In this chapter the general processes involved in controlling production and transformation of organic matter will be discussed as well as some of the associated stoichiometric changes of a few key biological elements (e.g., C, N, P, S). Stoichiometry is defined as the mass balance of chemical reactions as they relate to the law of definite proportions and conservation of mass (Sterner and Elser, 2002). For example, if we examine the average atomic ratios of C, N, and P in phytoplankton we see a relatively consistent ratio of 106:16:1 in most marine species. This is perhaps the best example of applied stoichiometric principles in natural ecosystems and is derived from the classic work of Alfred C. Redfield (1890–1983) (Redfield, 1958; Redfield et al., 1963). More specifically, Redfield compared the ratios of C, N, and P of dissolved nutrients in marine waters to that of suspended marine particulate matter (seston) (essentially phytoplankton) and found straight lines with equal slopes (figure 8.1; Redfield et al., 1963). This relationship suggested that marine biota were critical in determining the chemistry of the world ocean, clearly one of the most important historical findings linking chemical and biological oceanography (Falkowski, 2000). Moreover, the Redfield ratio has been further validated with recent data using improved analytical techniques (Karl et al., 1993; Hoppema and Goeyens, 1999). Other work has shown that there are predictable deviations from the Redfield ratio across a freshwater to open ocean marine gradient (figure 8.2; Downing, 1997). For example, N-to-P ratios in estuaries have commonly been shown to be lower and/or higher than the predicted Redfield ratio because of denitrification and anthropogenic nutrient enrichment processes, respectively. Inputs of vascular plant organic matter (e.g., mangroves, salt marshes, seagrasses) to estuarine systems presents another problem in causing deviations of C:N:P from the Redfield ratio. Vascular plants have been shown to deviate from this ratio in part because of relatively high amounts of C and N compared to algae due to a higher abundance of structural support molecules (e.g., cellulose, lignin) and defense antiherbivory (secondary) compounds (e.g., tannins), respectively (Vitousek et al., 1988).
Speciation of organosulfur compounds in carbonaceous chondrites