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              Bioenergetics                                                                               115

              reactions is directly linked to the transmembrane flow of  opportunity to unravel the mysteries of organisms from
              protons into the lumen of the membrane. This proton flow  planets other than Earth.
              results in the generation of an electrochemical proton gra-
              dient. Essentially, part of the light energy is conserved
              by formation of this gradient as well as by formation of  SEE ALSO THE FOLLOWING ARTICLES
              the strong reducing agent NADPH. The flow of protons
              provides the energy needed for the synthesis of the ter-  CARBOHYDRATES • CARBON CYCLE • CHROMATIN
              minal phosphate anyhydride bond of ATP, an example of  STRUCTURE AND MODIFICATION • ELECTRON TRANS-
              the conversion of the osmotic and electrical energy of the  FER REACTIONS • ENERGY FLOWS IN ECOLOGY AND
              proton gradient to chemical bond energy. The syntheses  IN THE ECONOMY • ENERGY TRANSFER,INTRAMOLECU-
              of ATP and NADPH capture some of the light energy. In  LAR • ION TRANSPORT ACROSS BIOLOGICAL MEMBRANES
              turn, ATP and NADPH drive the unfavorable reduction of  • LIPOPROTEIN/CHOLESTEROL METABOLISM • PROTEIN
              CO 2 by H 2 O to form carbohydrates and O 2 .     SYNTHESIS • THERMODYNAMICS
                Organisms, especially bacteria, have evolved novel
              bioenergetic mechanisms that are well suited to their en-
              vironments. For example, the bacterium Halobacter halo-  BIBLIOGRAPHY
              bium lives in salt marshes and requires NaCl at concentra-
              tions that kill other organisms. These halophilic bacteria  Cramer, W. A., and Knaff, D. B. (1990). “Energy Transduction in Bio-
              contain patches of a purple protein, halorhodopsin, on its  logical Membranes: A Text of Bioenergetics,” Springer-Verlag, New
                                                                 York.
              plasma membrane. Halorhodopsin is a light-driven proton
                                                                Garrett, R. H., and Grisham, C. M. (1999). “Biochemistry,” 2nd ed.,
              pump and its operation causes protons to be ejected from  Saunders College Publishing, Fort Worth.
              the cells. The resulting electrochemical proton gradient  McCarty, R. E. (1999). “Chemiosmotic Coupling,” In “Encyclopedia of
              may be used to drive ATP synthesis or the transport of  Molecular Biology” (T. Creighton, ed.), pp. 402–408, John Wiley and
              biochemicals. Given the diversity of the environments in  Sons, Inc., New York.
                                                                Nichols, D. G., and Ferguson, S. J. (1992). “Bioenergetics 2,” Academic
              which organisms grow, it is possible that biochemists will
                                                                 Press, London.
              uncover new ways in which organisms meet their ener-  Ort, D. R., and Yocum, C. F., eds. (1996). “Oxgenic Photosynthesis: The
              getic needs. Perhaps future bioenergeticists will have the  Light Reactions,” Kluwer Academic, Dordrechtshill, Norwell, MA.
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