The existence of unusually large fluctuations in the Neoproterozoic (1,000 -543 million years ago) carbon-isotopic record implies strong perturbations to the Earth's carbon cycle. To analyze these fluctuations, we examine records of both the isotopic content of carbonate carbon and the fractionation between carbonate and marine organic carbon. Together, these are inconsistent with conventional, steady-state models of the carbon cycle. The records can be well understood, however, as deriving from the nonsteady dynamics of two reactive pools of carbon. The lack of a steady state is traced to an unusually large oceanic reservoir of organic carbon. We suggest that the most significant of the Neoproterozoic negative carbon-isotopic excursions resulted from increased remineralization of this reservoir. The terminal event, at the ProterozoicCambrian boundary, signals the final diminution of the reservoir, a process that was likely initiated by evolutionary innovations that increased export of organic matter to the deep sea.T he coevolution of the biosphere and geosphere is reflected in large part by changes in the long-term carbon cycle (1). Past changes within the cycle are recorded in the isotopic content of carbonate and organic carbon buried in ancient sediments (2). Extraordinarily large fluctuations occur in the Neoproterozoic [1,000-543 million years ago (Ma)] carbon-isotopic record both immediately preceding the Cambrian diversification of complex animal life (3-5) and in the Ϸ200 million years before it (6). There is much interest in determining not only the cause of these isotopic events (3-9) but how, if at all, they are related to early animal evolution (10).Here we analyze a significant portion of the Neoproterozoic isotopic record by portraying it as a dynamical trajectory in a two-dimensional space indexed by the isotopic content of carbonate carbon and the fractionation between carbonate and marine organic carbon. This depiction, analogous to the construction of phase portraits in dynamical systems theory (11), provides specific predictions for a carbon cycle evolving quasistatically in a succession of steady states. We find that the Cenozoic portion (0-65 Ma) of the carbon-isotopic record satisfies these predictions; however, the Neoproterozoic record does not.The dynamics of a system with two sizeable and isotopically distinct pools of reactive carbon suffice to explain the Neoproterozoic records. Then as now one pool was oceanic and atmospheric CO 2 . Here we show that the other was probably oceanic organic carbon. The isotopic data indicate that this reservoir was large and that its average properties changed slowly. However, fluctuations in its size would have led to major variations in the isotopic record. Moreover, even if the size of the organic reservoir had been perfectly constant, changes in the fractionation associated with organic production would have led to isotopic fluctuations much greater than those predicted by steady-state theory.The fluctuations of greatest interest are those associ...