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Nutrient Limitation of Phytoplankton by Nitrogen and Phosphorus: Erosion of the Phosphorus Paradigm

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Nutrient Limitation of Phytoplankton by Nitrogen and Phosphorus: Erosion of the Phosphorus Paradigm NUTRIENT LIMITATION OF PHYTOPLANKTON BY NITROGEN AND PHOSPHORUS: EROSION OF THE PHOSPHORUS PARADIGM Wayne A. Wurtsbaugh Utah State University William M. Lewis, Jr. University of Colorado Nice, 26 January 2009 N vs P Limitation The Controversy That Won’t Die! Schindler, DW. 1977. Evolution of phosphorus limitation in lakes, Science Downing, JA & E McCauley. 1992. The nitrogen : phosphorus relationship in lakes. Limnology & Oceanography Elser, JJ et al. 2007. Global analysis of nitrogen and phosphorus limitation of primary producers in freshwater, marine and terrestrial ecosystems. Ecology Letters Schindler, DW et al. 2008. Eutrophication of lakes cannot be controlled by reducing nitrogen input: Results of a 37-year whole-ecosystem experiment. Proc. National Acad. Sci. Sterner, RH 2008. On the phosphorus limitation paradigm for lakes. International Review of Hydrobiology Lewis, WM, Jr. and WA Wurtsbaugh. 2008. Control of lacustrine phytoplankton by nutrients: Erosion of the phosphorus paradigm. International Review of Hydrobiology One foundation of the phosphorus Experimental Lakes Area (ELA) paradigm are whole-lake experiments suggesting that P North alone controls algal biomass alone controls algal biomass P, N, C David Schindler (1977, 2008, etc.) Concluded: N, C • Only phosphorus is important Divided Lake 226 South • If nitrogen is in short supply, nitrogen fixation by cyanobacteria will make up the nitrogen deficit: Schindler (1977) Evolution of phosphorus limitation in lakes, Science However, median N-fixation as a proportion of Cyanobacteria the total N necessary to support primary N > NH production is less than 5% (Howarth et al. 1988) 2 ——— 3 – Cyanobacteria rarely make up the deficit However,However, responseresponse toto nutrientnutrient additionsadditions inin allall ofof thethe ELAELA LakesLakes suggestsuggest aa differentdifferent conclusionconclusion 600 Adapted from Planktonic N- 500 Everett Fee (1979) fixation in most of +N P these eutrophic 400 lakes 300 200 +N +P 100 North C hlorophyll (% of predicted) P, N, C 0 1 1 2 2 2 2 3 3 3 3 1 1 4 4 4 4 5 5 5 5 6 6 Chlorophyll (% above predicted) Chlorophyll (% above -100 Lake Years { N, C Lake Years Lake 226 NorthDivided Lake 226 South No nitrogen fixation in plankton, but important in epiphytic periphyton (D. Schindler, personal communication) Small-scale bioassays also shed doubt on the phosphorus paradigm Waste Water Treatment Cutler Reservoir, Utah (Sept. 2008) Effluent Logan River WWTP Area Area Day 4 Results ManyMany lakeslakes showshow NN-- oror NPNP--limitationlimitation ) 500 400 Lake Bioassays Summary of 32 300 Bioassays in 8 Widely- 200 different Lakes in western US, Spain, Peru 100 (W. Wurtsbaugh) 0 1 3 5 7 91113151719212325272931333537394143454749515355575961636567697173757779818385878991939597991011031051071091111 Chlorophyll (% Above Control Above (% Chlorophyll -100 Chlorophyll (% of Controls (% Chlorophyll +N +P +NP +Mic -200 600 500 +N P 400 Experimental Lakes Area of Canada 300 +N +P (adapted from Fee 1979) 200 100 Chlorophyll (% of predicted) Chlorophyll (% of predicted) (% Chlorophyll 0 1 1 2 2 2 2 3 3 3 3 1 1 4 4 4 4 5 5 5 5 6 6 -100 Lake Years Regional Differences Atmospheric deposition of anthropogenic N may influence which nutrients are limiting 3.0 2.5 N-limitation 2.0 1.5 1.0 (Log +N/+P) (Log 0.5 Relative ResponseRelative P-limitation p = 0.07 0.0 Derived from Elser et al. (1990) & US 024681012National Atmospheric Deposition Program (EPA) Inorganic Nitrogen Deposition (kg/ ha/ yr) from NADP 37-Year