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Nutrient contributions from Dreissena spp. to Lyngbya wollei and Cladophora glomerata. Patricia Armenio Department of Environmental Sciences and the Lake Erie Center University of Toledo. What are local-scale interactions between Dreissena and benthic algae?. System level.
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Nutrient contributions from Dreissena spp. to Lyngbya wollei and Cladophora glomerata Patricia Armenio Department of Environmental Sciences and the Lake Erie Center University of Toledo
What are local-scale interactions between Dreissena and benthic algae? System level Increased light penetration Local level Resource importation Structural complexity Benthic Primary Production
Contribution to algae N & P Ammonium and phosphate CO2 Respiration Decomposition Feces and pseudofeces Organic matter Other nutrients? Structure Hard shells increase surface area P. Bichier
Do Dreissena contribute to benthic algal blooms? Not new to Lake Erie Have increased Decrease aesthetic value Health risks Food web structure What mechanisms from Dreissena facilitate these blooms?
Lyngbya wollei Cyanobacterium Common in southeastern US Recently washed up on shores of Lake Erie Requires relatively low light Does not attach to hard surfaces Toxic? P. Bichier J. Joyner
Cladophora glomerata • Green alga • Requires relatively high light and hard substrate • Blooms common from 1950s through early 1980s • Return of blooms since mid- 1990s not associated with P loading • Dreissena Higgins et al. 2008
2009 Lyngbya survey • 140 sites total • 113 sites had dreissenid substrate • 77 sites had Lyngbya • 72 sites had both • Only 5 sites that had Lyngbya did not have dreissenid substrate Panek et al.
Manipulative Experiments • Objective: test possible reasons why Dreissena may promote the growth of benthic algae and encourage blooms • N, P • C • Other nutrients • 10 others quantified • Structure
4 treatments 3400 Dreissena/m2 230 g/m2Lyngbya Low-P lake water 12 hour photoperiod One week N=40 Lyngbya Experiment Live Dreissena (N=10) Pottery shards (N=10) Empty Dreissena (N=10) Sand (N=10)
Fed Dreissena throughout experiment • Chlamydomonas reinhardtii • Phytoplankton • Labeled with stable isotope • 13C • 15N • Given to all tanks
Laboratory Analyses • C and N • CHN analyzer • P and other nutrients • ICP-OES • Chlorophyll a • UV-visible spectrophotometer • Phycocyanin • 10-AU fluorometer • Photosynthetic efficiency • DIVING-PAM fluorometer • 13C and 15N • Samples sent to UC Davis, CA
3 2.5 2 y = 0.1538x + 0.1367 1.5 dry weight (g) 2 R = 0.798 p<0.0001 1 0.5 0 0 5 10 15 20 wet weight (g) Lyngbya Positive correlation between wet weight and dry weight
70 60 50 40 % increase in wet weight 30 20 10 0 2.5 2 1.5 Dry weight (g) 1 0.5 0 live Dreissena empty Dreissena pottery shards sand substrate type Lyngbya All tanks increase in weight, but no difference among treatments Error bars = SE
0.5 0.45 0.4 0.35 0.3 Photosynthetic efficiency 0.25 0.2 0.15 0.1 0.05 0 live Dreissena empty Dreissena pottery shards sand substrate type 3 2.5 2 dry weight (g) 1.5 y = -2.8267x + 2.9382 1 R2 = 0.3987 p<0.0001 0.5 0 0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 photosynthetic efficiency Lyngbya No difference in photosynthetic efficiency, but decreases with more dry weight
500 450 400 350 Chlorophyll a (µg/g) 300 250 200 150 100 50 0 6000 5000 4000 Phycocyanin (µg/g) 3000 2000 1000 0 live Dreissena empty Dreissena pottery shards sand substrate type Lyngbya No difference in chlorophyll a, but higher phycocyanin in live Dreissena treatment A AB BC C
