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                     16   PA R T  I / Anatomy and Physiology

                     there is a continual movement toward equilibrium. When equi-  (positively charged) are more concentrated in the sarcoplasm than
                     librium is reached, forces driving ion movement are balanced, and  in the extracellular space. To balance the force of the concentra-
                     there is no additional net change in the ion distribution. The  tion difference, the inside of the myocardial cell would need to be
                     Nernst equation, discussed later, is useful in understanding the re-  approximately  94 mV compared with outside of the membrane.
                     lationship between electrical and diffusional forces driving ion  That charge,  94 mV, is known as the potassium ion equilibrium
                     movement. It is useful to remember that the permeability proper-  potential or the Nernst potential for potassium ion. The resting
                     ties of the living membrane change continually.    myocardial membrane is permeable to potassium ions. The large
                                                                        concentration gradient is maintained because the actual voltage
                     Diffusional Force                                  across the membrane between activation cycles, approximately
                     Particles in solution move, or diffuse, from an area of higher con-  –90 mV (inside negative), is close to the Nernst potential for
                     centration to an area of lower concentration. In the case of un-  potassium ions in the myocardial cell, and thus nearly sufficient to
                     charged, soluble molecules, diffusion proceeds until there is a uni-  retain potassium ions within the cell. The slow outward trickle of
                     form distribution of the molecules within the solution. The  potassium ions is corrected by a membrane pump that moves
                     solution is then said to be in equilibrium. At equilibrium, there is  potassium ions back into the cell (and moves sodium ions out of
                     still particle movement within the solution, but no net change in  the cell). If the resting potential were –94 mV, there would be no
                     overall particle distribution. Charged particles also diffuse. The  net potassium ion movement.
                     diffusion of charged particles is influenced not only by the con-  The following illustrates the Nernst equation calculation of the
                     centration gradient but also by the electrical field.  equilibrium potential for potassium ion:
                     Electrical Force and Current                                             RT   [K ] o

                     Like charges repel, and opposite charges attract. Positively charged  E K      Ln
                                                                                             FZ K  [K ] i
                     particles flow toward negatively charged particles and regions;
                     similarly, negatively charged particles are attracted to positive ions  where E K   equilibrium potential for K
                     and regions. The electrical (or electromotive) force difference be-  R   gas constant
                     tween regions is called the potential difference and is expressed in  T   absolute temperature
                     volts (1 mV   0.001 V). The net flow of charges is called current  F   the Faraday (number of coulombs per mole of charge)
                    (measured in amperes). Resistance is the opposition to the flow of  Z K    the valence of K ( 1)

                    current, measured in ohms. Ohm’s law (electromotive force    [K ] o   K concentration outside the cell (e.g., 4 mM)


                     current   resistance) describes the relation among current, volt-  [K ] i   K concentration inside the cell (e.g., 155 mM)


                     age, and resistance.                                  Converting from the natural log to the base 10 log and replac-
                       When charged particles have different concentrations in the  ing the constants measured at 37°C with numeric values, the
                     solutions separated by a cell membrane, and some of the particles  equation becomes approximately as follows:
                     are able to permeate the membrane and others are not, an electri-

                     cal force is established. This force influences the distribution of all        [K ] o
                     other charged particles. The potential difference across biologic   E K   61 log 10
                                                                                                   [K ] i
                     membranes is described by comparing the interior of the cell with
                     the external solution. In the typical quiescent or resting myocar-         4
                     dial cell, the potential difference is –70 to –90 mV; that is, the cell  E K   61 log 10   155    97 mV
                     interior is negative with respect to the exterior. When positively  According to the Nernst equation, the higher the potassium
                     charged ions move from the extracellular fluid to the intracellular  ion concentration in the external solution, the more depolarized
                     fluid, the current is said to be inward. With inward current, the  is the potassium equilibrium potential. If the resting membrane
                    cell interior becomes less negative, that is, it depolarizes. When  were highly permeable to potassium ion, then the higher the ex-
                    positively charged ions flow into the extracellular space from the  ternal potassium ion concentration, the more depolarized would
                    interior of the cell, the current is said to be outward; the cell re-  be the resting potential. If one were to perform such an experi-
                    polarizes or hyperpolarizes. Movement of negatively charged ions  ment, placing an intact muscle cell in a dish bathed in solutions
                    in one direction is electrically equivalent to an opposite-directed  with varying potassium ion concentrations as the potassium ion
                    movement of positively charged ions. Thus, the inward movement  concentration in the external solution is raised, the membrane be-
                    of an anion such as chloride is called an outward current. This,  comes more depolarized. When the concentration of potassium
                    too, causes repolarization or hyperpolarization.
                                                                        ion in the extracellular fluid becomes equal to the concentration
                       Nernst Equation Calculation of Equilibrium Potential  in the intracellular fluid, the membrane potential is 0 mV.
                     for Specific Ions. The Nernst equation is used to calculate the  In cardiac surgery, when it is important to have a heart with-
                    equilibrium potential for a particular ion. If the potential differ-  out electrical and mechanical activity, the organ is sometimes per-
                    ence across the membrane is the same as that calculated by the  fused with cool cardioplegic solution. The perfusate typically con-
                    Nernst equation for a particular ion, then the electrical force  tains 15 to 35 mM potassium. As would be predicted from the
                    would counterbalance the concentration difference of that specific  Nernst equation, the cell membranes depolarize. The depolarized
                    ion. If the membrane were permeable to the ion, there would be  cells no longer experience an action potential, resulting in a mo-
                    no net ion movement. An understanding of the equilibrium po-  tionless surgical field.
                    tential is basic to an understanding of the electrical characteristics  Each ion has a different equilibrium potential that depends on
                    of biologic membranes.                              the relative concentrations of that ion on the two sides of the
                       Potassium ion distribution across the sarcolemma provides a  membrane (see Table 1-2). In each case, the equilibrium potential
                    useful example in discussing the Nernst equation. Potassium ions  can be calculated using the Nernst equation. For example, given
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