We quantified the relationships among the specific growth rate, intracellular iron content, and steady state iron uptake rate for cultures of the marine diazotrophic cyanobacterium Trichodesmium (IMS 101) grown under differing conditions of Fe and N availability. The Fe quotas necessary to support a moderately Fe-limited growth rate (70% max) of 0.1 d Ϫ1 under diazotrophy and ammonium were 38 and 8 mol mol Ϫ1 , indicating a fivefold increased Fe cost for diazotrophy. This increased demand reflects the influences of both the ninefold lower marginal use efficiency and the greater maintenance Fe : C requirement at zero growth rate under diazotrophy (13.5 versus 5.2 mol mol Ϫ1 ). For diazotrophic growth at ϭ 0.1 d Ϫ1 , we estimate that 19%-53% of the cellular Fe is bound in nitrogenase. Trichodesmium is capable of luxury uptake of at least 13-fold greater amounts of Fe than needed for moderately Fe-limited growth. At least half of the populations sampled from the continental shelf of northern Australia had Fe : C ratios within the range of iron limitation of laboratory cultures. Bottle incubations showed that, after 3-4 d with added iron, a low Fe : C population (21 mol mol Ϫ1 ) had higher N 2 fixation rates than controls, whereas a high Fe : C population (78 mol mol
Ϫ1) showed no response. The cellular N : P quotas and N 2 fixation rates were compared to those previously reported for the Atlantic Ocean. Empirically, these data suggest a critical N : P ratio ϳ40-50 for the onset of P limitation of growth. Only 10% of the Trichodesmium populations from coastal Australia had N : P ratios within the putative P-limitation range (Ͼ40), whereas 40% the Atlantic populations fell within this range. These comparisons, along with those for N fixation versus colony Fe : C ratio, suggest that Fe limitation is more prevalent in the Australia Trichodesmium populations, while P limitation predominates in the Atlantic populations.Trichodesmium spp. is a genus consisting of five species of nitrogen-fixing cyanobacteria that inhabit (sub)tropical oligotrophic waters. These plankton provide new N for up to 50% of primary production in these waters (Karl et al. 1997) and, thus, can play a prominent role in the biogeochemical cycling of C and N (Carpenter and Romans 1991). N 2 fixation facilitates net C fixation and export from the euphotic zone (Karl et al. 1995) and may have pronounced global climate implications (Falkowski 1997). Iron has been proposed to be a primary factor limiting N 2 fixation and growth of Trichodesmium due to the high iron requirement for diazotrophic growth and the low availability of iron in oceanic 1 Present address: Department of Geosciences, Princeton University, Princeton, New Jersey 08544.2 Present address: Romberg Tiburon Center, San Francisco State University, 3152 Paradise Drive, Tiburon, California 94920.
AcknowledgmentsThe crew of the R/V Maurice Ewing, as well as Jim King and Rachel Foster, were instrumental in implementing our field sampling campaign. We thank Tracy Villareal for providing ...