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The brain is an intricate network, consisting of approximately 200 billion neurons and more than a trillion glia Cells. Each neuron receives and combines multiple inputs to determine, whether to transmit an action potential to the next target in its network (Neuron, Muscle, Gland, or Organ). Three Main Components of the Central Nervous System Neurons - basic signaling units of the nervous system. Glial cells - or Neuroglia (nerve glue) are 10 times more common and account for more than half the brain’s volume. Provide support functions both chemically and structurally. Vasculature The Neuron Doctrine - Cajal Connectional Specificity - cells are separate, and connections between cells are not random but pass information through specific pathways. Dynamic Polarization - some parts of the neuron are specialized for taking information in, while others are specialized for sending it out. Adapted from Kandel, E.R. Schwartz, J.H., and Jessell, T.M. (Eds.), Principles of Neural Science, 3rd edition. Norwalk, Connecticut: Appleton & Lange, 1991. Copyright © 1991 by Appleton & Lange. Variety of synapse possibilities -Axodendritic -Axoaxonic -Axosomatic -Dendrodendritic -Asymmetric (excitatory, glutamate or aspartate) -or Symmetric (inhibitory, GABA) Excitatory Neurons -Pyramidal Cells, projection -Spiny Stellate Cells, interneurons Inhibitory Neurons -Basket Cells, to somas and proximal dendrites, or shaft of apical dendrite, predominate layers III and V, horizontal inhibition -Chandelier Cells, synapse exclusively on the axon hillock of pyramidal cells, also known as axoaxonic cells, layer III, so may control corticocortical connections -Double Bouquet Cells, layers II, III, and V, association cortex, spines, columnar, limited horizontal spread Summary - Neurons -all neurological processes are dependent on complex cell-cell interactions between single neurons and/or groups of related neurons. -Neurons can be described according to their size, shape, neurochemical characteristics, location, and connectivity. -Structure and function are intimately related. W. W. Norton Netter, F.H., The CIBA Collection of Medical Illustrations. Vol I: Nervous System, Part 1: Anatomy and Physiology. Summit, NJ: CIBA Pharmaceutical Company, 1983. Adapted with permission of Novartis, formerly CIBA. Subcellular Organization: Organelles and Their Functions W. W. Norton SubCellular Anatomy • Cell body (Soma) - the nucleus does not divide after maturity. • Nucleus -enveloped compartment containing the genetic material of the cell. • Nucleolus - site of gene transcription. • Rough Endoplasmic Reticulum - extends throughout the cell for protein synthesis. • Ribosomes - site of protein production. • Smooth Endoplasmic Reticulum - tubular structure throughout the cell including nerve endings, site of the calcium pump and source of intracellular calcium, transportation of proteins. • Golgi Apparatus or Complex - important for protein modification (e.g., GlycoProteins) and as a source of vesicles, i.e., packaging. • Mitochondria - dispersed throughout the soma are part of oxidative phosphorylation coupling with electron transport, for ATP (energy) production; and may aid calcium concentrations at synapses, by high capacity low affinity transport. • Axons - contain output microtubules (25 nm) composed of tubulin, and Neurofilaments (10 nm); important for axoplasmic transport (fast - 400 mm/day). • Synaptic Vesicles - Neurotransmitter storage and release location. • Dendrites - input microtubules important for plasticity of synapse formation. • Synaptic Matrix - thickened cell membrane in region of synaptic transmission. Consists of extracellular matrix proteins. Slow axonal transport Fast axonal transport Neurophysiology Two types of synapses: Electrical (Gap Junctions) Chemical W. W. Norton