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INFLUENCE OF ALGAL PRODUCTION ON ECOSYSTEM METABOLISM, MICROBIAL ACTIVITY AND NUTRIENT DYNAMICS IN A CENTRAL INDIANA STREAM A THESIS SUBMITTED TO THE GRADUATE SCHOOL IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE MASTER OF SCIENCE BY ANDREA S. FITZGIBBON ADVISORS: DR. MELODY J. BERNOT & DR. KEVIN H. WYATT BALL STATE UNIVERSITY MUNCIE, INDIANA JULY 2014 INFLUENCE OF ALGAL PRODUCTION ON ECOSYSTEM METABOLISM, MICROBIAL ACTIVITY AND NUTRIENT DYNAMICS IN A CENTRAL INDIANA STREAM A THESIS SUBMITTED TO THE GRADUATE SCHOOL IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE MASTER OF SCIENCE BY ANDREA S. FITZGIBBON ADVISORS: DR. MELODY J. BERNOT & DR. KEVIN H. WYATT Committee Approval: ___________________________________________________________ ____________ Committee Chairperson Date ___________________________________________________________ ____________ Committee Member Date ___________________________________________________________ ____________ Committee Member Date Departmental Approval: ___________________________________________________________ ____________ Departmental Chairperson Date ___________________________________________________________ ____________ Dean of Graduate School Date BALL STATE UNIVERSITY MUNCIE, INDIANA JULY 2014 TABLE OF CONTENTS TABLE OF CONTENTS…………………………………………………………………..…….iii LIST OF FIGURES CHAPTER 1………………………………………………………...….…..iv LIST OF TABLES CHAPTER 1………………………………………………………...….…...vi LIST OF FIGURES CHAPTER 2……………………………………………………………….vii LIST OF TABLES CHAPTER 2………………………………………………………………...ix ABSTRACT……………………………………………………………………………………….1 ACKNOWLEGEMENTS…………………………………………………………………………3 CHAPTER 1: Comparison of algal and ecosystem metabolism and nutrient dynamics in a central Indiana stream ABSTRACT………………………………………………………………………………….……4 INTRODUCTION………………………………………………………………………….……..6 METHODS……………………………………………………………………………….……….8 RESULTS…………………………………………………………………………….………….14 DISCUSSION…………………………………………………………………….……………...21 CONCLUSIONS………………………………………………………………….…………......26 REFERENCES…………………………………………………………………….………….....27 FIGURES AND TABLES…………………………………………………………….………....32 CHAPTER 2: Influence of algal exudates on microbial activity and ecosystem dynamics ABSTRACT………………………………………………………………………………….…..44 INTRODUCTION………………………………………………………………………….……46 METHODS……………………………………………………………………………….……...48 RESULTS…………………………………………………………………………….………….53 DISCUSSION…………………………………………………………………….……….……..56 CONCLUSIONS………………………………………………………………….…………......59 REFERENCES…………………………………………………………………….………….....61 FIGURES AND TABLES…………………………………………………………….………....66 APPENDIXES…………………………………………………………………………………...75 iii LIST OF FIGURES Figure Page CHAPTER 1: Comparison of algal and ecosystem metabolism and nutrient dynamics in a central Indiana stream 1. Conceptual model of hypothesized algal mediated diurnal patterns of nutrients, (top panels) dissolved organic carbon (DOC), dissolve inorganic nitrogen (DIN); physiochemistry, (middle panels) dissolved oxygen (O ) and pH; and metabolism, 2 (bottom panels) primary production, respiration and decomposition. ……………….….32 2. Conceptual model of hypothesized seasonal water column and algae patterns of dissolved oxygen (top panel); nutrients (middle panel); and metabolism, (bottom panels) primary production and respiration. ……………………………………………………………..33 3. Water column and algae diurnal variation in dissolved oxygen (DO, mg O /L) 2 concentration and pH over time in July and December 2013. ………………………….34 4. Mean seasonal water column and algae dissolved oxygen concentrations. (n= 9 days per season; Winter = Dec - Feb, Spring = Mar - May, Summer = Jun - Aug, Fall = Sep - Nov). Error bars indicate SD. Dashed line indicates a 1:1 relationship. ………………..37 5. Mean seasonal water column and algae pH. (n= 9 days per season; Winter = Dec - Feb, Spring = Mar - May, Summer = Jun - Aug, Fall = Sep - Nov). Error bars indicate SD. Dashed line indicates a 1:1 relationship. …………………………………………...