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The second level, or cell-quota theory, allows organisms to vary their content of nutrient Q (and hence their yield of biomass from assimilated nutrient). It is increasingly referred to as the DROOP model after one of its authors (Droop, 1968, 1983). In principle, the quota should be defined as the ratio of nutrient to biomass (Droop, 1979) and will here be understood as the population (atomic) ratio of the nutrient element to carbon. A simplified version of the theory and some deductions from it, is given in Box 1 (refer page 348). The key equation (ignoring physical transports) is ... 

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