Archaeal membrane lipids known as glycerol dibiphytanyl glycerol tetraethers (GDGTs) are the basis of the TEX 86 paleotemperature proxy. Because GDGTs preserved in marine sediments are thought to originate mainly from planktonic, ammonia-oxidizing Thaumarchaeota, the basis of the correlation between TEX 86 and sea surface temperature (SST) remains unresolved: How does TEX 86 predict surface temperatures, when maximum thaumarchaeal activity occurs below the surface mixed layer and TEX 86 does not covary with in situ growth temperatures? Here we used isothermal studies of the model thaumarchaeon Nitrosopumilus maritimus SCM1 to investigate how GDGT composition changes in response to ammonia oxidation rate. We used continuous culture methods to avoid potential confounding variables that can be associated with experiments in batch cultures. The results show that the ring index scales inversely (R 2 = 0.82) with ammonia oxidation rate (ϕ), indicating that GDGT cyclization depends on available reducing power. Correspondingly, the TEX 86 ratio decreases by an equivalent of 5.4°C of calculated temperature over a 5.5 fmol·cell −1 ·d −1 increase in ϕ. This finding reconciles other recent experiments that have identified growth stage and oxygen availability as variables affecting TEX 86 . Depth profiles from the marine water column show minimum TEX 86 values at the depth of maximum nitrification rates, consistent with our chemostat results. Our findings suggest that the TEX 86 signal exported from the water column is influenced by the dynamics of ammonia oxidation. Thus, the global TEX 86 -SST calibration potentially represents a composite of regional correlations based on nutrient dynamics and global correlations based on archaeal community composition and temperature.T he glycerol dibiphytanyl glycerol tetraether (GDGT) membrane lipids of Archaea are abundant in marine water columns and sediments. The major source of GDGTs to ocean sediments is thought to be planktonic, ammonia-oxidizing Archaea (AOA) affiliated with the phylum Thaumarchaeota (formerly Marine Group I Crenarchaeota) (1, 2). Thaumarchaeota play a primary role in the nitrogen cycle, performing the first and rate-limiting step of nitrification-namely, the oxidation of ammonia to nitrite (3-5). Accordingly, Thaumarchaeota are most abundant at the base of, or below, the euphotic zone (6-9). Based on the phylogeny of their ammonia monooxygenase gene, the planktonic Thaumarchaeota are divided into two distinct clusters, the Water Column Cluster A that is most abundant in the epi-and upper mesopelagic (above ∼200 to ∼500 m, depending on location) and the Water Column Cluster B that dominates thaumarchaeal assemblages in the deeper mesopelagic and bathypelagic (7-10). These clusters putatively represent thaumarchaeal ecotypes adapted to high and low ammonium flux, respectively (11,12).Thaumarchaeota produce GDGTs containing from zero to four cyclopentane rings (GDGT-0 to GDGT-4) or four cyclopentane rings and one additional cyclohexane ring (e.g., in crenarchaeol;...