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Chapter 10 Functional Organization of Nervous Tissue Neuron Network Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Functions of the Nervous System • • The master controlling and communicating system of the body Functions 1. Sensory input: detects external and internal stimuli 2. Integration: processes and responds to sensory input 3. Control of Muscles and Glands 4. Homeostasis is maintained by regulating other systems 5. Center for Mental Activities Fig. 10.1 Parts of the Nervous System • Two anatomical divisions – Central Nervous System (CNS) • • – Peripheral Nervous System (PNS) • • • Brain and spinal cord Encased in bone Nervous tissue outside of the CNS Consists of sensory receptors and nerves The anatomical divisions perform different functions – – PNS detects stimuli and transmits information to the CNS and receives information from the CNS CNS processes, integrates, stores, and responds to information from the PNS Parts of the Nervous System • PNS has two divisions – – Sensory division transmits action potentials from sensory receptors to the CNS Motor division carries action potentials away from the CNS in cranial or spinal nerves (two subdivisions) • • Somatic nervous system innervates skeletal muscle Autonomic nervous system (ANS) innervates cardiac muscle, smooth muscle, and glands (three subdivisions) – – – Sympathetic division is most active during physical activity (fight or flight division) Parasympathetic division regulates resting functions (rest and digest division) Enteric nervous system controls the digestive system Parts of the Nervous System Central Nervous System (CNS) •Brain and Spinal Cord Peripheral Nervous System (PNS) •Nervous tissue outside the CNS •Sensory receptors and nerves Sensory Division •Transmits action potentials from sensory receptors to the CNS Motor Division •Carries action potentials away from the CNS in cranial nerves or spinal nerves Sympathetic Division Most active during physical activity Parasympathetic Division Regulates resting functions Enteric Nervous System Controls the Digestive System Autonomic Nervous System (ANS) •Innervates cardiac muscle, smooth muscle, and glands Somatic Nervous System •Innervates skeletal muscle Fig. 10.2 Cells of the Nervous System • The two principal cell types of the nervous system are: – Neurons: excitable cells that transmit electrical signals – Non-neural cells (Glial cells): cells that surround neurons. Account for over half of the brain’s weight • Less than 20% is extracellular space Neurons • Receive stimuli and transmit action potentials • Have three components: – The cell body (soma) is the primary site of protein synthesis – Dendrites are short, branched cytoplasmic extensions of the cell body that usually conduct electric signals toward the cell body – An axon is a cytoplasmic extension of the cell body that transmits action potentials to other cells Fig. 10.3 Neuron Structure • Cell Body (Soma) – Contains the nucleus and a nucleolus – Nissl substance is an aggregate of rough ER and free ribosomes • Primary site of protein synthesis – Golgi apparatus, mitochondria, and other organelles are present – Has no centrioles (hence its amitotic nature) – Clusters of cell bodies in the CNS are called nuclei and in the PNS ganglia Fig. 10.3 Neuron Structure • Axons (Nerve Fibers) – Trigger zone is the part of the neuron where the axon originates • Action potential is generated from the trigger zone – Slender processes of uniform diameter and may vary in length from a few millimeters to more than a meter – Usually, there is only one unbranched axon per neuron • Rare branches, if present, are called collateral axons – Presynaptic terminal: branched terminus of an axon (10,000 or more) – Synapse: junction between a nerve cell and another cell – Bundles of processes are called nerve tracts in the CNS and nerves in the PNS Fig. 10.3 Types of Neurons • Multipolar neurons have several dendrites and a single axon – Interneurons and motor neurons • Bipolar neurons have a single axon and dendrite – Components of sensory organs • Unipolar neurons have a single axon – Most sensory neurons Fig. 10.4 Glial Cells • Glial Cells (Supporting Cells): – – – – Provide a supportive scaffolding for neurons Segregate and insulate neurons Guide young neurons to the proper connections Promote health and growth • Glial Cells of the CNS – – – – Astrocytes Microglial Ependymal cells Oligodendrocytes • Glial Cells of the PNS – Satellite cells – Schwann cells Glial Cells of the CNS • Astrocytes – Most abundant, versatile, and highly branched – They cling to neurons and their synaptic endings, and cover capillaries – Functions: • Support and brace neurons and blood vessels – Anchor neurons to their nutrient supplies • Influence the functioning of the blood-brain barrier • Guide migration of young neurons • Process substances – mopping up leaked potassium ions – recycling neurotransmitters • Isolate damaged tissue and limit the spread of inflammation Astrocytes Fig. 10.5 Glial Cells of the CNS • Ependymal cells: range in shape from squamous to columnar and many are ciliated – They line the ventricles of the brain and the central canal of the spinal cord – Some are specialized (choroid plexuses) to produce cerebrospinal fluid (CSF) – Help to circulate CSF using their cilia Fig. 10.6 Glial Cells of the CNS • Microglia – Small, ovoid cells with spiny processes – Phagocytes that monitor