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habitats These coupled biogeochemical reactions are: aerobic respiration, denitri cation, Mn reduction, Fe reduction, fermentation, sulfate reduction, methanogenesis, and acetogenesis The hierarchy of the reactions is clear: aerobic respiration occurs at a pE of 1375 and its free energy yield exceeds that of the other reactions Thus, in a given aquatic or terrestrial habitat rich in carbon, microorganisms endowed with the physiological capacity to carry out aerobic respiration will have an advantage their ATP-generating ability exceeds that of the other microbial residents As long as the supply of oxygen is adequate, aerobic respiration will predominate Once the supply of oxygen is exhausted, nitrate-respiring organisms will become predominant and so on down the hierarchy This predictable sequence of physiological processes is reinforced at the level of gene regulation within individual microbial cells.

Generally speaking, the nal electron acceptor allowing the highest free energy yield inhibits expression of genes required for utilization of nal electron acceptors residing lower in the hierarchy (Saffarini et al, 2003; Gralnick et al, 2005) The reader should be aware that kinetic considerations, such as substrate uptake af nities by microorganisms and biochemical reaction rates, can also be important in determining which electronaccepting process will predominate in a given habitat Vital for anaerobic physiology and food chains (see Section 310), hydrogen gas production occurs via reduction of protons This process (mediated by hydrogenase enzymes) may be energetically unfavorable, but allows cells to dispose of excess reducing power, thereby maintaining redox balance (Hedderich and Forzi, 2005) The information in Table 37 accurately represents the energetics of key biogeochemical processes However, the information overlooks both the biochemical and ecological manifestations of the hierarchy.

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Examples of these, especially the involvement of anaerobic food chains are presented in Sections 310 and 73 75 The power of the information embodied by the thermodynamic compass (Figure 39) can be illustrated by its ability to successfully predict, in time and space, what biogeochemical reactions will occur A laboratory experiment, illustrating the sequence of physiological processes (also known as terminal electron acceptor processes, TEAPs) is described in Box 36 The rationale of the experiment is to provide the heterotrophic soil community of microorganisms with an excess supply of electron donor (potato starch) but limited supplies of common 2 electron acceptors (O2, NO3, Mn(OH)4, Fe(OH)3, SO4 , CO2) If water samples are removed and analyzed over time, the thermodynamically predicted patterns shown in Box 36 emerge The most energetically favorable reactions occur rst and the least favorable last.

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The sequence of 2 depleted electron acceptors is O2, NO3, Mn4+, Fe3+, and SO4 Each electron acceptor is converted to its reduced form In an aqueous medium, newly formed H2O would not be detectable (unless the original O2 were labeled with the stable isotope 18O, and analyses were performed with a.