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Human Anatomy & Physiology Ninth Edition CHAPTER 11 Fundamentals of the Nervous System and Nervous Tissue: Part 1 © Annie Leibovitz/Contact Press Images © 2013 Pearson Education, Inc. Figure 11.1 The nervous system’s functions. Sensory input Integration Motor output 6/27/2012 MDufilho 2 Divisions of the Nervous System • The Tale of Two Brains • 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 6/27/2012 MDufilho 3 Peripheral Nervous System (PNS) • Two functional divisions – Sensory (afferent) division • Somatic sensory fibers—convey impulses from skin, skeletal muscles, and joints to CNS • Visceral sensory fibers—convey impulses from visceral organs to CNS – Motor (efferent) division • Transmits impulses from CNS to effector organs – Muscles and glands • Two divisions – Somatic nervous system – Autonomic nervous system 6/27/2012 MDufilho 4 Motor Division of PNS: Somatic Nervous System • Somatic motor nerve fibers • Conducts impulses from CNS to skeletal muscle • Voluntary nervous system – Conscious control of skeletal muscles 6/27/2012 MDufilho 5 Motor Division of PNS: Autonomic Nervous System • Visceral motor nerve fibers • Regulates smooth muscle, cardiac muscle, and glands • Involuntary nervous system • Two functional subdivisions – Sympathetic – Parasympathetic – Work in opposition to each other 6/27/2012 MDufilho 6 Figure 11.2 Levels of organization in the nervous system. Central nervous system (CNS) Peripheral nervous system (PNS) Brain and spinal cord Cranial nerves and spinal nerves Integrative and control centers Communication lines between the CNS and the rest of the body Sensory (afferent) division Motor (efferent) division Somatic and visceral sensory nerve fibers Conducts impulses from receptors to the CNS Somatic sensory fiber Skin 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 Visceral sensory fiber Stomach Autonomic nervous system (ANS) Visceral motor (involuntary) Conducts impulses from the CNS to cardiac muscles, smooth muscles, and glands Skeletal muscle Motor fiber of somatic nervous system Sympathetic division Mobilizes body systems during activity Parasympathetic division Conserves energy Promotes housekeeping functions during rest Sympathetic motor fiber of ANS Heart Structure Function Sensory (afferent) division of PNS Motor (efferent) division of PNS 6/27/2012 Parasympathetic motor fiber of ANS MDufilho Bladder 7 Histology of Nervous Tissue • Highly cellular; little extracellular space – Tightly packed • Two principal cell types – Neurons (nerve cells)—excitable cells that transmit electrical signals – Neuroglia – small cells that surround and wrap delicate neurons • • • • • • 6/27/2012 Astrocytes (CNS) Microglial cells (CNS) Ependymal cells (CNS) Oligodendrocytes (CNS) Satellite cells (PNS) Schwann cells (PNS) MDufilho 8 Supporting Cells: Neuroglia • The supporting cells (neuroglia or glial cells): – Provide a supportive scaffolding for neurons – Segregate and insulate neurons – Assist with repair after damage – Guide young neurons to the proper connections – Promote health and growth 6/27/2012 MDufilho 9 Resting Membrane Potential (Vr) • Potential difference across the membrane of a resting cell – Approximately –70 mV in neurons (cytoplasmic side of membrane is negatively charged relative to outside) • Generated by ????? 6/27/2012 MDufilho 10 Role of Membrane Ion Channels: 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, as in sensory receptors 6/27/2012 MDufilho 11 Figure 11.6 Operation of gated channels. Chemically gated ion channels Open in response to binding of the appropriate neurotransmitter Voltage-gated ion channels Open in response to changes in membrane potential Neurotransmitter chemical attached to receptor Receptor Membrane voltage changes Chemical binds Closed 6/27/2012 Open MDufilho Closed Open 12 Resting Membrane Potential: Differences in Ionic Composition - Review • ECF has higher concentration of ___than ICF – Balanced chiefly by ________________ • ICF has higher concentration of _____than ECF – Balanced by _________________________ • ___plays most important role in membrane potential PLAY 6/27/2012 A&P Flix™: Resting Membrane Potential MDufilho 13 Differences in Plasma Membrane Permeability - Review • Impermeable large ______________ • Slightly permeable to _____(through leakage channels) – ________diffuses into cell down concentration gradient • 25 times more permeable to ____than sodium (more leakage channels) – _________diffuses out of cell down concentration gradient • Quite permeable to _____ 6/27/2012 