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Physiology of the Nervous System / 215


                                      Na+         Na+        Na+      3Na+
  VetBooks.ir                  Extracellular         Na+                       Na+
                                 space

                                          (b)
                           Cell                                          (a)
                        membrane


                               Cytoplasm                          ADP
                                           K+         K+                ATP
                                                           K+
                                   Large, negatively
                                (c) charged molecules  K+                  2K+

               Figure 11-2.  The neuron’s cell membrane at rest is polarized by the separation of charges. (a) Positively
               charged sodium ions are actively pumped out of the neuron in exchange for potassium ions (in a ratio of
               3Na for 3 K). (b) Potassium ions slowly leak out of the neuron in response to their electrical and chemical
               gradients. (c) Large molecules with many negative charges are trapped inside the cell because they are too
               large to cross the membrane.

                  Signaling between neurons usually
               involves temporarily changing permea­
               bility of the cell membrane to ions. For                    Action potential
               example, when protein channels that allow
               sodium ion to pass through them are                       gNa
               opened, sodium enters the cell in response
               to its chemical and electrical gradients;
               this influx of positive ions depolarizes     Voltage/permeability
               (make less negative) the interior of the                    gK
               neuron. In the receptive zone, this happens
               in localized regions of the cell membrane
               in response to the signals  from other
               neurons in proportion to the strength of        Resting potential
               the  signal,  a  so‐called  graded potential.
               These kinds of depolarizations spread           1 ms
               only a small distance from the site at which
               they are generated, and become weaker    Figure 11-3.  Change in sodium and potassium
               the further they spread.                 permeability during neuron action potential. gNa,
                                                        conductance for sodium; gK, conductance for
                  At the junction of the axon with the cell   potassium. Early influx of sodium (gNa) drives the
               body, the  axon hillock, depolarization   steepest part of the depolarization of the action
               has a different effect, however. It is here that   potential. Sodium flow begins to decline and
               the action potential, a large, self‐propagating   the  conductance for potassium (gK) increases.
               wave of depolarization, is initiated.    Outflow of potassium is a significant contributor
                  The membrane of the axon hillock and   to the return of the membrane to its resting,
               axon contains voltage‐gated sodium chan-  hyperpolarized state.
               nels. These channels are closed at normal
               resting membrane potentials but rapidly   depolarization (Fig. 11‐3). At the peak of the
               open when the membrane potential is      action potential (maximal  depolarization),
               depolarized to threshold voltage. This   the sodium channels close and additional
               opening results in the inward movement   voltage‐gated potassium channels are
               of sodium and a large, rapid membrane    opened. The closing of the sodium channels
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