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Thermodynamics of Biochemical Reactions

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This book is about the thermodynamics of enzyme-catalyzed reactions that make up the metabolism of living organisms. It is not an introductory text, but the fundamental principles of thermodynamics are reviewed. The reader does need some background in thermodynamics, such as that provided by a first course in physical chemistry. The book uses a generalized approach to thermodynamics that makes it possible to calculate the effects of changing pH, free concenrations of metal ions that are bound by reactants, and steady-state concentrations of coenzymes. This approach can be extended to other types of work that may be involved in a living organism.

The concepts involved in this approach are simple, but the equations become rather complicated. Biochemical reactions are written in terms of reactants like ATP that are made up of sums of species, and they are referred to as biochemical reactions to differentiate them from the underlying chemical reactions that are written in terms of species. The thermodynamics of biochemical reactions is independent of the properties of the enzymes that catalyze them. However, the fact that enzymes may couple reactions that might otherwise occur separately increases the number of constraints that have to be considered in thermodynamics.

Biochemical thermodynamics is complicated for several reasons: (1) Biochemical reactants consist of sums of species whenever a reactant has a pK within about two units of the pH of interest or binds metal ions reversibly. (2) Species of a biochemical reactant are often ions, and the activity coefficients of ions are functions of ionic strength. (3) Enzyme catalysis may introduce constraints in biochemical reactions in addition to balances of atoms of elements. (4) Metabolism is sufficiently complicated that it is important to find ways to obtain a more global view. (5) In biochemistry other kinds of work, such as electric work, elongation work, and surface work may be involved. It is remarkable that the same basic reactions are found in all living systems. The most important thing about these reactions is that they provide the means to carry out the oxidation of organic matter in a sequence of steps that store energy that is needed for the synthesis of organic molecules, mechanical work, and other functions required for life.
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