the health of neurons Fig. 10.7 Glial Cells of the CNS • Oligodendrocytes: form myelin sheaths around the axons of several CNS neurons Fig. 10.8 Glial Cells of the PNS • Schwann cells: form a myelin sheath around part of the axon of a PNS neuron • Satellite cells: support and nourish neuron cell bodies within ganglia Fig. 10.9 Myelinated and Unmyelinated Axons • Myelinated axons – Plasma membrane of Schwann cells or Oligodendrocytes repeatedly wraps around a segment of an axon to form the myelin sheath – Myelin is a whitish, fatty (protein-lipid), segmented sheath around most long axons – It functions to: • Protect the axon • Electrically insulate fibers from one another • Increase the speed of nerve impulse transmission • Node of Ranvier – Gaps in the myelin sheath Fig. 10.10 Myelinated and Unmyelinated Axons • Unmyelinated axons – Rest in invaginations of Schwann cell (PNS) or Oligodendrocytes (CNS) – Conduct action potentials slowly Fig. 10.10 Fig. 10.10 Organization of Nervous Tissue • Nervous tissue can be grouped into white matter and gray matter – White matter • Consists of myelinated axons • Propagates action potentials • Forms nerve tracts in the CNS and nerves in the PNS – Gray Matter • Collections of neuron cell bodies or unmyelinated axons • Forms cortex and nuclei in the CNS and ganglia in the PNS • Axons synapse with neuron cell bodies, which are functionally the site of integration in the nervous system Electric Signals • Electric signals produced by cells are called action potentials – When action potentials are received from sensory cells it can result in the sensations of sight, hearing, and touch – Complex mental activities, such as conscious thought, memory, and emotions, result from action potentials – Contraction of muscles and the secretion of certain glands occur in response to action potentials • Electrical properties of cells result from – Ionic concentration differences across the plasma membrane – Permeability characteristics of the plasma membrane Electric Signals • Concentration Differences Across the Plasma Membrane – – Sodium ions (Na+), calcium ions (Ca2+), and chloride ions (Cl-) are in much greater concentration outside the cell than inside Potassium ions (K+) and negatively charged molecules, such as proteins, are in much greater concentration inside the cell than outside • Negatively charged proteins are synthesized inside the cell and cannot diffuse out of it Electric Signals • Concentration gradients of ions result mainly from 1. The Na+-K+ pump • Moves ions by active transport • Potassium ions are moved into the cell, and Na+ are moved out of it 2. Permeability characteristics of the plasma membrane are determined by • Leak channels (always open) – Potassium ion leak channels are more numerous than Na+ leak channels; thus, the plasma membrane is more permeable to K+ than to Na+ when at rest • Gated ion channels – Include ligand-gated ion channels, voltage-gated ion channels, and other gated ion channels Electric Signals • Gated Ion Channels – Open and close in response to stimuli • Ligand-gated ion channels – Open or close with the binding of a specific ligand (neurotransmitter) » Ligand is a molecule that binds to a receptor » A receptor is a protein or glycoprotein that has a receptor site to which a ligand can bind – Common in tissues such as nervous and muscle tissue, as well as glands • Voltage-gated ion channels – Open and close in response to small voltage changes across the plasma membrane – Common in tissues such as nervous and muscle tissues • Other gated ion channels – Open and close in response to physical deformation of receptors – Touch receptors (mechanical stimulation) and temperature receptors (temperature changes) of the skin Electric Signals • Establishing the Resting Membrane Potential – Resting membrane potential • Charge difference across the plasma membrane when the cell is not being stimulated • The inside of the cell is negatively charged, compared with the outside of the cell – Due mainly to the tendency of positively charged K+ to diffuse out of the cell – Opposed by the negative charge that develops inside the plasma membrane Electric Signals • Changing the Resting Membrane Potential – Depolarization is a decrease in the resting membrane potential caused by • • • • A decrease in the K+ concentration gradient A decrease in membrane permeability to K+ An increase in membrane permeability to Na+ or Ca2+ A decrease in extracellular Ca2+ concentrations – Hyperpolarization is an increase in the resting membrane potential caused by • • • • • An increase in the K+ concentration gradient An increase in membrane permeability to K+ An increase in membrane permeability to ClA decrease in membrane permeability to Na+ An increase in extracellular Ca2+ concentrations Graded Potentials • Are small changes in the resting membrane potential • Confined to a small area of the plasma membrane – An increase in membrane permeability to Na+ can cause graded depolarization – An increase in membrane permeability to K+ or Cl- can result in graded hyperpolarization • Decreases in magnitude as the distance from the stimulation increases • The term graded potential is used because a stronger stimulus produces a greater potential change than a weaker stimulus Fig. 10.15 • Graded potentials can summate, or add together Action Potentials • Are larger changes in the resting membrane potential that spread over the entire surface of the cell – Occurs when