…...38 6. Mean seasonal water column and algae total organic carbon concentrations. (n = 9 days per season; Winter = Dec - Feb, Spring = Mar - May, Summer = Jun - Aug, Fall = Sep - Nov). Error bars indicate SD. Dashed line indicates a 1:1 relationship. …………….…39 7. Mean seasonal water column and algae nitrate concentrations. (n= 9 days per season; Winter = Dec - Feb, Spring = Mar - May, Summer = Jun - Aug, Fall = Sep - Nov). Error bars indicate SD. Dashed line indicates a 1:1 relationship. ……………………………40 8. Mean seasonal water column and algae phosphate concentrations. (n= 9 days per season; Winter = Dec - Feb, Spring = Mar - May, Summer = Jun - Aug, Fall = Sep - Nov). Error bars indicate SD. Dashed line indicates a 1:1 relationship. ……………………………41 iv 9. Mean daily rates of (A) GPP, (B) ER and (C) NEP measured in White River, Yorktown, Indiana from January through December 2013 (n = 2-3 days per month). Error bars indicate SD. Shaded area (A) indicated mean dry algal biomass (n = 12). ……………42 v LIST OF TABLES Table Page CHAPTER 1: Comparison of algal and ecosystem metabolism and nutrient dynamics in a central Indiana stream 1. Physiochemical characteristics of the water column (Water) and algae in the White River, Yorktown, Indiana from January through December 2013. Dissolved oxygen (DO), pH, temperature and conductivity were measured continuously at 10 minute intervals over approximately 72 hours (n = 125-432). Width (n = 1), flow and depth (n = 5) were measured during the sampling event. Discharge (n = 1) was calculated from width, flow and depth. Parentheses indicate standard error. …………………………………………35 2. Cumulative mean water column (Water) and algae, total organic carbon (TOC), nitrate (NO ), phosphate (PO ) and ammonium (NH ) concentrations. Phosphate concentrations 3 4 4 were the below detection limit (0.01 mg/L) September through December. Standard error in parentheses, N= 48-144. ……………………………………………………………...36 3. Multiple regression model covariates for water and algal gross primary production (GPP) and ecosystem respiration (ER). ……………………………………………....….……..43 vi . LIST OF FIGURES Figure Page CHAPTER 2: Influence of algal exudates and benthic substrata on microbial activity 1. Mean dissolved oxygen across mesocosm treatments (algae, exudate and control) over three sampling events (n = 126 for algae treatment, n = 63 for exudate and control treatments). Error bars indicate SE. Note y-axis begins at 7.00 mg O / L. Dissolved 2 oxygen differs by treatment (p = 0.027) but not by substrate (p= 0.241) with no interaction between main effects (p = 0.723). Pairwise differences indicated by letters above bars. ………………………………………………………………………………66 2. Mean pH of mesocosm (A) treatment (algae, exudate and control) and (B) substrate (sediment, leaves, sponge and glass) over three sampling events (n = 126 for algae treatment, n = 63 for exudate and control treatments, n = 60 for each substrate). Error bars indicate SE. Note y-axis begins at 8.00. pH differs by treatment and substrate (p < 0.001) with no interaction between main effects (p = 0.397). Pairwise differences indicated by letters above bars. ………………………………………………………….68 3. Mean total organic carbon (TOC) of mesocosm (A) treatment (algae, exudate and control) and (B) substrate (sediment, leaves, sponge and glass) over two sampling events (n = 32 for algae treatment, n = 16 for exudate and control treatments, n = 16 for each substrate). Error bars indicate SE. TOC differs by treatment (p < 0.0001) and substrate (p < 0.0001) with an interaction between main effects (p = 0.0073). Pairwise differences indicated by letters above bars. ……………………………………………………….