Whole-Lake Bioassay Experiment (Schindler et al. 2008) NP NP P only Once eutrophic, ) 3 adding only P has maintained high algal levels for 12 yr Nitrogen-fixing Algal Biomass (mg/m cyanobacteria Lake 227, ELA “Long-term” Bioassay Results Logan River Area Day 20 Results Cyanobacteria (Anabaena) associated Logan River with “benthic” walls of flasks Area Day 4 Results Meta-analysis of Elser et al. (2007) also indicates that periphyton respond better to P-alone additions than do the algae in pelagic zones 1.4 1.2 Is strong response to 1.0 P limited primarily to 0.8 eutrophic lakes & 0.6 “eutrophic” biofilms? 0.4 Response Ratio Response Ratio ( ln Treatment / Control ) / Control ( ln Treatment 0.2 0.0 +N +P123 +NP +N +P +NP Lake Lake Benthos Pelagic ConclusionsConclusions • Both N and P can be important in controlling eutrophication: We need a more balanced and integrated approach for understanding eutrophication whether we’re studying freshwaters or marine ecosystems. • Pelagic nitrogen-fixing cyanobacteria are limited by more than just phosphorus, but we still do not have a good understanding of this process in either freshwaters, estuaries or the oceans ConclusionsConclusions • P may be more effective in promoting N- fixation in eutrophic situations: • Eutrophic lakes • “Eutrophic” benthic areas of lakes (or flasks) • If so, control factors for eutrophication and oligotrophication may not be symmetrical: N and P necessary Oligotrophic ----------Æ Eutrophic Remove only P? Oligotrophic <------------ Eutrophic ConclusionsConclusions • Management of eutrophication must consider: • Current limiting nutrient in system • Cost-effectiveness of removing P, N • May be most efficient to make a nutrient limiting by removing it from effluent, even though it might not initially be limiting MerciMerci DidnDidn’’tt presentpresent thethe followingfollowing slidesslides Regional Differences Clearly Evident Nitrogen Deposition -2 -1 3 NE U.S. (mg m y ) watersheds g N / L) μ From Galloway et al. 2004 N loss from 13 Andean Watersheds N Concentration ( N Concentration Perakis, S.S. and L.O. Hedin. 2002. Nitrogen loss from unpolluted South American forests mainly via dissolved organic compounds Nature 415, 416- 419. 300 A foundation of phosphorus 200 paradigm is the stronger 100 50 , µg/L correlation between TP and algal a 30 biomass in suites of lakes 20 10 5 Chlorophyll 3 2 -The simulated data at right shows a 1 situation when the cell quota of of 1 2 3 5 10 20 30 50 100 300 500 1000 the N:P in algae was set at 16:1 such Total P, µg/L that neither nutrient was more 300 limiting than the other, but with 200 reasonable++ assumptions concerning 100 , µg/L 50 a the concentrations of DON, DOP, 30 DIN and SRP. 20 10 5 -In this case the natural variability of Chlorophyll 3 the relatively non-available DON 2 causes a much high scatter in the 1 30 50 100 200 300 500 1000 2000 3000 TN-chlorophyll relationship Total N, µg/L Elser et al. 2007. Global analysis of nitrogen and phosphorus limitation of primary producers in freshwater, marine and terrestrial ecosystems. Ecology Letters 10: 1-8 N > 140 ±S.E. N > 100 N > 500 Response Ratio (Total N) 10 Log Log10 (Total P) Sterner 2008 CanCan nitrogennitrogen fixingfixing cyanobacteriacyanobacteria makemake upup thethe NN--deficiency?deficiency? For eutrophic lakes showing N fixation in the plankton, the median contribution to total load that could be attributed to N fixation is near 22%, and the median fixation as a proportion of the total N necessary to support primary production is less than 5%, according to the data compiled by HOWARTH et al. (1988). -- Lewis & Wurtsbaugh (2008) Limited by some other nutrient (e.g. Fe, Wurtsbaugh and Horne 1983) Light limitation (energetic constraints) Turbulence Grazing losses.
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