“…The equilibrium view predicts a ratio too high, and the growth-rate approach (see also Williams, 2006) colleagues (2004a, 2004b) build on classical chemostat models of the ocean, incorporating growth according to a standard law (Droop, 1974) and Liebig's Law of the Minimum (which states simply that growth will be limited by whichever essential nutrient is in shortest supply), to show that, in general, the system's dynamics will reach an equilibrium in which a single factor is limiting. When this dynamic is embedded into an evolutionary framework, however, in which organisms must expend their available carbon on making nitrogen-rich proteins (the primary machines responsible for uptake of nutrients) or phosphorusrich ribosomes (which play a key role in growth), the system evolves to colimitation, in which neither nutrient is in (Karl, 2002) Figure 1B, C).…”