Problems of adverse pH and bacterial strategies to combat it

This chapter aims to survey the problems faced by bacteria found in environments of adverse pH, to review strategies used to combat those problems and to ask how those strategies are implemented. At acid or alkaline pH, bacteria are challenged not just by excess of H+ or OH- but also by excess of metal ions (aluminium, heavy metals at acidic pH, Na+ at alkaline pH), as well as shortages. Bacteria attempt to maintain their intracellular pH by minimizing membrane permeability to H+ and other ions, buffering the cytoplasm, ameliorating the external pH through catabolism or selective substrate utilization, and developing ionic pumping systems. The amelioration of pH depends on the availability of substrate, and is unlikely in most naturally stressful environments. Ion pumping is expensive energetically, although the cost to growth is unknown. The response to adverse pH involves sensing systems and responsive regulatory systems. The adaptive acid tolerance response is now well known in and other bacteria, but is there a widespread adaptive alkali tolerance response? What and where are the sensors? Whether they sense intracellular pH, extracellular pH or delta pH is unclear, although an external sensory input seems essential. Is there one major sensory system responsive to pH or multiple systems with back-up mechanisms? What and where are the regulators? Is there one central regulator controlling all the responses or are there cascades of responses?