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Nervous Tissue overview of the nervous system properties of neurons
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. overview of the nervous system properties of neurons supportive cells (neuroglia) electrophysiology of neurons synapses neural integration Neurofibrils Axon (d) Figure 12.4d
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Overview of Nervous System
nervous system carries out its task in three basic steps: Sense organs receive information They transmit coded messages to the spinal cord and the brain The brain and spinal cord processes this information, relates it to past experiences, and determine what response is appropriate to the circumstances and issues commands to muscles and gland cells to carry out such a response
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Two Major Anatomical Subdivisions of Nervous System
central nervous system (CNS) brain and spinal cord enclosed in bony coverings enclosed by cranium and vertebral column peripheral nervous system (PNS) all the nervous system except the brain and spinal cord composed of nerves and ganglia nerve – a bundle of nerve fibers (axons) wrapped in fibrous connective tissue ganglion – a knot-like swelling in a nerve where neuron cell bodies are concentrated
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Subdivisions of Nervous System
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Central nervous system (CNS) Peripheral nervous system (PNS) Brain Spinal cord Nerves Ganglia Figure 12.1
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Subdivisions of Nervous System
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Central nervous system Peripheral nervous system Spinal cord Sensory division Motor division Brain Visceral sensory division Somatic sensory division Visceral motor division Somatic motor division Figure 12.2 Sympathetic division Parasympathetic division
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Functional Classes of Neurons
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Peripheral nervous system Central nervous system 1 Sensory (afferent) neurons conduct signals from receptors to the CNS. 2 Interneurons (association neurons) are confined to the CNS. 3 Motor (efferent) neurons conduct signals from the CNS to effectors such as muscles and glands. Figure 12.3
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Functional Types of Neurons
sensory (afferent) neurons specialized to detect stimuli transmit information about them to the CNS begin in almost every organ in the body and end in CNS afferent – conducting signals toward CNS interneurons (association) neurons lie entirely within the CNS process, store, and retrieve information and ‘make decisions’ that determine how the body will respond to stimuli 99% of all neurons are interneurons motor (efferent) neuron send signals out to muscles and gland cells (the effectors) motor because most of them lead to muscles efferent neurons conduct signals away from the CNS
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Structure of a Neuron Figure 12.4a Figure 12.4a
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. soma – the control center of the neuron also called neurosoma, cell body, or perikaryon dendrites – vast number of branches coming from a few thick branches from the soma primary site for receiving signals from other neurons axon (nerve fiber) – originates from a mound on one side of the soma called the axon hillock cylindrical, relatively unbranched for most of its length distal end of the axon has the Telodendria– extensive complex of fine branches Dendrites Soma Nucleus Nucleolus Trigger zone: Axon hillock Initial segment Axon collateral Axon Direction of signal transmission Internodes Node of Ranvier Myelin sheath Schwann cell Figure 12.4a Synaptic knobs 12-8 Figure 12.4a (a)
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Universal Properties of Neurons
excitability (irritability) respond to environmental changes called stimuli conductivity neurons respond to stimuli by producing electrical signals that are quickly conducted to other cells at distant locations secretion when electrical signal reaches end of nerve fiber, a chemical neurotransmitter is secreted that crosses the gap and stimulates the next cell
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Variation in Neuron Structure
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. multipolar neuron one axon and multiple dendrites most common most neurons in the brain and spinal cord bipolar neuron one axon and one dendrite olfactory cells, retina, inner ear unipolar neuron single process leading away from the soma sensory from skin and organs to spinal cord anaxonic neuron many dendrites but no axon help in visual processes Dendrites Axon Multipolar neurons Dendrites Axon Bipolar neurons Dendrites Axon Unipolar neuron Dendrites Figure 12.5 Anaxonic neuron
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Myelin in PNS, Schwann cell spirals repeatedly around a single nerve fiber neurilemma – thick outermost coil of myelin sheath contains nucleus and most of its cytoplasm external to neurilemma is basal lamina and a thin layer of fibrous connective tissue – endoneurium in CNS – oligodendrocytes reaches out to myelinate several nerve fibers in its immediate vicinity nerve fibers in CNS have no neurilemma or endoneurium
