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1 EFFECT of DISTURBANCE on ECOSYSTEM PROCESSES By EFFECT OF DISTURBANCE ON ECOSYSTEM PROCESSES By SEUNGJUN LEE A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA 2010 1 © 2010 Seungjun Lee 2 To my family 3 ACKNOWLEDGMENTS I owe my deepest gratitude to Dr. Mark Brown for his guidance as an advisor during my PhD program. His great passion and unique philosophy on advisement have driven me to freely explore the intellectual world, develop my own research project, and mature as an independent scholar over the last four and a half years. The completion of this dissertation would not have been possible without his intellectual and financial support. I was fortunate to have as one of my committee members Dr. Clay Montague, who was always delighted to provide me with invaluable ideas and suggestions on microcosm experiments. His extensive knowledge and experience on microcosms were crucial to the development and completion of my research project. Dr. Matt Cohen enriched my research project with original questions and suggestions, and encouraged me to acquire diverse theoretical perspectives. Dr. Paul Zwick guided me in the right direction until the completion of the program with fundamental questions and comments regarding my theories and thoughts. My system-oriented thinking has been fostered by teaching and advisement from Drs. Brown, Montague, Cohen, and Zwick, who have been the best systems ecologists I have ever met. I have been honored to work with all of my previous and current colleagues of the Systems Ecology group at the University of Florida. Diverse thoughts and discussions they shared with me in the systems seminars, research meetings, and individual talks have been cornerstones for this dissertation. In particular, David Pfahler helped me by reviewing and commenting on the final revision of this dissertation. Most importantly, I would like to thank my family. I thank my parents for their patience and support with unconditional love. I would like to address my special thanks to Hyewon for her love, ideas, and emotional support as a wife, colleague, and friend 4 throughout my doctoral study. She shared the best and worst moments of the journey of the last several years with me, and her never-failing support and encouragement have been essential to the completion of this dissertation. The birth and growth of my son, Shimok, during the last year and a half have given me great joy and inspiration for the self-organization of man and nature. The construction of the microcosm systems was supported in part by Glick and Wilkes scholarships awarded through the department of Environmental Engineering Sciences at the University of Florida. I thank Sejin Youn, who is working with Dr. Jean- Claude Bonzongo, for gladly providing me with purified water for the maintenance of microcosms. Charles H. Meyer, a tutor at the University of Florida Reading and Writing Center, helped me with proofreading during the revision of this dissertation. 5 TABLE OF CONTENTS page ACKNOWLEDGMENTS.................................................................................................. 4 LIST OF TABLES............................................................................................................ 9 LIST OF FIGURES........................................................................................................ 10 LIST OF ABBREVIATIONS........................................................................................... 13 ABSTRACT ................................................................................................................... 14 CHAPTER 1 INTRODUCTION .................................................................................................... 16 Statement of the Problem ....................................................................................... 16 Research Questions ............................................................................................... 17 Studies on Ecological Disturbance ......................................................................... 18 Progress in the Theory of Ecological Disturbance............................................ 18 Disturbance and Ecosystem Structures ........................................................... 22 Disturbance and Ecosystem Processes ........................................................... 26 Water Motion Disturbance................................................................................ 28 Ecosystem Theories and Disturbance .................................................................... 29 Self-Organization and Hypotheses on Ecosystem-Level Strategies................. 29 Ecosystem Traits during Succession................................................................ 31 Pulsing Paradigm ............................................................................................. 34 Microcosm Studies ................................................................................................. 35 Microcosm Overview ........................................................................................ 35 Ecosystem-Level Traits and Measurements..................................................... 38 A pH Method for Continuous Monitoring of Ecosystem-Level Metabolism ....... 39 Simulation Models of Disturbance .......................................................................... 40 Research Plan ........................................................................................................ 43 2 METHODS.............................................................................................................. 51 Microcosm Experiments.......................................................................................... 52 System Design ................................................................................................. 52 Sampli ng .......................................................................................................... 53 Maintenance and Measurements ..................................................................... 53 Disturbance Regimes ....................................................................................... 55 Data Processing and Analysis ................................................................................ 57 Data Processing: From Raw Data to Gross Primary Productivity (GPP) and Ecosystem Respiration Rate (ER)................................................................. 57 Ranges of pH and Alkalinity ............................................................................. 60 6 Analysis of Disturbance Intensity and Frequency Effects................................. 61 Analysis of GPP and ER Pulsing Pattern ......................................................... 62 Analysis of Resistance and Resilience............................................................. 63 Analysis of Uncertainty in the Resultant Patterns............................................. 64 Simulation Models .................................................................................................. 65 Model of a Microcosm ...................................................................................... 66 Simulation of Disturbance................................................................................. 66 Validation of Models ......................................................................................... 68 3 RESULTS ............................................................................................................... 78 Structures and Processes of Microcosms............................................................... 78 Microcosm Features......................................................................................... 78 Ranges of pH and Alkalinity ............................................................................. 79 GPP and ER of Microcosms............................................................................. 80 Effect of Disturbance Intensity ................................................................................ 81 GPP and Disturbance Intensity ........................................................................ 81 Pulsing Patterns of GPP and ER under Disturbance........................................ 84 Effect of Disturbance Frequency............................................................................. 85 Effect of Disturbance under Different Ecosystem Maturities................................... 87 Relationship between MGPP and Disturbance Regime...................................... 88 Resistance and Resilience of GPP and ER...................................................... 90 GPP and ER Pulsing ........................................................................................ 90 Uncertainty of Resultant Patterns ........................................................................... 91 Uncertainty from [GCO2] Correction ................................................................. 91 Uncertainty from pH Measurement Error.......................................................... 93 Simulation Models .................................................................................................. 95 Model Equations and Calibrations.................................................................... 95 Simulation Results............................................................................................ 95 Validation of the Models ..................................................................................
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