400 A 350 B B B 300 250 200 Carbon (mg/g) 150 100 50 0 live Dreissena empty Dreissena pottery shards sand substrate type Lyngbya Dreissena increased carbon content in tissue
6 5 4 Phosphorus (mg/g) 3 2 1 0 45 40 35 30 Nitrogen (mg/g) 25 20 15 10 5 0 live Dreissena empty Dreissena pottery shards sand substrate type Lyngbya Dreissena helped retain macronutrient content in tissue A B B B A B B B
120 100 80 C:P ratio 60 40 20 0 12 10 8 C:N ratio 6 4 2 0 live Dreissena empty Dreissena pottery shards sand substrate type Lyngbya No P deficiency, but N deficiency in all treatments except live Dreissena B B B • <143 is no Pdeficiency • >9.4 C:N is a N deficiency • No deficiency in live Dreissena treatment A B B B A Kahlert 1998
4 3.5 3 2.5 Potassium (mg/g) 2 1.5 1 0.5 0 4 3.5 3 2.5 Sulfur (mg/g) 2 1.5 1 0.5 0 live Dreissena empty Dreissena pottery shards sand substrate type Lyngbya Dreissena helped retain nutrient content in tissue A B B B A B B B
160 B B B 140 A 120 100 Calcium (mg/g) 80 60 40 20 0 live Dreissena empty Dreissena pottery shards sand substrate type Lyngbya However…
-25 A -20 B B B -15 δ13C (‰) -10 -5 0 live Dreissena empty Dreissena pottery shards sand substrate type Lyngbya Stable isotope: less 13C in live Dreissena treatment, no significant difference in 15N
3 B B B 2.5 A 2 δ 15N (‰) 1.5 1 0.5 0 live Dreissena empty Dreissena pottery shards sand substrate type Two week preliminary experiment with Cladophora: the 13C difference goes away and there becomes a significant difference in 15N
Summary for Lyngbya • Dreissena did not increase the biomass of Lyngbya • Increased phycocyanin • Supplied several nutrients, prevented a decrease in others • Decreased Ca • Decreased 13C • Did not respond to substrates
4 treatments 3400 Dreissena/m2 160 g/m2 algae Low-P lake water One week 12 hour photoperiod N=40 Fed Chlamydomonas Cladophora Experiment Live Dreissena (N=10) Pottery shards (N=10) Empty Dreissena (N=10) Sand (N=10)
Extra data for Cladophora • Took algal samples after 24 hours and 48 hours • Tissue nutrient content • Stable isotope • Water samples from each tank at end of experiment compared to initial water • Nutrient
3 2.5 2 dry weight (g) 1.5 y = 0.1387x + 0.4254 R2 = 0.6933 1 p<0.0001 0.5 0 0 2 4 6 8 10 12 14 16 18 wet weight (g) Cladophora Positive correlation between wet weight and dry weight
80 70 60 50 % increase in wet weight 40 30 20 10 0 12 A B 10 bottom biomass 8 wet weight (g) 6 4 2 0 live Dreissena pottery shards sand empty Dreissena substrate type Cladophora All tanks increase in weight, significant difference when other treatments are grouped
2 1.8 1.6 1.4 bottom biomass 1.2 dry weight (g) 1 0.8 0.6 0.4 0.2 0 empty Dreissena pottery shards sand live Dreissena substrate type Cladophora Separation in biomass, more with live Dreissena
Cladophora Results • More variability than Lyngbya • No difference in pigments among treatments • Chl a or b • No difference in photosynthetic efficiency among treatments
0.8 0.7 0.6 0.5 Photosynthetic efficiency 0.4 0.3 0.2 0.1 0 first last day Cladophora Photosynthetic efficiency significantly decreased at end
340 320 300 Carbon (mg/g) 280 live Dreissena empty Dreissena 260 pottery shards sand 240 220 200 day 1 day 2 day 7 Cladophora Nutrient tissue was significantly different between days, but not treatments
4.5 4 3.5 3 2.5 Phosphorus (mg/g) 2 1.5 40 1 live Dreissena 35 empty Dreissena 0.5 30 pottery shards sand 0 25 20 Nitrogen (mg/g) 15 10 5 0 day 1 day 2 day 7 Cladophora Trend of higher nutrient concentration with live Dreissena
180 160 140 120 100 C:P ratio 80 60 40 20 0 live Dreissena empty Dreissena 16 pottery shards sand 14 12 10 8 6 C:N ratio 4 2 0 day 1 day 2 day 7 Cladophora No P deficiency, but N deficiency in all treatments • <143 is no P deficiency • >9.4 C:N is a N deficiency Kahlert 1998
