1. 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

2. Increased light penetration System level Benthic Primary Production Local level Resource importation Structural complexity What are local-scale interactions between Dreissena and benthic algae?

3. Contribution to algae N & P –Ammonium and phosphate CO 2 –Respiration –Decomposition Feces and pseudofeces Organic matter Other nutrients? Structure –Hard shells increase surface area P. Bichier

4. 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?

5. 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? J. Joyner P. Bichier

6. 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

7. 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.

8. 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

9. Experimental set-up

10. Live Dreissena (N=10) Empty Dreissena (N=10) Sand (N=10) Lyngbya Experiment 4 treatments 3400 Dreissena/m 2 230 g/m 2 Lyngbya Low-P lake water 12 hour photoperiod One week N=40 Pottery shards (N=10)

11. Fed Dreissena throughout experiment Chlamydomonas reinhardtii –Phytoplankton –Labeled with stable isotope 13 C 15 N –Given to all tanks

12. 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 13 C and 15 N –Samples sent to UC Davis, CA

13. Positive correlation between wet weight and dry weight y = 0.1538x + 0.1367 R 2 = 0.798 0 0.5 1 1.5 2 2.5 3 05101520 wet weight (g) dry weight (g) p<0.0001 Lyngbya

14. All tanks increase in weight, but no difference among treatments 0 0.5 1 1.5 2 2.5 live Dreissena empty Dreissena pottery shards sand substrate type Dry weight (g) 0 10 20 30 40 50 60 70 % increase in wet weight Error bars = SE Lyngbya

15. No difference in photosynthetic efficiency, but decreases with more dry weight substrate type 0 0.05 0.1 0.15 0.2 0.25 0.3 0.35 0.4 0.45 0.5 live Dreissena empty Dreissena pottery shards sand Photosynthetic efficiency y = -2.8267x + 2.9382 R 2 = 0.3987 p<0.0001 0 0.5 1 1.5 2 2.5 3 00.10.20.30.40.50.60.7 photosynthetic efficiency dry weight (g) Lyngbya

16. No difference in chlorophyll a, but higher phycocyanin in live Dreissena treatment 0 50 100 150 200 250 300 350 400 450 500 Chlorophyll a (µg/g) 0 1000 2000 3000 4000 5000 6000 live Dreissena empty Dreissena pottery shards sand substrate type Phycocyanin (µg/g) A C BC AB Lyngbya

17. Dreissena increased carbon content in tissue 0 50 100 150 200 250 300 350 400 live Dreissena empty Dreissena pottery shards sand substrate type Carbon (mg/g) A BBB Lyngbya

18. Dreissena helped retain macronutrient content in tissue 0 5 10 15 20 25 30 35 40 45 live Dreissena empty Dreissena pottery shards sand substrate type Nitrogen (mg/g) 0 1 2 3 4 5 6 Phosphorus (mg/g) A B B B A B BB Lyngbya

19. No P deficiency, but N deficiency in all treatments except live Dreissena <143 is no P deficiency >9.4 C:N is a N deficiency –No deficiency in live Dreissena treatment 0 2 4 6 8 10 12 C:N ratio live Dreissena empty Dreissena pottery shards sand substrate type 0 20 40 60 80 100 120 C:P ratio A B B B A BBB Kahlert 1998 Lyngbya

20. Dreissena helped retain nutrient content in tissue 0 0.5 1 1.5 2 2.5 3 3.5 4 Potassium (mg/g) 0 0.5 1 1.5 2 2.5 3 3.5 4 live Dreissena empty Dreissena pottery shards sand substrate type Sulfur (mg/g) A B B B A B B B Lyngbya

21. However… 0 20 40 60 80 100 120 140 160 live Dreissena empty Dreissena pottery shards sand substrate type Calcium (mg/g) A B B B Lyngbya

22. Stable isotope: less 13 C in live Dreissena treatment, no significant difference in 15 N -25 -20 -15 -10 -5 0 live Dreissena empty Dreissena pottery shards sand substrate type δ 13 C (‰) A B BB Lyngbya

23. Two week preliminary experiment with Cladophora: the 13 C difference goes away and there becomes a significant difference in 15 N 0 0.5 1 1.5 2 2.5 3 live Dreissena empty Dreissena pottery shards sand substrate type δ 15N (‰) A B B B

24. Summary for Lyngbya Dreissena did not increase the biomass of Lyngbya Increased phycocyanin Supplied several nutrients, prevented a decrease in others Decreased Ca Decreased 13 C Did not respond to substrates

25. Live Dreissena (N=10) Empty Dreissena (N=10) Sand (N=10) Cladophora Experiment 4 treatments 3400 Dreissena/m 2 160 g/m 2 algae Low-P lake water One week 12 hour photoperiod N=40 Fed Chlamydomonas Pottery shards (N=10)

26. 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

27. y = 0.1387x + 0.4254 R2R2 = 0.6933 0 0.5 1 1.5 2 2.5 3 024681012141618 wet weight (g) dry weight (g) p<0.0001 Positive correlation between wet weight and dry weight Cladophora

28. All tanks increase in weight, significant difference when other treatments are grouped 0 10 20 30 40 50 60 70 80 % increase in wet weight live Dreissena empty Dreissena pottery shards sand substrate type 0 2 4 6 8 10 12 wet weight (g) bottom biomass B A Cladophora

29. 2 0 0.2 0.4 0.6 0.8 1 1.2 1.4 1.6 1.8 live Dreissena empty Dreissena pottery shards sand substrate type dry weight (g) bottom biomass Separation in biomass, more with live Dreissena Cladophora

30. Cladophora Results More variability than Lyngbya No difference in pigments among treatments –Chl a or b No difference in photosynthetic efficiency among treatments

31. Photosynthetic efficiency significantly decreased at end 0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 firstlast day Photosynthetic efficiency Cladophora

32. Nutrient tissue was significantly different between days, but not treatments live Dreissena empty Dreissena pottery shards sand Cladophora

33. Trend of higher nutrient concentration with live Dreissena live Dreissena empty Dreissena pottery shards sand day 1day 2day 7 0 5 10 15 20 25 30 35 40 Nitrogen (mg/g) Cladophora

34. No P deficiency, but N deficiency in all treatments <143 is no P deficiency >9.4 C:N is a N deficiency Kahlert 1998 live Dreissena empty Dreissena pottery shards sand Cladophora

35. K showed more distinct pattern at end, but was not significantly different 0 5 10 15 20 25 30 Potassium (mg/g) 0 5 10 15 20 25 Sulfur (mg/g) live Dreissena empty Dreissena pottery shards sand day 1day 2day 7 Cladophora

36. Marginally significantly difference in tissue calcium, significant decrease in water 0 50 100 150 200 day 1day 2 day 7 Calcium (mg/g) 0 5 10 15 20 25 30 35 40 45 Calcium (mg/l) initialend Cladophora

37. 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 13 C and 15 N Did not respond to structure

38. Overall Conclusions: growth Dreissena increased growth in Cladophora, but not Lyngbya –Cladophora responded to increased nutrients Dilution Attachment

39. 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

40. 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

41. 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

42. Implications Dreissena increased algal biomass of one species and contributed several important nutrients to benthic algae –Supports nearshore shunt hypothesis

43. 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

44. 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

45. 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