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Neural Signaling Chapter 40 Learning Objective 1 • Describe the processes involved in neural signaling: reception, transmission, integration, and action by effectors Neural Signaling 1 (1) Reception of information • by a sensory receptor (2) Transmission by an afferent neuron • to the central nervous system (CNS) (3) Integration by interneurons • in the central nervous system (CNS) Neural Signaling 2 (4) Transmission by an efferent neuron • to other neurons or effector (5) Action by effectors • the muscles and glands Peripheral Nervous System (PNS) • Made up of • • sensory receptors neurons outside the CNS Response to Stimulus External stimulus (e.g., vibration, movement, light, odor) Internal stimulus (e.g., change in blood pH or blood pressure) RECEPTION Detection by external sense organs Detection by internal sense organs TRANSMISSION Sensory (afferent) neurons transmit information Fig. 40-1a, p. 846 Central Nervous System (brain and spinal cord) INTEGRATION Interneurons sort and interpret information TRANSMISSION Motor (efferent) neurons transmit impulses ACTION BY EFFECTORS (muscles and glands) e.g., animal runs away e.g., espiration rate increases; blood pressure rises Fig. 40-1b, p. 846 External stimulus (e.g., vibration, movement, light, odor) Internal stimulus (e.g., change in blood pH or blood pressure) RECEPTION Detection by internal sense organs Detection by external sense organs TRANSMISSION Sensory (afferent) neurons transmit information Central Nervous System (brain and spinal cord) INTEGRATION Interneurons sort and interpret information TRANSMISSION Motor (efferent) neurons transmit impulses ACTION BY EFFECTORS (muscles and glands) e.g., animal runs away e.g., espiration rate increases; blood pressure rises Stepped Art Fig. 40-1, p. 846 KEY CONCEPTS • Neural signaling involves reception, transmission, integration, and action by effectors Learning Objective 2 • What is the structure of a typical neuron? • Give the function of each of its parts Neurons • Specialized to • • • receive stimuli transmit electrical and chemical signals Cell body • contains nucleus and organelles Dendrites • Many branched dendrites • • extend from cell body of neuron specialized to receive stimuli and send signals to the cell body Axons 1 • A single long axon • • • extends from neuron cell body forms branches (axon collaterals) Transmits signals into terminal branches • which end in synaptic terminals Axons 2 • Myelin sheath • • • surrounds many axons insulates Schwann cells • form the myelin sheath in the PNS Axons 3 • In the CNS • • sheath is formed by other glial cells Nodes of Ranvier • gaps in sheath between successive Schwann cells Neuron Structure Dendrites covered with dendritic spines Cell body Axon collateral Cytoplasm of Schwann cell Synaptic terminals Axon Nucleus Myelin sheath Nucleus Axon Nodes of Ranvier Schwann Terminal cell branches Fig. 40-2, p. 847 Nerves and Ganglia • Nerve • • • several hundred axons wrapped in connective tissue Ganglion • mass of neuron cell bodies in the PNS Nerve Structure Ganglion Cell bodies Myelin sheath Artery Vein Axon (a) Fig. 40-3a, p. 848 100 µm (b) Fig. 40-3b, p. 848 Learn more about the structure of neurons and nerves by clicking on the figures in ThomsonNOW. Learning Objective 3 • Name the main types of glial cells • Describe the functions of each Glial Cells • Support and nourish neurons • Are important in neural communication Glial Cell Types 1 • Astrocytes • • • • physically support neurons regulate extracellular fluid in CNS (by taking up excess potassium ions) communicate with one another (and with neurons) induce and stabilize synapses Glial Cell Types 2 • Oligodendrocytes • • form myelin sheaths around axons in CNS Schwann cells • form sheaths around axons in PNS Glial Cell Types 3 • Microglia • • Phagocytic cells Ependymal Cells • • • line cavities in the CNS contribute to formation of cerebrospinal fluid serve as neural stem cells KEY CONCEPTS • Neurons are specialized to receive stimuli and transmit signals; glial cells are supporting cells that protect and nourish neurons and that can modify neural signals Learning Objective 4 • How does the neuron develop and maintain a resting potential? Neural Signals • Electrical signals transmit information • along axons • Plasma membrane of resting neuron (not transmitting an impulse) is polarized • Inner surface of plasma membrane is negatively charged • relative to extracellular fluid Resting Potential • Potential difference of about -70 mV • • across the membrane Magnitude of resting potential (1) differences in ion concentrations (Na+, K+) inside cell relative to extracellular fluid (2) selective permeability of plasma membrane to these ions Axon 40 20 0 –20 –40 –60 –80 –70 mV Time Amplifier Plasma membrane + – – + Electrode placed inside the cell + – – + + – – + Electrode placed outside the cell + – + – – + + – – + – + (a) Measuring the resting potential of a neuron. Fig. 40-4a, p. 850 Ions • Pass through specific passive ion channels • • • K+ leak out faster than Na+ leak in Cl- accumulate at inner surface of plasma membrane Large anions (proteins) • • cannot cross plasma membrane contribute negative charges Sodium–Potassium Pumps • Maintain gradients that determine resting potential • • transport 3 Na+ out for each 2 K+ in Require ATP Extracellular fluid 3 Na+ Na+ K+ Diffusion out K+ K+ CI Na+ – + + + + + – – – – – – K+ K+ CI– Diffusion in A Na /K pump K+ K+ Na+ _ CI– Plasma membrane Na+ K+ Na+ Na+ Na+ + K+ 2 K+ _ A _ CI CI– A – Cytoplasm (b) Permeability of the neuron membrane. CI– Fig. 40-4b, p. 850 KEY CONCEPTS • The resting potential of a neuron is maintained by differences in concentrations of specific ions inside the cell relative to the extracellular fluid and by selective permeability of the plasma membrane to these ions Learning Objective 5 • Compare a graded potential with an action potential • Describe the production and transmission of each Membrane Potential • Membrane is depolarized • • if stimulus causes membrane potential to become less negative Membrane is hyperpolarized • if membrane potential becomes more negative than resting potential Graded Potential • A local response • Varies in magnitude • • depending on strength of applied stimulus Fades out • within a few millimeters of point of origin Action Potential 1 • Action potential is a wave of depolarization • • that moves down the axon Generated when • • • voltage across the membrane declines to a critical point (threshold level) voltage-activated ion channels open Na+ ions flow into the neuron Voltage-Activated Ion Channels Extracellular fluid Activation gate Cytoplasm Inactivation gate (a) Sodium channels. (b) Potassium channels. Fig. 40-6, p. 852 Voltage-Activated Ion Channels During an Action Potential Membrane potential (mV) Spike Depolarization Repolarization Threshold level Resting state Time (milliseconds) (a) Action potential. Fig. 40-7a, p. 853 Axon Extracellular fluid Sodium channel Potassium channel Cytoplasm 1 Resting state. 2 Depolarization. 3 Repolarization. 4 Return to resting state. (b) The action of the ion channels in the plasma membrane determines the state of the neuron. Fig. 40-7b, p. 853 Action Potential 2 • An all-or-none response • • • no variation in strength of a single impulse either membrane potential exceeds threshold level or it does not Once begun, an action potential is selfpropagating Repolarization • As an action potential moves down an axon, repolarization occurs behind it Transmission of an Action Potential Stimulus Axon Area of depolarization Potassium channel Action potential Sodium channel (1) Action potential is transmitted as wave of depolarization that travels down axon. At region of depolarization, Na+ diffuse into cell. Fig. 40-8a, p. 854 Area of repolarization Area of depolarization Action potential (2) As action potential progresses along axon, repolarization occurs quickly behind it. Fig. 40-8b, p. 854 Refractory Periods • During depolarization, the axon enters an absolute refractory period • • when it can’t transmit another action potential When enough gates controlling Na+ channels have been reset, the neuron enters a relative refractory period • when the threshold is higher Learn more about ion channels and action potentials by clicking on the figures in ThomsonNOW. KEY CONCEPTS • Depolarization of the neuron plasma membrane to threshold level generates an action potential, an electrical signal that travels as a wave of depolarization along the axon Learning Objective 6 • Contrast continuous conduction with saltatory conduction Continuous Conduction • Involves entire axon plasma membrane • Takes place in unmyelinated neurons Saltatory Conduction • Depolarization skips along axon from one node of Ranvier to the next • • • more rapid than continuous conduction takes place in myelinated neurons Nodes of Ranvier • • sites where axon is not covered by myelin Na+ channels are concentrated Saltatory Conduction Area of action potential 1 Saltatory conduction Nodes of Ranvier Axon Schwann cell 2 Fig. 40-9a, p. 855 3 4 Direction of depolarization Fig. 40-9b, p. 855 Learning Objective 7 • Describe the actions of the neurotransmitters identified in the chapter Synapses • Junctions between two neurons • • or between a neuron and effector Most synapses are chemical • some are electrical synapses Synaptic Transmission • A presynaptic neuron releases neurotransmitter (chemical messenger) from its synaptic vesicles Neurotransmitters 1 • Acetylcholine • • triggers contraction of skeletal muscle Biogenic amines • • • norepinephrine, serotonin, dopamine