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The response of heterotrophic bacteria in a mesoscale iron (Fe) enrichment was measured in the northeast subarctic Pacific Ocean in July 2002. Addition of FeSO4 increased the dissolved Fe concentration in the fertilized patch to 2–3 nmol L−1 and triggered an increase in concentration of Fe(III)-binding ligands that complexed all of the dissolved Fe. Two to three days later, leucine incorporation rate and specific growth rate of bacteria doubled. Physiological markers of bacterial Fe nutritional state varied during the experiment as the microbial community assimilated the added Fe. Cellular uptake rate of an iron-siderophore complex, 55Fe-ferrioxamine B (FB), increased twofold to fourfold over background values and then declined by day 4.5. The fastest rate of Fe-FB uptake on day 2.5 coincided roughly with a transient increase in outer-membrane, 55Fe-FB-binding protein(s) in bacteria and with the peak in ligand concentration. Maximum potential uptake rate of inorganic Fe (Vmax) was 8 zmol Fe bacterium−1 h−1 prior to Fe enrichment and then decreased by a factor of four within 2.5 d of fertilizing the patch as bacteria became Fe sufficient. Vmax gradually increased by day 6.5 as the bacterial community re-entered iron deficiency. Similar changes in growth and Fe uptake kinetics were observed after a second Fe addition. Heterotrophic bacteria in the subarctic Pacific were Fe-deficient and responded directly to Fe addition by up-regulating pathways for Fe-siderophore acquisition and assimilating complexed Fe. The observation that increases in Fe uptake pathways and production were synchronous is consistent with the hypothesis that bacterial growth was directly limited by Fe.

Centric diatoms isolated from open ocean environments require higher concentrations of Cu for growth than their coastal counterparts. In artificial seawater medium containing ,1 nmol L 21 Cu, three coastal species maintained near maximum rates of growth, but the oceanic clones were unable to survive. Copper limitation was more severe in the diatoms grown in low- than in high-Fe seawater, suggesting that Cu and Fe were interacting essential resources. The interactive effect was in part the result of a Cu requirement for Fe transport. Thalassiosira weissflogii and Thalassiosira oceanica had lower Fe quotas and slower rates of Fe uptake when [Cu] was reduced in the medium. Brief exposure of Cu-limited cells to 10 nmol L 21 Cu increased the instantaneous Fe uptake rate by 1.5 times in T. oceanica. Steady-state uptake rates of both species at high, growth-saturating concentrations of Fe were also Cu dependent. Oceanic species appeared to have an additional Cu requirement that was independent of Fe acquisition and likely responsible for their higher requirements compared to coastal species. Evidence for the importance of Cu in natural communities of phytoplankton was obtained from an incubation experiment performed in the Fe-limited basin of the Bering Sea. Addition of 2 nmol L 21 Cu doubled the phytoplankton net growth rate compared to the untreated controls and, in the presence of extra Fe, increased the growth rate compared to the samples amended with Fe alone. The results suggest that Cu may be an important micronutrient for phytoplankton growth in low-Fe regions of the sea because of its role in Fe acquisition. Paradoxically, oceanic diatoms may be more susceptible to the effects of low Cu concentrations than coastal species.

We have discovered that coastal and oceanic diatoms require more manganese (Mn) to grow in iron (Fe) -deficient than in Fe-sufficient seawater. At low inorganic concentrations, like those of the open sea, Fe and Mn can thus colimit Thalassiosira pseudonana and T. oceanica so that maximum rates of cell division are achieved only when both resources are added simultaneously to cultures. Colimited diatoms amended with either Fe or Mn alone show unique physiological responses, which implies that the observed interaction between Fe and Mn is not caused by a substitution of one metal for the other. Iron deficiency increases the Mn quota of T. pseudonana by three times compared with controls and enhances the production of reactive oxygen species by 1.7 times in T. weissflogii. Both diatoms respond to this oxidative stress by increasing the activities of the antioxidant enzyme superoxide dismutase (SOD). The Mn content of the SODs increases by 1.8 to 2.8 times when Fe is limiting, which suggests that the SODs contain Mn and may account for part of the observed increase in the Mn quota. Such an increased biochemical requirement may elevate the Mn content of low Fe diatoms, and possibly other phytoplankton, resulting in high Mn: Fe ratios in surficial particulate matter in Fe-limited regions of the sea.