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Energy Production and Metabolism  Heat is transferred in all chemical reactions.
                                       The amount of heat produced upon conversion
       Energy is the ability of a system to perform  of a given substance into product X is the same,
       work; both are expressed in joules (J). A poten-  regardless of the reaction pathway or whether
       tial difference (potential gradient) is the so-  the system is closed or open, as in a biological
       called driving “force” that mobilizes the matter  system. For caloric values, see p. 228.
       involved in the work. Water falling from height  Enthalpy change (∆H) is the heat gained or
       X (in meters) onto a power generator, for ex-  lost by a system at constant pressure and is re-
       ample, represents the potential gradient in
                                       lated to work, pressure, and volume (∆H = ∆U
    Fundamentals and Cell Physiology  formed can be determined by multiplying the  ∆H is positive in endothermic reactions. The
                                       + p ! ∆V). Heat is lost and ∆H is negative in ex-
       mechanical work. In electrical and chemical
       work, potential gradients are provided respec-
                                       othermic reactions, while heat is gained and
       tively by voltage (V) and a change in free en-
                  – 1
       thalpy ∆G (J ! mol ). The amount of work per-
                                       second law of thermodynamics states that the
                                       total disorder (randomness) or entropy (S) of a
       potential difference (intensity factor) by the
                                       closed system increases in any spontaneous
                                       process, i.e., entropy change (∆S) # 0. This
       corresponding capacity factor. In the case of
                                       must be taken into consideration when at-
       the water fall, the work equals the height the
                                       tempting to determine how much of ∆H is
       water falls (m) times the force of the falling
       water (in N). In the other examples, the
                                       freely available. This free energy or free en-
       times the amount of charge (C). Chemical work
                                       chemical reaction. The heat produced in the
       performed = ∆G times the amount of sub-
                                       process is the product of absolute temperature
    1  amount work performed equals the voltage (V)  thalpy (∆G) can be used, for example, to drive a
                                       and entropy change (T · ∆S).
       stance (mol).
         Living organisms cannot survive without an  Free enthalpy (∆G) can be calculated using
       adequate supply of energy. Plants utilize solar  the Gibbs-Helmholtz equation:
       energy to convert atmospheric CO 2 into oxy-  ∆G ! ∆H – T · ∆S.  [1.24]
       gen and various organic compounds. These, in  ∆G and ∆H are approximately equal when ∆S
       turn, are used to fill the energy needs of  approaches zero. The maximum chemical
       humans and animals. This illustrates how  work of glucose in the body can therefore be
       energy can be converted from one form into  determined based on heat transfer, ∆H,
       another. If we consider such a transformation  measured during the combustion of glucose in
       taking place in a closed system (exchange of  a calorimeter (see p. 228 for caloric values).
       energy, but not of matter, with the environ-  Equation 1.24 also defines the conditions
       ment), energy can neither appear nor disap-  under which chemical reactions can occur. Ex-
       pear spontaneously. In other words, when  ergonic reactions (∆G $ 0) are characterized
       energy is converted in a closed system, the  by the release of energy and can proceed spon-
       total energy content remains constant. This is  taneously, whereas endergonic reactions (∆G
       described in the first law of thermodynamics,  # 0) require the absorption of energy and are
       which states that the change of internal energy  not spontaneous. An endothermic reaction
       (= change of energy content, ∆U) of a system  (∆H # 0) can also be exergonic (∆G $ 0) when
       (e.g. of a chemical reaction) equals the sum of  the entropy change ∆S is so large that ∆H–
       the work absorbed (+W) or performed (–W) by  T · ∆S becomes negative. This occurs, for ex-
       a system and the heat lost (–Q) or gained (+Q)  ample, in the endothermic dissolution of crys-
       by the system. This is described as:  talline NaCl in water.
         ∆U ! heat gained (Q) " work performed  Free enthalpy, ∆G, is a concentration-de-
         (W) [J] and            [1.22]  pendent variable that can be calculated from
         ∆U ! work absorbed (W) " heat lost  the change in standard free enthalpy (∆G ) and
                                                                0
         (Q) [J].               [1.23]  the prevailing concentrations of the sub-
                                                      0
       (By definition, the signs indicate the direction  stances in question. ∆G is calculated assum-
       of flow with respect to the system under con-  ing for all reaction partners that concentration
   38  sideration.)                    = 1 mol/L, pH = 7.0, T = 298 K, and p = 1013 hPa.
                                                                   !
       Despopoulos, Color Atlas of Physiology © 2003 Thieme
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