“…One of the most important processes is the reversible reaction between orthophosphate and pyrophosphate catalyzed by pyrophosphatase, which exchanges all O atoms in PO 4 with ambient water O, resulting in a temperature dependent equilibrium fractionation [ Blake et al ., ; Longinelli and Nuti , ]. This isotopic equilibration between PO 4 and water has been observed in many environments as an indicator of biological P turnover (i.e., PO 4 uptake, intracellular cycling, and PO 4 release into environment) [ Kolodny et al ., ; McLaughlin et al ., , ; Paytan et al ., ]. Moreover, many extracellular enzymes catalyze reactions that involve breaking of P bonds and cause kinetic isotopic effects [ Blake et al ., ; Colman et al ., ; Liang and Blake , ], providing a potential means to identify P sources and transformations.…”