by Talib F. Abbas In fact, the word glia is Greek for glue. However, today theses cells are recognized for their role in communication within the CNS in partnership with neurons. There are two major types of glial cells in the vertebrate nervous system: microglia and macrogli. Microglia are scavenger cells that resemble tissue macrophages. Macroglia: oligodendrocytes, Schwann cells, and astrocytes. Fibrous astrocytes, which contain many intermediate filaments, are found primarily in white matter. Protoplasmic astrocytes are found in gray matter. Electrical potentials exist across the membranes of virtually all cells of the body. In addition, some cells, such as nerve and muscle cells, are capable of generating rapidly changing electrochemical impulses at their membranes, and these impulses are used to transmit signals along the nerve or muscle membranes. In other types of cells, such as glandular cells, macrophages, and ciliated cells, local changes in membrane potentials also activate many of the cells’ functions. “Diffusion Potential” Caused by an Ion Concentration Difference on the Two Sides of the Membrane. In , the potassium concentration is great inside a nerve fiber membrane but very low outside the membrane. Let us assume that the membrane in this instance is permeable to the potassium ions but not to any other ions. Because of the large potassium concentration gradient from inside toward outside, there is a strong tendency for extra numbers of potassium ions to diffuse outward through the membrane. As they do so, they carry positive electrical charges to the outside, thus creating electropositivity outside the membrane and electronegativity inside because of negative anions that remain behind and do not diffuse outward with the potassium. [K+x] > [K+y], that is, more osmotically active particles are on side X than on side Y. Donnan and Gibbs showed that in the presence of a nondiffusible ion. [K+x]/ [K+y] =[Cl–y]/ [Cl–x] Cross-multiplying, [K+x] + [Cl–x] = [K+y] + [Cl–y] This is the Gibbs–Donnan equation. It holds for any pair of cations and anions of the same valence. The interior of the cell is negative relative to the exterior, and chloride ions are pushed out of the cell along this electrical gradient. An equilibrium is reached between Cl– influx and Cl– . ECl = RT/ FZCl ln [Clo–][Cli–] where ECl = equilibrium potential for Cl– R = gas constant T = absolute temperature F = the faraday (number of coulombs per mole of charge). ZCl = valence of Cl– (–1) [Clo–] = Cl– concentration outside the cell [Cli–] = Cl– concentration inside the cell the concentration gradient is outward and the electrical gradient inward. In mammalian spinal motor neurons, EK is –90 mV. Because the resting membrane potential is –70 mV, there is somewhat more K+ in the neurons than can be accounted for by the electrical and chemical gradients. The situation for Na+ is quite different from that for K+ and Cl– . The direction of the chemical gradient for Na+ is inward, to the area where it is in lesser concentration, and the electrical gradient is in the same direction. ENa is +60 mV Because neither EK nor ENa is equal to the membrane potential, one would expect the cell to gradually gain Na+ and lose K+ if only passive electrical and chemical forces were acting across the membrane. “uphill” against an electrical or pressure gradient), the process is called active transport. some energy source must cause excess movement of potassium ions to the inside of cells and excess movement of sodium ions to the outside of cells. The basic mechanism for transport of a substance through a cellular sheet shows that the epithelial cells are connected together tightly at the luminal pole by means of junctions called “kisses.” The brush border on the luminal surfaces of the cells is permeable to both sodium ions and water. The resting membrane potential of large nerve fibers when not transmitting nerve signals is about –90 millivolts. That is, the potential inside the fiber is 90 millivolts more negative than the potential in the extracellular fluid on the outside of the fiber. all cell membranes of the body have a powerful Na+-K+ that continually pumps sodium ions to the outside of the cell and potassium ions to the inside, this is an electrogenic pump because more positive charges are pumped to the outside than to the inside (three Na+ ions to the outside for each two K+ ions to the inside), leaving a net deficit of positive ions on the inside; this causes a negative potential inside the cell membrane. Nerve signals are transmitted by action potentials, which are rapid changes in the membrane potential that spread rapidly along the nerve fiber membrane. The cell membranes of nerves, like those of other cells, contain many different types of ion channels. Some of these are voltage- gated and others are ligand-gated. In response to a depolarizing stimulus, some of the voltage-gated Na+ channels become active, and when the threshold potential is reached, the voltagegated Na+channels overwhelm the K+ and other channels and an action potential results (a positive feedbackloop) . The membrane potential moves toward the equilibrium potential for Na+(+60 mV) but does not reach it during the action potential, primarily because the increase in Na+ conductance is short-lived. The Na+ channels rapidly enter aclosed state called the inactivated state. In addition, the direction of the electrical gradient for Na+ is reversed during the overshoot because the membrane potential is reversed, and this limits Na+ influx. A third factor producing repolarization is the opening of voltagegated K+ channels. The slow return of the K+ channels to the closed state also explains the after-hyperpolarization, followed by a return to the resting membrane potential. Thus, voltagegated K + channels bring the action potential to an end and causeclosure of their gates through anegative feedback process. Two types of physicochemical disturbances are produced: local, nonpropagated potentials called, depending on their location, synaptic, generator, Or electrotonic potentials; and propagated potentials, the action potentials (or nerve impulses ). The impulse is normally transmitted (conducted) along the axon to its termination. Nerves are not “telephone wires” that transmit impulses passively; conduction of nerve impulses, although rapid, is much slower than that of electricity. The transmission of each action potential along a nerve fiber reduces very slightly the concentration differences of sodium and potassium inside and outside the membrane, because sodium ions diffuse to the inside during depolarization and potassium ions diffuse to the outside during repolarization. For a single action potential, this effect is so minute that it cannot be measured. Indeed, 100,000 to 50 million impulses can be transmitted by large nerve fibers before the concentration differences reach the point that action potential conduction ceases. Even so, with time, it becomes necessary to re-establish the sodium and potassium membrane concentration differences. This is achieved by action of the Na+-K+ pump in the same way as described previously in the chapter for the original establishment of the resting potential. Epineurium all-or-none action potentials. maximal stimulus Neurotrophins
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