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350       Tissues and organs



             Resting potential and action                     uli (or more rarely electrical stimuli). Binding
             potential                                        of a neurotransmitter to an ionotropic recep-
                                                              tor results in a brief local increase in the
                                                              membrane potential from –60 mV to about
             A. Resting potential
                                                              +30 mV. Although the membrane potential
             A characteristic property of living cells is the  quickly returns to the initial value within a
             uneven distribution of positively and nega-      few milliseconds (ms) at its site of origin, the
             tively charged ions on the inside and outside    depolarization is propagated because neigh-
             of the plasma membrane. This gives rise to a     boring membrane areas are activated during
             membrane potential (see p. 126)—i. e., there is  this time period.
             electrical voltage between the two sides of         [1] The process starts with the opening of
                                                                               +
             the membrane, which can only balance out         voltage-gated Na channels (see p. 222). Due
             when ion channels allow the unevenly distrib-    to their high equilibrium potential (see A), Na +
             uted ions to move.                               ions flow into the cell and reverse the local
                At rest, the membrane potential in most       membrane potential (depolarization).
                                                                            +
             cells is –60 to –90 mV. It mainly arises from       [2] The Na channels immediately close
                                 +
             the activity of Na /K  +  transporting ATPase    again, so that the inflow of positive charges
                  +
                     +
             (“Na /K ATPase”), which occurs on practically    is only very brief.
             all animal cells. Using up ATP, this P-type         [3] Due to the increase in the membrane
                                                     +
             enzyme (see p. 220) “pumps” three Na ions        potential, voltage-dependent K   +  channels
                                                                                                         +
                                                                          +
                                                    +
             out of the cell in exchange for two K ions.      open and K ions flow out. In addition, Na /
                                                               +
                           +
             Some of the K ions, following the concentra-     K ATPase (see A) pumps the Na+ ions that
             tion gradient, leave the cell again through      have entered back out again. This leads to
             potassium channels. As the protein anions        repolarization of the membrane.
             that predominate inside the cell cannot fol-        [4] The two processes briefly lead to the
                                          –
             lowthem, and inflowofCl ions from the            charge even falling below the resting poten-
                                                                                            +
             outside is not possible, the result is an excess  tial (hyperpolarization). The K channels also
             of positive charges outside the cell, while      close after a few milliseconds. The nerve cell
             anions predominate inside it.                    is then ready for re-stimulation.
                An equilibrium potential exists for each of      Generally, it is always only a very small
             the ions involved. This is the value of the      part of the membrane that is depolarized dur-
             membrane potential at which there is no net      ing an action potential. The process can there-
             inflow or outflow of the ions concerned. For     fore be repeated again after a short refractory
               +
             K ions, the resting potential lies in the range  period, when the nerve cell is stimulated
                                                     +
             of the membrane potential, while for Na ions     again. Conduction of the action potential on
             it is much higher at +70 mV. At the first op-    the surface of the nerve cell is based on the
                           +
             portunity, Na ions will therefore spontane-      fact that the local increase in the membrane
             ously flow into the cell. The occurrence of      potential causes neighboring voltage-gated
             action potentials is based on this (see B).      ion channels to open, so that the membrane
                Nerve cell membranes contain ion chan-        stimulation spreads over the whole cell in the
                               –
                                        2+
                        +
                            +
             nels for Na ,K ,Cl ,and Ca . These channels      form of a depolarization wave.
             are usually closed and open only briefly to let
             ions pass through. They can be divided into
             channels that are regulated by membrane po-
                                                   +
             tentials (“voltage-gated”—e. g., fast Na chan-
             nels; see p. 222) and those regulated by
             ligands (“ligand-gated”—e. g., nicotinic acetyl-
             choline receptors; see p. 222).


             B. Action potential
             Action potentials are special signals that are
             used to transmit information in the nervous
             system. They are triggered by chemical stim-


           Koolman, Color Atlas of Biochemistry, 2nd edition © 2005 Thieme
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