Microbial growth requires the polymerization of biochemical building blocks into proteins, nucleic acids, polysaccharides, and lipids. The building blocks must come preformed in the growth medium or must be synthesized by the growing cells. Additional biosynthetic demands are placed by the requirement for coenzymes that participate in enzymatic catalysis. Biosynthetic polymerization reactions demand the transfer of anhydride bonds from adenosine triphosphate (ATP). Growth demands a source of metabolic energy for the synthesis of anhydride bonds and for the maintenance of transmembrane gradients of ions and metabolites.
Metabolism has two components, catabolism and anabolism (Figure 6-1). Catabolism encompasses processes that harvest energy released from the breakdown of compounds (eg, glucose), and using that energy to synthesize ATP. In contrast, anabolism, or biosynthesis, includes processes that utilize the energy stored in ATP to synthesize and assemble the subunits, or building blocks, of macromolecules that make up the cell. The sequence of building blocks within a macromolecule is determined in one of two ways. In nucleic acids and proteins, it is template-directed: DNA serves as the template for its own synthesis and for the synthesis of the various types of RNA; messenger RNA serves as the template for the synthesis of proteins. In carbohydrates and lipids, on the other hand, the arrangement of building blocks is determined entirely by enzyme specificities. Once the macromolecules have been synthesized, they self-assemble to form the supramolecular structures of the cell, eg, ribosomes, membranes, cell wall, flagella, and pili.
The relationship between catabolism and anabolism. Catabolism encompasses processes that harvest energy released during disassembly of compounds, using it to synthesize adenosine triphosphate (ATP); it also provides precursor metabolites used in biosynthesis. Anabolism, or biosynthesis, includes processes that utilize ATP and precursor metabolites to synthesize and assemble subunits of macromolecules that make up the cell. (Reproduced with permission from Nester EW, Anderson DG, Roberts CE, Nester MT [editors]: Microbiology: A Human Perspective, 6th ed. McGraw-Hill, 2009, p. 127. © The McGraw-Hill Companies, Inc.)
The rate of macromolecular synthesis and the activity of metabolic pathways must be regulated so that biosynthesis is balanced. All of the components required for macromolecular synthesis must be present for orderly growth, and control must be exerted so that the resources of the cell are not expended on products that do not contribute to growth or survival.
This chapter contains a review of microbial metabolism and its regulation. Microorganisms represent extremes of evolutionary divergence, and a vast array of metabolic pathways is found within the group. For example, any of more than half a dozen different metabolic pathways may be used for assimilation of a relatively simple compound, benzoate, and a single pathway for benzoate assimilation may be regulated by any of more than half a dozen control mechanisms. Our goal is to illustrate the principles that underlie metabolic ...