important in regulating mood dopamine is also important in motor function Neurotransmitters 2 • Some amino acids • • • glutamate (excitatory neurotransmitter in brain) GABA (widespread inhibitory neurotransmitter) Neuropeptides (opioids) • • endorphins (e.g. beta-endorphin) enkephalins Neurotransmitters 3 • Nitric oxide (NO) • • gaseous neurotransmitter transmits signals from postsynaptic neuron to presynaptic neuron (opposite direction from other neurotransmitters) Learning Objective 8 • Trace the events that take place in synaptic transmission • Draw diagrams to support your description Synaptic Transmission • Calcium ions cause synaptic vesicles to fuse with presynaptic membrane • • releases neurotransmitter into synaptic cleft Neurotransmitter diffuses across the synaptic cleft • combines with specific receptors on a postsynaptic neuron Synaptic Transmission Synaptic vesicles Plasma membrane of postsynaptic neuron 0.25 µm (a) The TEM shows synaptic terminals filled with synaptic vesicles. Fig. 40-10a, p. 858 Fig. 40-10bc, p. 858 Axon of presynaptic neuron Voltage-gated Ca2+ channel Synaptic terminal 1 Synaptic vesicle Ca2+ 2 Neurotransmitter molecule 3 4 Ligand-gated channels Postsynaptic neuron 5 Receptor for neurotransmitter Postsynaptic membrane (b) How a neural impulse is transmitted across a synapse. Fig. 40-10b, p. 858 Ca2+ Synaptic terminal Presynaptic membrane Synaptic cleft Na+ Postsynaptic membrane (c) Neurotransmitter binds with receptor. Ligand-gated channel opens, resulting in depolarization. Fig. 40-10c, p. 858 Neurotransmitter Receptors • Many are proteins that form ligand-gated ion channels • Others work through a second messenger such as cAMP Learn more about synaptic transmission by clicking on the figure in ThomsonNOW. KEY CONCEPTS • Neurons signal other cells by releasing neurotransmitters at synapses Learning Objective 9 • Compare excitatory and inhibitory signals and their effects Binding of Neurotransmitter to a Receptor • Binding causes either • • • excitatory postsynaptic potential (EPSP) or inhibitory postsynaptic potential (IPSP) Depending on the type of receptor EPSPs and IPSPs • EPSPs • • bring neuron closer to firing IPSPs • move neuron farther away from its firing level Learning Objective 10 • Define neural integration • Describe how a postsynaptic neuron integrates incoming stimuli and “decides” whether or not to fire Neural Integration • Process of summing (integrating) incoming signals • Summation • process of adding and subtracting incoming signals Summation • Each EPSP or IPSP is a graded potential • • • vary in magnitude depending on strength of stimulus applied Summation of several EPSPs • brings neuron to critical firing level Temporal Summation • Occurs when repeated stimuli cause new EPSPs to develop before previous EPSPs have decayed Spatial Summation • Occurs when several closely spaced synaptic terminals release neurotransmitter simultaneously • stimulating postsynaptic neuron at several different places Neural Integration Postsynaptic membrane potential (mV) Threshold level Resting potential Time (msec) (a) Subthreshold (no summation). (b) Temporal summation. (c) Spatial summation. (d) Spatial summation of EPSPs and IPSPs. Fig. 40-11, p. 860 KEY CONCEPTS • During integration, incoming neural signals are summed; temporal and spatial summation can bring a neuron to threshold level Learning Objective 11 • Distinguish among convergence, divergence, and reverberation • Explain why each is important Neural Circuits • Complex neural circuits are possible because of associations such as convergence and divergence Convergence • A single neuron is affected by converging signals from two or more presynaptic neurons • Allows CNS to integrate incoming information from various sources Divergence • A single presynaptic neuron stimulates many postsynaptic neurons • allowing widespread effect Neural Circuits (a) Convergence of neural input. Several presynaptic neurons synapse with one postsynaptic neuron. (b) Divergence of neural output. A single presynaptic neuron synapses with many postsynaptic neurons. Fig. 40-12, p. 861 Reverberating Circuits • Important in • • • • rhythmic breathing mental alertness short-term memory Depend on positive feedback • new impulses generated again and again until synapses fatigue Reverberating Circuits 1 2 (a) Simple reverberating circuit. An axon collateral of the second neuron turns back on its own dendrites, so the neuron continues to stimulate itself. Fig. 40-13a, p. 861 Interneuron Axon collateral 1 2 3 (b) Reverberating circuit with interneuron. An axon collateral of the second neuron synapses with an interneuron. The interneuron synapses with the first neuron in the sequence. New impulses are triggered again and again in the first neuron, causing reverberation. Fig. 40-13b, p. 861