The surface stress theory of microbial morphogenesis.

From the physics of the situation, one might conclude that the osmotic pressure within most prokaryotes creates a sufficiently high tension in the wall that organisms are at risk of ripping themselves apart. The Surface Stress Theory holds that they avoid this, and are able to carry out certain morphogenetic processes by linking the cleavages of appropriate bonds to enzymes that are sensitive to the stress in the bonds under attack. This tends to maintain the internal pressure and couples wall growth to cytoplasmic growth. Mechanisms with widely different geometry function for different organisms, but they have in common the requirement that new murein be covalently linked, and usually in an unextended conformation. Organisms differ in the site of wall addition and site of cleavage. In the Gram-positive Streptococcus, septum formation, and septal splitting occurs with little stretching of the unsplit septum. In Gram-positive bacilli, the cylinder grows by the inside-to-outside mechanism, and the poles appear to be formed by a split-and-stretch mechanism. Gram-negative rods, with their much thinner wall, resist a spherical shape and are capable of cell division by altering the biochemical mechanism so that initially one-third to one-fifth of the pressure-volume work required to increase the area of the side wall is needed to increase that in a developing pole. The growth of hyphae is a separate case; it requires that much less work is needed to force growth of the apex relative to the side wall. Some other bacterial shapes also can be explained by the theory. But at present, it is only a theory, although it is gradually becoming capable of accounting for current observations in detail. Its importance is that it prescribes many experiments that now need to be done.