Principles of functional and structural organization in the bacterial cell: 'compartments' and their enzymes

Abstract

Most bacteria lack obvious compartmentation, i.e., structural partition of the cell into functional entities (organelles) formed by a closed biological membrane. Nevertheless, these organisms exhibit sophisticated regulation and interactions of their catabolic and anabolic pathways; they are able to exploit a great variety of carbon and energy sources, and they conserve and transform energy in an efficient manner. In a less stringent sense, 'compartments' are also present in bacteria if one accepts that bacterial 'compartments' are not necessarily surrounded by a membrane, but are rather defined as mere functional entities characterized by their structural components, their enzymes and other functional proteins such as binding proteins. This view would mean that the bacterial cell can be described as a highly organized structured system comprised of these functional entities. Regulated transport processes within 'compartments' and across boundaries involving low and high molecular mass compounds, solutes, and ions take place within the 'framework' constituted by this structured system. Special emphasis is given to the fact that many of the transport processes take place involving the functional entity 'energized membrane'. This 'framework', the structural basis for the functional potential of a bacterial cell, can be studied by electron microscopy. Advanced sample preparation techniques and imaging modes are available which keep the danger of artefact formation low; they can be applied at cellular and macromolecular levels. Recent developments in immunoelectron microscopy and affinity labelling techniques provide tools which allow to unequivocally locate enzymes and other antigens in the cell and to identify polypeptide chains in enzyme complexes. Application of these approaches in studies on cellular and macromolecular organization of bacteria and their enzyme systems confirmed some old views but also extended our knowledge. This is exemplified by a description of selected enzyme complexes located in the bacterial cytoplasm, in the cytoplasmic membrane or attached to it, in the periplasmic space, and attached to the cell wall or set free into the surrounding medium.