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Myelin many Schwann cells or oligodendrocytes are needed to cover one nerve fiber myelin sheath is segmented nodes of Ranvier – gap between segments internodes – myelin covered segments from one gap to the next initial segment – short section of nerve fiber between the axon hillock and the first glial cell trigger zone – the axon hillock and the initial segment play an important role in initiating a nerve signal
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nodes of Ranvier and internodes
Myelin Sheath in PNS Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. One schann cell covers one axon. Schwann cell nucleus Axoplasm Axolemma Neurilemma Figure 12.4c Myelin sheath (c) nodes of Ranvier and internodes 12-13
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Myelination in CNS Figure 12.7b
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. One oligodendrocytes covers multiple axons Oligodendrocyte Myelin Nerve fiber Figure 12.7b (b)
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Myelination in PNS Figure 12.7a Schwann cell Axon Basal lamina
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Schwann cell Axon Basal lamina Endoneurium Nucleus (a) Neurilemma Myelin sheath Figure 12.7a
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Electrical Potentials and Currents
electrical potential – a difference in the concentration of charged particles between one point and another electrical current – a flow of charged particles from one point to another in the body, currents are movement of ions, such as Na+ or K+ through gated channels in the plasma membrane gated channels enable cells to turn electrical currents on and off resting membrane potential (RMP) – is the charge difference across the plasma membrane of a cell -70 mV in a resting, unstimulated neuron negative value means there are more negatively charged particles on the inside of the membrane than on the outside
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Resting Membrane Potential
RMP exists because of unequal electrolyte distribution between extracellular fluid (ECF) and intracellular fluid (ICF) RMP results from the combined effect of three factors: ions diffuse down their concentration gradient through the membrane plasma membrane is selectively permeable and allows some ions to pass easier than others electrical attraction of cations and anions to each other
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Creation of Resting Membrane Potential
potassium ions (K+) found inside the cell K+ is about 40 times as concentrated in the ICF as in the ECF have the greatest influence on RMP leaks out until electrical charge of cytoplasmic anions attracts it back in and equilibrium is reached and net diffusion of K+ stops Sodium ions (Na+) found outside the cell Na+ is about 12 times as concentrated in the ECF as in the ICF resting membrane is much less permeable to Na+ than K+ Large anions in the ICF that keep the RMP negative. Na+/K+ pumps out 3 Na+ for every 2 K+ it brings in works continuously to compensate for Na+ and K+ leakage, and requires great deal of ATP 70% of the energy requirement of the nervous system
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Ionic Basis of Resting Membrane Potential
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. ECF Na+ 145 mEq/L K mEq/L Figure 12.11 K+ channel Na+ channel Na mEq/L K mEq/L Large anions that cannot escape cell ICF Na+ concentrated outside of cell (ECF) K+ concentrated inside cell (ICF)
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Excitation of a Neuron by a Chemical Stimulus
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Dendrites Soma Trigger zone Axon Current ECF Ligand Receptor Plasma membrane of dendrite Na+ ICF Figure 12.12
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Action Potentials action potential – dramatic change produced by voltage-regulated ion gates in the plasma membrane trigger zone where action potential is generated threshold – critical voltage to which local potentials must rise to open the voltage-regulated gates Between -55mV and -50mV when threshold is reached, neuron ‘fires’ and produces an action potential more and more Na+ channels open in in the trigger zone in a positive feedback cycle creating a rapid rise in membrane voltage – spike when rising membrane potential passes 0 mV, Na+ gates are inactivated when all closed, the voltage peaks at about +30 mV membrane now positive on the inside and negative on the outside polarity reversed from RMP - depolarization by the time the voltage peaks, the slow K+ gates are fully open K+ repelled by the positive intracellular fluid now exit the cell their outflow repolarizes the membrane shifts the voltage back to negative numbers returning toward RMP K+ gates stay open longer than the Na+ gates slightly more K+ leaves the cell than Na+ entering negative overshoot – hyperpolarization or afterpotential
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Action Potentials Figure 12.13a
only a thin layer of the cytoplasm next to the cell membrane is affected in reality, very few ions are involved action potential is often called a spike – happens so fast characteristics of action potential versus a local potential follows an all-or-none law if threshold is reached, neuron fires at its maximum voltage if threshold is not reached it does not fire nondecremental - do not get weaker with distance irreversible - once started goes to completion and can not be stopped Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. 4 +35 3 5 Depolarization Repolarization Action potential mV Threshold 2 –55 Local potential 1 7 –70 Resting membrane potential 6 Hyperpolarization Time (a) Figure 12.13a