…69 4. Mean nitrate (NO ) concentration by mesocosm (A) treatment (algae, exudate and 3 control) and (B) substrate (sediment, leaves, sponge and glass) over two sampling events (n = 64 for algae treatment, n = 32 for exudate and control treatments, n = 32 for each substrate). Error bars indicate SE. NO differs by treatment (p = 0.0002) and substrate (p 3 < 0.0001) with an interaction between main effects (p = 0.0003). Pairwise differences indicated by letters above bars. …………………………………………………………70 5. Mean phosphate (PO ) concentration by mesocosm (A) treatment (algae, exudate and 4 control) and (B) substrate (sediment, leaves, sponge and glass) over two sampling events (n = 64 for algae treatment, n = 32 for exudate and control treatments, n = 32 for each substrate). Error bars indicate SE. PO differs by treatment (p = 0.0003) and substrate (p 4 vii = 0.0003) with no interaction between main effects (p < 0.0001). Pairwise differences indicated by letters above bars. …………………………………………………………71 6. Nutrient ratios of total organic carbon (TOC) to nitrate (NO ) and phosphate (PO ) 3 4 among treatments (algae, exudate and control). Error bars indicate SE. TOC: NO did not 3 differ by treatment (p = 0.263) nor did TOC: PO (p = 0.126). Dashed lines indicate 4 differences in nutrient availability; arrows indicate high or low amounts of nutrient available. ………………………………………………………………………………...72 7. Mean respiration by mesocosm substrate type (sediment, leaves, sponge and glass; n = 20). Error bars indicate SE. Respiration differs by substrate (p < 0.001) but not by treatment (p = 0.997) with no interaction between main effects (p = 0.803). Pairwise differences indicated by letter above bars. ……………………………………………....73 8. Relationship between total organic carbon (TOC) concentration and microbial respiration by treatment (algae, exudate and control) (p < 0.01). ………………………………..….74 viii LIST OF TABLES Table Page CHAPTER 2: Influence of algal exudates and benthic substrata on microbial activity 1. Mean mesocosm temperature (°C) across algal and substrate treatments over three sampling events (n = 126 for algae treatment, n = 63 for exudate and control treatments and n = 60 for substrate treatments). Numbers in parentheses are SE. Temperature differs by treatment (p = 0.010) but not by substrate (p = 0.573) with no interaction between main effects (p = 0.371). Pairwise difference denoted by letters. ………………………67 ix ABSTRACT THESIS: Influence of algal production on ecosystem metabolism, microbial activity and nutrient dynamics in a central Indiana stream STUDENT: Andrea S. Fitzgibbon DEGREE: Masters of Science COLLEGE: Sciences and Humanities DATE: July 2014 PAGES: 155 One primary function in most ecosystems is photosynthesis which provides energy for consumers. In mid-order streams, benthic algae are primary producers fulfilling this role and their influence on stream biogeochemistry changes with the diurnal patterns of production. During the day, photosynthesis increases stream oxygen concentrations facilitating shifts in pH and nutrient concentrations. However, little is known about diurnal patterns of nutrient concentrations. Algal production regulates nutrient cycling through assimilation of inorganic nutrients and release of nitrogen and carbon exudates. Though nutrient assimilation by algae in streams is well understood, understanding of algal nutrient release is limited. Nutrient release by algae is a significant gap in our knowledge of ecosystem dynamics, especially in nutrient saturated streams. To understand algal dynamics in a nutrient saturated stream, algal production in a mid-order stream was measured via monthly sampling for one year using microelectrodes to quantify algal biofilm chemical gradients of dissolved oxygen and pH. Ecosystem energetics as gross primary production (GPP) and ecosystem respiration (ER) were also calculated. Nutrients were analyzed from water column and algal pore water samples. Dissolved oxygen concentrations and pH between the algae and water column followed similar diurnal curves although the amplitude of these changes varied seasonally. Nutrient and total organic carbon 1

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CHAPTER 1: Comparison of algal and ecosystem metabolism and nutrient .. streams is well understood, understanding of algal nutrient release is
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