30 25 20 15 Potassium (mg/g) 10 5 live Dreissena 0 empty Dreissena pottery shards sand 25 20 15 Sulfur (mg/g) 10 5 0 Cladophora K showed more distinct pattern at end, but was not significantly different day 1 day 2 day 7
45 40 35 30 200 Calcium (mg/l) 25 20 150 15 Calcium (mg/g) 10 100 5 0 50 0 day 7 day 1 day 2 Cladophora Marginally significantly difference in tissue calcium, significant decrease in water initial end
Summary for Cladophora • Showed a biomass increase in the live Dreissena treatment • No significant difference among pigments • Showed trends of higher nutrient concentrations in live Dreissena treatment • C, N, P, K, S • Dreissena decreased Ca concentration • **Not yet received results for 13C and 15N • Did not respond to structure
Overall Conclusions: growth • Dreissena increased growth in Cladophora, but not Lyngbya • Cladophora responded to increased nutrients • Dilution • Attachment
Overall Conclusions: growth • Dreissena increased growth in Cladophora, but not Lyngbya • Cladophora responded to increased nutrients • Dilution • Attachment • Not enough time for Lyngbya • Lyngbya growth does not always positively respond to N and P • Increased PC and nutrients indicate that Lyngbya was photosynthetically healthier with Dreissena
Overall Conclusions: nutrients • Dreissena can retain nutrient concentrations in benthic algae • Can translate to increase biomass over time • More than just C, N, P • Higher nutrient content can mean higher quality of algae for grazers • Decrease Ca concentration, but unlikely to affect algae in natural system
Overall Conclusions: structure • Not found to be as important as nutrient contributions • No difference among empty shell, pottery shard, or sand alone treatments • Algae were floating in the water • Cladophora bulk biomass was not able to reattach • More attachment with live Dreissena
Implications • Dreissena increased algal biomass of one species and contributed several important nutrients to benthic algae • Supports nearshore shunt hypothesis
Implications • Dreissena increased algal biomass of one species and contributed several important nutrients to benthic algae • Nutrient reduction policies help control algal blooms, but Dreissena aggregate N and P which benefits benthic algae • May promote blooms despite reduced loading • Target nutrient levels may have to be lower than previously believed • Nutrient reductions may be ineffective • Dreissena help benthic algae acquire C • Benthic algae have a low affinity for nutrients in water column
Implications • Dreissena increased algal biomass of one species and contributed several important nutrients to benthic algae • Nutrient reduction policies help control algal blooms, but Dreissena aggregate N and P which benefits benthic algae • Algae can store nutrients • Helpful when nutrient availability is low • Create longer presence of algae
Acknowledgements • Advisor: Dr. Christine Mayer • Committee: Dr. Scott Heckathorn, Dr. Thomas Bridgeman, Dr. Rex Lowe • Department of Environmental Sciences and the Lake Erie Center • Dr. Jonathan Frantz & USDA ARS team at UT • Plant Science Research Center • Lab/Field/Other Help • Mike Bur and Patrick Kocovsky, USGS • Kristen DeVanna, Justin Chaffin, Sasmita Mishra, Dominic Armenio, Sarah Panek, Shellie Phillips, Rachel Kuhanek, Nate Manning, Peter Bichier, Mike Kuebbeler, Todd Crail, Chris Bronish, Sam Guffey, Blake Quinton, Steve Timmons, Jeremy Pritt, Dale Lorenzen III • Funding sources • Lake Erie Protection Fund, SG 384-10 • EPA Lake Erie Algal Source Tracking