which contains synaptic vesicles – Receptor region on the postsynaptic neuron Chemical synapses transmit signals from one neuron to another using neurotransmitters. Presynaptic neuron Presynaptic neuron Postsynaptic neuron 1 Action potential arrives at axon terminal. 2 Voltage-gated Ca2+ channels open and Ca2+ enters the axon terminal. Mitochondrion Ca2+ Ca2+ Ca2+ 3 Ca2+ entry causes neurotransmittercontaining synaptic vesicles to release their contents by exocytosis. Axon terminal Ca2+ Synaptic cleft Synaptic vesicles 4 Neurotransmitter diffuses across the synaptic cleft and binds to specific receptors on the postsynaptic membrane. Postsynaptic neuron Ion movement Enzymatic degradation Graded potential Reuptake Diffusion away from synapse 5 Binding of neurotransmitter opens ion channels, resulting in graded potentials. 6 Neurotransmitter effects are terminated by reuptake through transport proteins, enzymatic degradation, or diffusion away from the synapse. Figure 11.17 Postsynaptic Potentials • Types of postsynaptic potentials – EPSP—excitatory postsynaptic potentials – IPSP—inhibitory postsynaptic potentials Excitatory Synapses and EPSPs • Neurotransmitter binding opens chemically gated channels • Allows simultaneous flow of Na+ and K+ in opposite directions • Na+ influx greater than K+ efflux net depolarization called EPSP (not AP) • EPSP help trigger AP if EPSP is of threshold strength – Can spread to axon hillock, trigger opening of voltagegated channels, and cause AP to be generated Membrane potential (mV) Figure 11.18a Postsynaptic potentials can be excitatory or inhibitory. +30 0 Threshold –55 –70 An EPSP is a local depolarization of the postsynaptic membrane that brings the neuron closer to AP threshold. Neurotransmitter binding opens chemically gated ion channels, allowing Na+ and K+ to pass through simultaneously. Stimulus 10 20 30 Time (ms) Excitatory postsynaptic potential (EPSP) © 2013 Pearson Education, Inc. Inhibitory Synapses and IPSPs • Reduces postsynaptic neuron's ability to produce an action potential – Makes membrane more permeable to K+ or Cl– • If K+ channels open, it moves out of cell • If Cl- channels open, it moves into cell – Neurotransmitter hyperpolarizes cell • Inner surface of membrane becomes more negative • AP less likely to be generated Membrane potential (mV) Figure 11.18b Postsynaptic potentials can be excitatory or inhibitory. +30 0 Threshold An IPSP is a local hyperpolarization of the postsynaptic membrane that drives the neuron away from AP threshold. Neurotransmitter binding opens K+ or Cl– channels. –55 –70 Stimulus 10 20 30 Time (ms) Inhibitory postsynaptic potential (IPSP) © 2013 Pearson Education, Inc. Synaptic Integration: Summation • A single EPSP cannot induce an AP • EPSPs and IPSPs can summate to influence postsynaptic neuron • Most neurons receive both excitatory and inhibitory inputs from thousands of other neurons – Only if EPSP's predominate and bring to threshold AP Two Types of Summation • Temporal summation – One + presynaptic neurons transmit rapid-fire impulses • Spatial summation – Postsynaptic neuron stimulated simultaneously by large number of terminals at same time Neurotransmitters • Most neurons make two or more neurotransmitters, which are released at different stimulation frequencies • 50 or more neurotransmitters have been identified • Classified by chemical structure and by function Chemical Classes of Neurotransmitters • Acetylcholine (Ach) – Released at neuromuscular junctions and some ANS neurons • Biogenic amines include: – Broadly distributed in the brain – Play roles in emotional behaviors and the biological clock • Catecholamines – Dopamine, norepinephrine (NE), and epinephrine • Indolamines – Serotonin and histamine Chemical Classes of Neurotransmitters • Amino acids include: – GABA—Gamma ()-aminobutyric acid – Glycine – Glutamate • Peptides (neuropeptides) include: • Substance P – Mediator of pain signals • Endorphins – Act as natural opiates; reduce pain perception • Gut-brain peptides – Somatostatin and cholecystokinin Chemical Classes of Neurotransmitters • Purines such as ATP: – Act in both the CNS and PNS – Produce fast or slow responses – Induce Ca2+ influx in astrocytes – Provoke pain sensation Chemical Classes of Neurotransmitters • Gases and lipids – Nitric oxide (NO) • Synthesized on demand • Involved in learning and memory – Carbon monoxide (CO) is a regulator of cGMP in the brain – Endocannabinoids • Lipid soluble; synthesized on demand from membrane lipids • Involved in learning and memory Functional Classification of Neurotransmitters • Neurotransmitter effects may be excitatory (depolarizing) and/or inhibitory (hyperpolarizing) – Determined by the receptor type of the postsynaptic neuron – Acetylcholine • Excitatory at neuromuscular junctions in skeletal muscle • Inhibitory in cardiac muscle Neurotransmitter Actions • Direct action – Neurotransmitter binds to channel-linked receptor and opens ion channels – Promotes rapid responses • Examples: ACh and amino acids • Indirect action – Neurotransmitter binds to a G protein-linked receptor and acts through an intracellular second messenger – Promotes long-lasting effects • Examples: biogenic amines, neuropeptides, and dissolved gases Neural Integration: Neuronal Pools • Functional groups of neurons that: – Integrate incoming information – Forward the processed information to other destinations • Simple neuronal pool – Single presynaptic fiber branches and synapses with several neurons in the pool – Discharge zone—neurons most closely associated with the incoming fiber – Facilitated zone—neurons farther away from incoming fiber Presynaptic (input) fiber Facilitated zone Discharge zone Facilitated zone Figure 11.21 Types of Circuits in Neuronal Pools • • • • Diverging circuit Converging Reverberating Parallel after-discharge Patterns of Neural Processing • Serial processing – Input travels along one pathway to a specific destination – Works in an all-or-none manner to produce a specific response – Example: Reflexes • rapid, automatic responses to stimuli that always cause the same response • Reflex arcs (pathways) have five essential components: receptor, sensory neuron, CNS integration center, motor neuron, and effector Stimulus 1 Receptor Interneuron 2 Sensory neuron 3 Integration center 4 Motor neuron 5 Effector Spinal cord (CNS) Response Figure 11.23 Patterns of Neural Processing • Parallel processing – Input travels along several pathways – One stimulus promotes numerous responses – Important for higher-level mental functioning • Example: a smell may remind one of the odor and associated experiences