Contributors to the resting membrane potential: 1) Selective membrane permeability (ion channels, negative proteins) 2) Diffusion (equilibrium potential) 3) Electrostatic Forces (equilibrium potential) 4) Sodium-potassium pump (3 sodium ions out, 2 potassium in) W. W. Norton -70 mV 0 mV W. W. Norton Contributors to the resting membrane potential: 1) Selective membrane permeability (ion channels, negative proteins) 2) Diffusion (equilibrium potential) 3) Electrostatic Forces (equilibrium potential) 4) Sodium-potassium pump (3 sodium ions out, 2 potassium in) absolute refractory period versus refractory period 2.16A Adapted from Kandel, E.R. Schwartz, J.H., and Jessell, T.M. (Eds.), Principles of Neural Science, 3rd edition. Norwalk, Connecticut: Appleton & Lange, 1991. Copyright © 1991 by Appleton & Lange. 2.16B Adapted from Kuffler, S., and Nicholls, J., From Neuron to Brain. Sunderland, MA: Sinauer Associates, 1976. W. W. Norton W. W. Norton Trigeminal Ganglion Cell: this is about 2 seconds of activity that was recorded from a ganglion cell after the maxillary (upper) incisor tooth of an anesthetized rat was tapped 5 times. Listen for 5 distinct "bursts" of action potentials. Trigeminal Ganglion Cell: this is about 2 seconds of activity that was recorded from a rat ganglion cell after a single whisker (vibrissa) was moved and held in position. Listen for the rapid steady burst of action potentials. This neuron was firing about 100 action potentials every second. Adapted from Kandel, E.R. Schwartz, J.H., and Jessell, T.M. (Eds.), Principles of Neural Science, 3rd edition. Norwalk, Connecticut: Appleton & Lange, 1991. Copyright © 1991 by Appleton & Lange. Ion Channels - - Ion channels are polypeptides, formed by amino acids, and any two amino acids can form a peptide, but many come to gather to form small and large proteins. Two main types, active transporters (Na+/K+ pump), and ion channels (gated and non-gated) Amino acid sequence primary Strings of amino acids coiled into helical shape - secondary Folding of helix can give rise to globular proteins - tertiary Ion channel - complex 3-D structure made of globular proteins - quaternary W. W. Norton Doyle, D., Cabral, J., Pfuetzner, R., Kuo, A., Gulbis, J., Cohen, S., Chait, B., and MacKinnon, R., (1998). The structure of the potassium channel; Molecular basis of K+ conduction and selectivity; Science 280 (5360); 69-77. 2.26 Doyle, D., Cabral, J., Pfuetzner, R., Kuo, A., Gulbis, J., Cohen, S., Chait, B., and MacKinnon, R., (1998). The structure of the potassium channel; Molecular basis of K?? conduction and selectivity; Science 280 (5360); 69-77. W. W. Norton Receptors are specialized ion channels and can be directly or indirectly coupled Direct - allows ions in, ionotropic Indirect - secondary affect using Gproteins, second messengers, intracellular signaling, metabotropic receptors W. W. Norton Adapted from Kandel, E.R. Schwartz, J.H., and Jessell, T.M. (Eds.), Principles of Neural Science, 3rd edition. Norwalk, Connecticut: Appleton & Lange, 1991. Copyright © 1991 by Appleton & Lange. W. W. Norton W. W. Norton W. W. Norton Proteins SG synaptogamin SB synaptobrevin ST - syntaxin S25 - SNAP-25 Adapted from Purves D., Augustine, G., and Fitzpatrick, D. (2001). Neuroscience, 2nd edition. Sunderland, MA: Sinaur Associates. SG synaptogamin, green SB synaptobrevin, blue ST - syntaxin, red S25 - SNAP-25, purple Adapted from Purves D., Augustine, G., and Fitzpatrick, D. (2001). Neuroscience, 2nd edition. Sunderland, MA: Sinaur Associates. SG synaptogamin, green SB synaptobrevin, blue ST - syntaxin, red S25 - SNAP-25, purple Adapted from Purves D., Augustine, G., and Fitzpatrick, D. (2001). Neuroscience, 2nd edition. Sunderland, MA: Sinaur Associates. QuickTime™ and a TIFF (LZW) decompressor are needed to see this picture. Cyclic Nucleotide Metabolism - cAMP SIGMA-ALDRICH