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Sodium and Potassium Gates
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. K+ Na+ K+ gate Na+ gate 35 35 mV mV 1 Na+ and K+ gates closed 2 Na+ gates open, Na+ enters cell, K+ gates beginning to open –70 –70 Resting membrane potential Depolarization begins Figure 12.14 35 35 mV mV 3 Na+ gates closed, K+ gates fully open, K+ leaves cell 4 Na+ gates closed, K+ gates closing –70 –70 Depolarization ends, repolarization begins Repolarization complete
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The Refractory Period Figure 12.15
during an action potential and for a few milliseconds after, it is difficult or impossible to stimulate that region of a neuron to fire again. refractory period – the period of resistance to stimulation two phases of the refractory period absolute refractory period no stimulus of any strength will trigger AP relative refractory period only especially strong stimulus will trigger new AP refractory period is occurring only at a small patch of the neuron’s membrane at one time other parts of the neuron can be stimulated while the small part is in refractory period Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Absolute refractory period Relative refractory period +35 mV Threshold –55 Resting membrane potential –70 Time Figure 12.15
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Nerve Signal Conduction Unmyelinated Fibers
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. unmyelinated fiber has voltage-regulated ion gates along its entire length and Na+ and K+ gates open and close producing a new action potential all the way down the axon. This slows down the signal being sent. Dendrites Cell body Axon Signal – – – Action potential in progress – – – – – – – – – – – – – – – Refractory membrane – – – – – – – – – – – – – – – Excitable membrane – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – + + Figure 12.16 – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – – + +
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Saltatory Conduction Myelinated Fibers
voltage-gated channels needed for APs fast Na+ diffusion occurs between nodes signal weakens under myelin sheath, but still strong enough to stimulate an action potential at next node saltatory conduction – the nerve signal seems to jump from node to node increasing speed of signal being sent. Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Figure 12.17a Na+ inflow at node generates action potential (slow but nondecremental) Na+ diffuses along inside of axolemma to next node (fast but decremental) Excitation of voltage- regulated gates will generate next action potential here (a)
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Saltatory Conduction Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. + + – – + + + + + + + + – – + + – – – – – – – – – – + + – – – – – – – – + + – – + + + + + + + + + + + + – – + + + + + + – – – – + + – – – – – – – – – – + + – – – – – – + + + + – – + + + + + + + + + + + + – – + + + + – – – – – – + + – – – – – – – – – – + + – – – – + + + + + + – – + + + + Action potential in progress Refractory membrane Excitable membrane (b) Figure 12.17b much faster than conduction in unmyelinated fibers
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Synapses a nerve signal can go no further when it reaches the end of the axon triggers the release of a neurotransmitter stimulates a new wave of electrical activity in the next cell across the synapse electrical synapses do exist advantage of quick transmission no delay for release and binding of neurotransmitter Gap junctions in cardiac and smooth muscle and some neurons disadvantage is they cannot integrate information and make decisions ability reserved for chemical synapses in which neurons communicate by releasing neurotransmitters synapse between two neurons 1st neuron in the signal path is the presynaptic neuron releases neurotransmitter 2nd neuron is postsynaptic neuron responds to neurotransmitter presynaptic neuron may synapse with a dendrite, soma, or axon of postsynaptic neuron neuron can have an enormous number of synapses in cerebellum of brain, one neuron can have as many as 100,000 synapses
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Synaptic Relationships Between Neurons
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Soma Synapse Axon Presynaptic neuron Direction of signal transmission Postsynaptic neuron (a) Axodendritic synapse Axosomatic synapse Figure 12.18 Axoaxonic synapse (b)
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© Omikron/Science Source/Photo Researchers, Inc
Synaptic Knobs Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Axon of presynaptic neuron Synaptic knob Soma of postsynaptic neuron © Omikron/Science Source/Photo Researchers, Inc Figure 12.19
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Structure of a Chemical Synapse
synaptic knob of presynaptic neuron contains synaptic vesicles containing neurotransmitter ready to release neurotransmitter on demand postsynaptic neuron membrane contains proteins that function as receptors and ligand-regulated ion gates
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Structure of a Chemical Synapse
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Microtubules of cytoskeleton Axon of presynaptic neuron Mitochondria Postsynaptic neuron Synaptic knob Synaptic vesicles containing neurotransmitter Synaptic cleft Neurotransmitter receptor Figure 12.20 Neurotransmitter release Postsynaptic neuron presynaptic neurons have synaptic vesicles with neurotransmitter and postsynaptic have receptors and ligand-regulated ion channels