MDufilho 14 Resting Membrane Potential – Review • More potassium diffuses out than sodium diffuses in – Cell more ________inside – Establishes resting membrane potential • ___________________stabilizes resting membrane potential – Maintains concentration gradients for Na+ and K+ – __Na+ pumped out of cell; two ___pumped in 6/27/2012 MDufilho 15 Membrane Potential Changes Used as Communication Signals • Membrane potential changes when – Concentrations of ions across membrane change – Membrane permeability to ions changes • Changes produce two types signals – Graded potentials • Incoming signals operating over short distances – Action potentials • Long-distance signals of axons • Changes in membrane potential used as signals to receive, integrate, and send information 6/27/2012 MDufilho 16 Figure 11.9a Depolarization and hyperpolarization of the membrane. Membrane potential (voltage, mV) Depolarizing stimulus +50 Inside positive 0 Inside negative Depolarization –50 –70 Resting potential –100 0 1 2 3 4 Time (ms) 5 6 7 Depolarization: The membrane potential moves toward 0 mV, the inside becoming less negative (more positive). 6/27/2012 MDufilho 17 Changes in Membrane Potential • Terms describing membrane potential changes relative to resting membrane potential • Hyperpolarization – An increase in membrane potential (away from zero) – Inside of cell more negative than resting membrane potential) – Reduces probability of producing a nerve impulse 6/27/2012 MDufilho 18 Figure 11.9b Depolarization and hyperpolarization of the membrane. Membrane potential (voltage, mV) Hyperpolarizing stimulus +50 0 –50 Resting potential –70 Hyperpolarization –100 0 1 2 3 4 Time (ms) 5 6 7 Hyperpolarization: The membrane potential increases, the inside becoming more negative. 6/27/2012 MDufilho 19 Graded Potentials • Short-lived, localized changes in membrane potential – Magnitude varies with stimulus strength – Stronger stimulus more voltage changes; farther current flows • Either depolarization or hyperpolarization • Triggered by stimulus that opens gated ion channels • Current flows but dissipates quickly and decays – Graded potentials are signals only over short distances 6/27/2012 MDufilho 20 Figure 11.10 The spread and decay of a graded potential. Stimulus Depolarized region Plasma membrane Depolarization: A small patch of the membrane (red area) depolarizes. Membrane potential (mV) Depolarization spreads: Opposite charges attract each other. This creates local currents (black arrows) that depolarize adjacent membrane areas, spreading the wave of depolarization. Active area (site of initial depolarization) –70 Resting potential Distance (a few mm) 6/27/2012 Membrane potential decays 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. MDufilho 21 Figure 11.11 The action potential (AP) is a brief change in membrane potential in a “patch” of membrane that is depolarized by local currents. The big picture Resting state Membrane potential (mV) 1 The key players 2 Voltage-gated Na+ channels Voltage-gated K+ channels Outside cell Outside cell Depolarization +30 3 3 Repolarization 0 Action potential 2 4 Hyperpolarization Inactivation gate Inside Activation cell gate Closed Opened Closed Opened The events Threshold –55 –70 1 1 2 3 Time (ms) Sodium channel 1 4 0 0 Action potential Na+ permeability K+ permeability 2 –55 –70 1 0 6/27/2012 1 4 1 2 3 Time (ms) Relative membrane permeability +30 3 Potassium channel 4 Activation gates Inactivation gate The AP is caused by permeability changes in the plasma membrane: Membrane potential (mV) Inactivated Inside cell 4 MDufilho 1 Resting state 4 Hyperpolarization 2 Depolarization 3 Repolarization 22 Threshold • Not all depolarization events produce APs • For axon to "fire", depolarization must reach threshold – That voltage at which the AP is triggered • At threshold: – Membrane has been depolarized by 15 to 20 mV – Na+ permeability increases – Na influx exceeds K+ efflux – The positive feedback cycle begins 6/27/2012 MDufilho 23 Membrane potential (mV) Figure 11.12 Propagation of an action potential (AP). Voltage at 2 ms +30 Voltage at 0 ms Voltage at 4 ms –70 Recording electrode Time = 0 ms. Action potential has not yet reached the recording electrode. Resting potential Time = 2 ms. Action potential peak reaches the recording electrode. Time = 4 ms. Action potential peak has passed the recording electrode. Membrane at the recording electrode is still hyperpolarized. Peak of action potential Hyperpolarization 6/27/2012 MDufilho 24 Stimulus voltage Membrane potential (mV) Figure 11.13 Relationship between