a graded potential causes depolarization of the plasma membrane to a level called threshold – Occur in an all-or-none fashion and are of the same magnitude, no matter how strong the stimulus – Occurs in three phases • Depolarization phase • Repolarization phase • Afterpotential Action Potentials • Depolarization Phase – Inside of the membrane becomes more positive – Na+ diffuses into the cell through voltage-gated ion channels • Repolarization Phase – Return of the membrane potential toward the resting membrane potential – Voltage-gated Na+ channels close – Voltage-gated K+ channels open and K+ diffuses out of the cell • Afterpotential – Brief period of hyperpolarization following repolarization Fig. 10.16 Refractory Periods • Absolute refractory period – Time during an action potential when a second stimulus (no matter how strong) cannot initiate another action potential • Relative refractory period – Time during which a stronger-than-threshold stimulus can evoke another action potential Action Potential Frequency • The number of action potentials produced per unit of time in response to stimuli – It is directly proportional to stimulus strength and to the size of the graded potential • Subthreshold stimulus: graded potential • Threshold stimulus: a single action potential • Submaximal stimulus: action potential frequency increases as the strength of the stimulus increases • Maximal or a supramaximal stimulus: produces a maximum frequency of action potentials Propagation of Action Potentials • An action potential generates ionic currents – Currents stimulate voltage-gated Na+ channels in adjacent regions of the plasma membrane to open – Producing new action potentials • Reversal of the direction of action potential propagation is prevented by the absolute refractory period • Occurs most rapidly in myelinated, largediameter axons Propagation of Action Potentials • In an unmyelinated axon, action potentials are generated immediately adjacent to previous action potentials • In a myelinated axon, action potentials are generated at successive Nodes of Ranvier Fig. 10.21 The Synapse • The synapse is the junction between two cells where communication takes place – Presynaptic cell: transmits signal towards a synapse – Postsynaptic cell: receives the signal • Two types of synapses – Electrical synapse – Chemical synapse Electrical Synapses • Gap junctions in which tubular proteins called connexons allow ionic currents to move between cells • An action potential in one cell generates an ionic current that causes an action potential in an adjacent cell • Action potentials are conducted rapidly between cells allowing for synchronized activity • Common in cardiac muscle and in many types of smooth muscle where coordinated contractions are essential Chemical Synapses • Have three anatomical components – The enlarged ends of the axon are the presynaptic terminals containing synaptic vesicles – The postsynaptic membranes contain receptors for the neurotransmitter – The synaptic cleft, a space, separates the presynaptic and postsynaptic membrane Fig. 10.22 Chemical Synapse Activity 1. Action potentials arriving at the presynaptic terminal cause voltagegated Ca2+ channels to open 2. Calcium ions diffuse into the cell and cause synaptic vesicles to release neurotransmitters 3. Neurotransmitters diffuse from the presynaptic terminal across the synaptic cleft 4. Neurotransmitters combine with their receptor sites and cause ligand-gated ion channels to open. Ions diffuse into the cell (shown) or out of the cell (not shown) and cause a change in membrane potential Fig. 10.22 Chemical Synapse Activity • The effect of the neurotransmitter on the postsynaptic membrane is stopped in two ways – The neurotransmitter is broken down by an enzyme – The neurotransmitter is taken up by the presynaptic terminal • A list of neurotransmitters and neuromodulators is presented in Table 10.4 – Neuromodulators are substances released from neurons that can presynaptically or postsynaptically influence the likelihood that an action potential will be generated Chemical Synapse Activity • Excitatory and inhibitory postsynaptic potentials – An excitatory postsynaptic potential (EPSP) is a depolarizing graded potential of the postsynaptic membrane – An inhibitory postsynaptic potential (IPSP) is a hyperpolarizing graded potential of the postsynaptic membrane – Presynaptic inhibition decreases neurotransmitter release – Presynaptic facilitation increases neurotransmitter release Spatial and Temporal Summation • Presynaptic action potentials through neurotransmitters produce graded potentials in postsynaptic neurons. The graded potential can summate to produce an action potential at the trigger zone – Spatial summation occurs when two or more presynaptic terminals simultaneously stimulate a postsynaptic neuron – Temporal summation occurs when two or more action potentials arrive in succession at a single presynaptic terminal • Inhibitory and excitatory presynaptic neurons can converge on a postsynaptic neuron – An action potential is produced at the trigger zone when the graded potential is produced as a result of the sum of the EPSPs and IPSPs reaching threshold Neuronal Pathways and Circuits • Convergent pathways have many neurons synapsing with a few neurons • Divergent pathways have a few neurons synapsing with many neurons • Oscillating circuits have collateral branches of postsynaptic neurons synapsing with presynaptic neurons