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Neurotransmitters and Related Messengers
more than 100 neurotransmitters have been identified fall into four major categories according to chemical composition acetylcholine in a class by itself formed from acetic acid and choline amino acid neurotransmitters include glycine, glutamate, aspartate, and -aminobutyric acid (GABA) monoamines synthesized from amino acids by removal of the –COOH group retaining the –NH2 (amino) group major monoamines are: epinephrine, norepinephrine, dopamine (catecholamines) histamine and serotonin neuropeptides
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Function of Neurotransmitters at Synapse
they are synthesized by the presynaptic neuron they are released in response to stimulation they bind to specific receptors on the postsynaptic cell they alter the physiology of that cell
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Effects of Neurotransmitters
a given neurotransmitter does not have the same effect everywhere in the body multiple receptor types exist for a particular neurotransmitter 14 receptor types for serotonin receptor governs the effect the neurotransmitter has on the target cell
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Synaptic Transmission
neurotransmitters are diverse in their action some excitatory some inhibitory some the effect depends on what kind of receptor the postsynaptic cell has some open ligand-regulated ion gates some act through second-messenger systems three kinds of synapses with different modes of action excitatory cholinergic synapse inhibitory GABA-ergic synapse excitatory adrenergic synapse synaptic delay – time from the arrival of a signal at the axon terminal of a presynaptic cell to the beginning of an action potential in the postsynaptic cell 0.5 msec for all the complex sequence of events to occur
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Excitatory Cholinergic Synapse
cholinergic synapse – employs acetylcholine (ACh) as its neurotransmitter ACh excites some postsynaptic cells skeletal muscle inhibits others describing excitatory action nerve signal approaching the synapse, opens the voltage-regulated calcium gates in synaptic knob Ca2+ enters the knob triggers exocytosis of synaptic vesicles releasing ACh empty vesicles drop back into the cytoplasm to be refilled with ACh reserve pool of synaptic vesicles move to the active sites and release their ACh ACh diffuses across the synaptic cleft binds to ligand-regulated gates on the postsynaptic neuron gates open allowing Na+ to enter cell and K+ to leave pass in opposite directions through same gate as Na+ enters the cell it spreads out along the inside of the plasma membrane and depolarizes it producing a local potential called the postsynaptic potential if it is strong enough and persistent enough it opens voltage-regulated ion gates in the trigger zone causing the postsynaptic neuron to fire
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Excitatory Cholinergic Synapse
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Presynaptic neuron Presynaptic neuron 3 Ca2+ 1 2 ACh Na+ – – – – 4 – – – + + + + + + + Figure 12.22 5 K+ Postsynaptic neuron
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Cessation of the Signal
mechanisms to turn off stimulation to keep postsynaptic neuron from firing indefinitely neurotransmitter molecule binds to its receptor for only 1 msec or so then dissociates from it if presynaptic cell continues to release neurotransmitter one molecule is quickly replaced by another and the neuron is restimulated stop adding neurotransmitter and get rid of that which is already there stop signals in the presynaptic nerve fiber getting rid of neurotransmitter by: diffusion neurotransmitter escapes the synapse into the nearby ECF astrocytes in CNS absorb it and return it to neurons reuptake synaptic knob reabsorbs amino acids and monoamines by endocytosis break neurotransmitters down with monoamine oxidase (MAO) enzyme some antidepressant drugs work by inhibiting MAO degradation in the synaptic cleft enzyme acetylcholinesterase (AChE) in synaptic cleft degrades ACh into acetate and choline choline reabsorbed by synaptic knob
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Kinds of Neural Circuits
diverging circuit one nerve fiber branches and synapses with several postsynaptic cells one neuron may produce output through hundreds of neurons converging circuit input from many different nerve fibers can be funneled to one neuron or neural pool opposite of diverging circuit reverberating circuits neurons stimulate each other in linear sequence but one cell restimulates the first cell to start the process all over diaphragm and intercostal muscles parallel after-discharge circuits input neuron diverges to stimulate several chains of neurons each chain has a different number of synapses eventually they all reconverge on a single output neuron after-discharge – continued firing after the stimulus stops
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Neural Circuits Figure 12.30 Diverging Converging Input Output Output
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Diverging Converging Input Output Output Input Reverberating Parallel after-discharge Figure 12.30 Input Output Input Output
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