The importance of trace metal nutrients for marine phytoplankton and bacteria along Line-P. Jay T. Cullen, Maite Maldonado (UBC), Erin Lane (UBC)
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1 The importance of trace metal nutrients for marine phytoplankton and bacteria along Line-P Jay T. Cullen, Maite Maldonado (UBC), Erin Lane (UBC)
2 Outline Trace Metals and Marine Biogeochemical Cycles Iron and microbial production in the Gulf of Alaska Dissolved Fe distributions Limitation of algal growth Fe acquisition by marine microbes Bacterial carbon use efficiencies The Future: Trace Metal Interactions and Global Elemental Cycles Fe limitation and Cd biogeochemistry Role of Cu in Fe uptake by eukaryotic phytoplankton
3 CO 2 Trace Metals, Enzymes and Biogeochemical Cycles N 2 Atmosphere Ocean CO 2(aq) Fe, Zn, Co, Cd Biota Fe Mo V HCO 3 - CO 3 2- Deep Ocean NO 3 - Thermocline
4 Primary Metal Requirements For Microbially Mediated N Transformations Morel and Price 2005 Science
5 Known Biochemical Functions of Selected Trace Elements in Marine Microorganisms Morel et al Treatise on Geochem.
6 Surface Nitrate Concentrations Showing HNLC Regions
7 The Distribution of Dissolved Fe at OSP Aug. 5, 1987 Martin et al DSR
8 High Nutrient-Low Chlorophyll Conditions and Dissolved Fe Along Line P (Wong, Johnson) LaRoche et al Nature
9 Fe Limits Algal Growth in Shipboard Incubation Studies at OSP Martin and Fitzwater 1988 Nature
10 Organic Complexation of Dissolved Fe after Rue and Bruland 1995 Marine Chemistry
11 Fe acquisition by phytoplankton Fe model Fe? Transporter Seawater Fe chemistry with strong L L < 0.001% Inorganic Fe(III) photochemical & thermal Fe(III)L in out (Anderson & Morel 1982, Hudson & Morel 1990, Sunda & Huntsman 1995) organically bound > 99.99%
12 Utilization of Organically Complexed Fe by Marine Microbes and Fe-light Co-limitation Maldonado and Price 1999 DSR Eukaryotic reduction of FeL at cell surface allows access to strong organic Fe complexes Physiological response of marine microbes to Fe limitation New model of Fe acquisition by marine phytoplankton
13 Revised Model for Fe Acquisition by Fe-limited eukaryotic phytoplankton Fe inorganic Transporter Fe R+ e- Reductase L R in out HIGH-AFFINITY Fe Transport System Fe(III)L Organically bound
14 Marine Diatoms Economize Cellular Fe When Resource is Scarce LaRoche et al Nature
15 Substitution of Flavodoxin for Ferredoxin in Diatoms Along Line P LaRoche et al Nature
16 Fe Requirements and Physiology of Heterotrophic Marine Bacteria Azam 1998 Science
17 Fe Requirements and Physiology of Heterotrophic Marine Bacteria Azam 1998 Science
18 Fe Requirements and Physiology of Heterotrophic Marine Bacteria at OSP Tortell, Maldonado and Price 1996 Nature 1. Heterotrophic bacteria have higher Fe:C than eukaryotic phytoplankton 2. Constitute ~50% of total biogenic Fe 3. Account for 20-45% of Fe uptake
19 Bacterial ETS Activity and Carbon Use Efficiencies Affected By Fe-Limitation Tortell, Maldonado and Price 1996 Nature
20 Fe Requirements and Physiology of Heterotrophic Marine Bacteria at OSP Tortell, Maldonado and Price 1996 Nature 4. Bacterial Fe quotas in the field imply Felimitation 5. Fe-limited cultures have lower ETS activity, growth rates and C use efficiencies In Fe-limited ecosystems heterotrophic bacteria may respire more organic carbon reducing the amount available to higher trophic levels.