stimulus strength and action potential frequency. Action potentials +30 –70 Threshold Stimulus 0 Time (ms) 6/27/2012 MDufilho 25 Figure 11.14 Absolute and relative refractory periods in an AP. Relative refractory period Absolute refractory period Membrane potential (mV) Depolarization (Na+ enters) +30 0 Repolarization (K+ leaves) Hyperpolarization –70 Stimulus 0 6/27/2012 MDufilho 1 2 Time (ms) 3 4 5 26 Figure 11.15 Action potential propagation in nonmyelinated and myelinated axons. Stimulus Size of voltage Stimulus Voltage-gated ion channel In bare plasma membranes, voltage decays. Without voltage-gated channels, as on a dendrite, voltage decays because current leaks across the membrane. In nonmyelinated axons, conduction is slow (continuous conduction). Voltage-gated Na+ and K+ channels regenerate the action potential at each point along the axon, so voltage does not decay. Conduction is slow because it takes time for ions and for gates of channel proteins to move, and this must occur before voltage can be regenerated. Stimulus Myelin sheath 6/27/2012 In myelinated axons, conduction is fast (saltatory conduction). Myelin keeps current in axons (voltage doesn’t decay much). APs are generated only in the myelin sheath gaps and appear to jump rapidly from gap to gap. MDufilho Myelin sheath Myelin sheath gap 1 mm 27 The Synapse • Nervous system works because information flows from neuron to neuron • Neurons functionally connected by synapses – Junctions that mediate information transfer • From one neuron to another neuron • Or from one neuron to an effector cell 6/27/2012 MDufilho 28 Figure 11.16 Synapses. Axodendritic synapses Dendrites Axosomatic synapses Cell body Axoaxonal synapses Axon Axon Axosomatic synapses 6/27/2012 Cell body (soma) of postsynaptic neuron MDufilho 29 Varieties of Synapses: Chemical Synapses • Specialized for release and reception of chemical neurotransmitters • Typically composed of two parts – Axon terminal of presynaptic neuron • Contains synaptic vesicles filled with neurotransmitter – Neurotransmitter receptor region on postsynaptic neuron's membrane • Usually on dendrite or cell body • Two parts separated by synaptic cleft – Fluid-filled space • Electrical impulse changed to chemical across synapse, then back into electrical 6/27/2012 MDufilho 30 Varieties of Synapses: Electrical Synapses • Less common than chemical synapses – Neurons electrically coupled (joined by gap junctions that connect cytoplasm of adjacent neurons) • Communication very rapid • May be unidirectional or bidirectional • Synchronize activity – More abundant in: • Embryonic nervous tissue • Nerve impulse remains electrical 6/27/2012 MDufilho 31 Figure 11.17 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. 3 Ca2+ entry causes synaptic vesicles to release neurotransmitter by exocytosis Mitochondrion Synaptic cleft Axon terminal 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. 6/27/2012 MDufilho 32 Postsynaptic Potentials • Neurotransmitter receptors cause graded potentials that vary in strength with – Amount of neurotransmitter released and – Time neurotransmitter stays in area • Types of postsynaptic potentials – EPSP—excitatory postsynaptic potentials – IPSP—inhibitory postsynaptic potentials 6/27/2012 MDufilho 33 Table 11.2 Comparison of Graded Potentials and Action Potentials (1 of 4) 6/27/2012 MDufilho 34 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) 6/27/2012 MDufilho 35 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) 6/27/2012 MDufilho 36 Table 11.2 Comparison of Graded Potentials and Action Potentials (4 of 4) 6/27/2012 MDufilho 37 Synaptic Integration: Summation • A single EPSP cannot induce an AP • EPSPs can summate to influence postsynaptic neuron • IPSPs can also summate • Most neurons receive both excitatory and inhibitory inputs from thousands of other neurons – Only if EPSP's predominate and bring to threshold AP 6/27/2012 MDufilho 38 Figure 11.19 Neural integration of EPSPs and IPSPs. E1 E1 E1 E1 Membrane potential (mV) E2 l1 0 Threshold of axon of postsynaptic neuron Resting potential –55 –70 E1 E1 E1 E1 Time Time No summation: 2 stimuli separated in time cause EPSPs that do not add together. Temporal summation: 2 excitatory stimuli close in time cause EPSPs that add together. E1 + E2 Time Spatial summation: 2 simultaneous stimuli at different locations cause EPSPs that add together. l1 E1 + l1 Time Spatial summation of EPSPs and IPSPs: Changes in membane potential can cancel each other out. Excitatory synapse 1 (E1) Excitatory synapse 2 (E2) Inhibitory synapse (I1) 6/27/2012 MDufilho 39