21 Fe Biogeochemistry in Surface Waters of HNLC Areas Tortell et al FEMS
22 Trace Metal Biota Interactions 1. There is a reciprocal relationship Trace metal distributions and availability affect the microbial community structure Microbes can alter the chemical speciation and distribution of trace metals 2. The biogeochemical cycles of C, N and P (Si) can be modulated by trace metals
23 Outline Trace Metals and Marine Biogeochemical Cycles Iron and microbial production in the Gulf of Alaska Dissolved Fe distributions Limitation of algal growth Fe acquisition by marine microbes Bacterial carbon use efficiencies The Future: Trace Metal Interactions and Global Elemental Cycles Fe limitation and Cd biogeochemistry Role of Cu in Fe uptake by eukaryotic phytoplankton
24 Global Dissolved Cadmium Versus Phosphate Relationship 1.2 dissolved Cd (nmol L -1 ) Yeats and Westerlund (1991) Bruland (1980) Bruland (1983) Danielsson and Westerlund (1983) Frew and Hunter (1992) Nolting and De Baar (1994) Frew and Hunter (1995) Kremling and Pohl (1989) Bruland et al. (1978) Knauer and Martin (1981) Ellwood (2004) Fitzwater et al. (2000) Abe (2001) PO 4 3- (μmol L -1 ) Cullen 2006 L&O
25 The Effect of Fe Bioavailability on the Cd:P Composition of Phytoplankton Cullen 2006 L&O
26 Dissolved Cd:PO 4 Lower in HNLC Fe-Limited System dissolved Cd (nmol L -1 ) Bruland (1983) Bruland et al. (1978) Chen et al. (2005 ) Abe (2002a) This study Abe (2002b) Kremling and Pohl (1989) Yeats and Westerlund (1991) Ellwood (2004) Fitzwater et al. (2000) Martin et al. (1989) Frew and Hunter (1995) This study PO4 3- (μmol L -1 ) Cullen 2006 L&O
27 An Fe link to the Cd:PO 4 kink? 1.5 Fe-limited algal growth in surface waters Cd (nmol kg -1 ) Elevated Cd:PO 4 in nutricline PO 4 (µmol kg -1 ) 3 4
28 Mechanism: Non-specific Fe Transporter (Lane and Maldonado, UBC) Up-regulated under Fe-limited Fe deficient plants often accumulate Cd Cd inhibits Fe (II) transport Searched the recently sequenced T. pseudonana genome and found Nramps Plasma membrane IRT1 Plant FET4 Nramps Yeast Curie et al., 2000 ; Eide et al., 1996 Dix et al., 1997
29 The role of Cu in the high-affinity Fe transport system of marine diatoms (Maldonado et al. in press L&O) Investigations historically (even just 5 years ago) have focused on toxic effects that shape microbial community structure (Barber and Ryther 1969, Moffett et al.) Granger and Ward 2002 L&O Cu limitation and N 2 O production by denitrifiers What physiological processes are affected by Cu limitation in marine diatoms?
30 Mechanism of Fe acquisition by Fe-limited marine diatoms Fe(III) R+ Transporter e- Reductase Fe(III) Cu Oxidase O 2 Fe(II) R in out Fe(III)L 4Fe 2+ + O 2 + 4H + = 4Fe H 2 O 4Fe H + + O 2 = 4Fe 3+ + H 2 O 2
31 Effect of Cu-limitation on Fe transport by Fe-limited Thalassiosira oceanica 6 5 High Cu Low Cu Particulate 55 Fe (*10-21 mol Fe mm -2 ) Time (hours) 100 nm Fe : 1 µm siderophore (DFB) 6 fold difference in Fe uptake rates
32 Fe(II) concentration (μm) Effects of Cu addition on Fe(II) oxidation by Fe-Cu-limited T. oceanica ph 6.6 Time (hours) Control (no cells) Cells (low Cu) Cells (high Cu)
33 Where do we go from here? We are poised to develop more mechanistic understandings of the cellular level processes that can affect basin scale distribution of trace elements and CNPSi biogeochemical cycling The subarctic Pacific is an ideal natural laboratory and the Line P program has proven the ideal platform to probe metal biota interactions
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