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Morphological & Physiological Leaf Adaptations

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Morphological & Physiological Leaf Adaptations MORPHOLOGICAL AND PHYSIOLOGICAL LEAF ADAPTATIONS TO SEASONAL AND DIURNAL ABIOTIC STRESS FOR TWO BARRIER ISLAND SAND DUNE SPECIES By HEATHER M. JOESTING A Dissertation Submitted to the Graduate Faculty of WAKE FOREST UNIVERSITY GRADUATE SCHOOL OF ARTS AND SCIENCES In Partial Fulfillment of the Requirements for the Degree of DOCTOR OF PHILOSOPHY in the Department of Biology August 2009 Winston-Salem, NC Approved by: William K. Smith, Ph.D., Advisor Examining Committee: Miles R. Silman, Ph.D. Ronald V. Dimock, Jr., Ph.D. Kathleen A. Kron, Ph.D. Tara L. Greaver, Ph.D. ACKNOWLEDGEMENTS I would like to first thank my parents, Mel Pitts and Dale Joesting, for their love and support for the past decade of my college adventures as well as in all the decisions I have made for the last 31 years. I would also like to thank my wonderful supportive significant other Matt Marenberg for being my rock through the entire dissertation process and my research assistant during long hot days at the beach. I would also like to acknowledge my grandfather Elden Lee Pitts who instilled the love of science and nature in my mother and I and whose stubborn and inquisitive mind I’ve inherited. Without the love and support of these people, I would never have made it this far. I would also like to thank my advisor William K. Smith for all the guidance he has given me during this dissertation research, for always being available to answer any question I had, no matter how trivial, and for giving me the opportunity to conduct research in a habitat so near and dear to my heart. I also thank my committee members, Drs. Ronald V. Dimock, Jr., Miles R. Silman, Kathy A. Kron, and Tara L. Greaver for all their assistance and advice. These people have made me a better scientist, and I have learned so much from them. Also, thanks to the members of the Smith lab past and present, namely Z. Carter Berry for always being up for a sushi lunch and his many reviews of manuscripts and graphs, Joseph White for assistance in the field, and Cristin Walters for always being up for a “chat” break, as well as the many undergraduates who assisted in the field and the laboratory. I’d also like to thank my extended family of friends here in Winston Salem who have made the past five years some of the best in my life. Finally, I’d like to thank my two cats, Winston and Taylor, for laying on my computer when I tried to write and laying on articles when I tried to read, and my dog Tela for being the best companion a girl could ask for on my frequent trips to the beach. ii TABLE OF CONTENTS ACKNOWLEDGEMENT……………………………………………………………...………….ii LIST OF TABLES…………………………………………………………………………………v LIST OF FIGURES……………………………………………………………………………....vii ABSTRACT……………………………………………………………………………………….x CHAPTER I An introduction to barrier islands and sand dune ridge and swale systems……………....1 CHAPTER II Introduction to study site and species……………………………………………………30 CHAPTER III Seasonal and diurnal leaf orientation, bifacial sunlight incidence, and leaf structure in the sand dune herb Hydrocotyle bonariensis ABSTRACT………………………………………………………………….…53 INTRODUCTION………………………………………………………………54 MATERIALS AND METHODS……………………………………………….55 RESULTS…………………………………………………………………….....60 DISCUSSION…………………………………………………………...………63 CHAPTER IV Influence of leaf orientation on leaf photosynthetic gas exchange in the sand dune herb Hydrocotyle bonariensis ABSTRACT…………………………………………………………………….86 INTROUDCTION………………………………………………………………87 MATERIALS AND METHODS……………………………………………….88 RESULTS……………………………………………………………………….92 DISCUSSION…………………………………………………………………...94 iii CHAPTER V Do leaf characteristics predict leaf photosynthetic gas exchange along a small-scale stress gradient? ABSTRACT………………………………………………………………….108 INTRODUCTION…………………………………………………………....109 MATERIALS AND METHODS…………………………………………….111 RESULTS…………………………………………………………………….116 DISCUSSION………………………………………………………………...118 CHAPTER VI Influence of leaf structure, leaf orientation, and abiotic environment on leaf physiology in two barrier island sand dune species with different growth habits ABSTRACT…………………………………………………………………..141 INTRODUCTION…………………………………………………………….142 MATERIALS AND METHODS……………………………………………..143 RESULTS……………………………………………………………………..150 DISCUSSION…………………………………………………………………154 CHAPTER 7 Conclusions and future research………………………………………………………..177 CURRICULUM VITAE………………………………………………………………………...185 iv LIST OF TABLES Table III – 1: Predicted and measured leaf structure and orientation for Hydrocotyle bonariensis based on conceptual Structure-Orientation model………………………………..……..73 Table III – 2: Effect of time and month on leaf orientation and incident sunlight level on adaxial and abaxial leaf surfaces for Hydrocotyle bonariensis..…..………………………….…74 Table III – 3: Relationships between leaf orientation and incident sunlight on adaxial and abaxial leaf surfaces for Hydrocotyle bonariensis………………………………..…………...…75 Table IV – 1: Mean leaf temperature of control inclined and experimental horizontal Hydrocotyle bonariensis leaves…………………………….………………………………………...100 Table V – 1: Predicted and measured leaf structure and orientation for Hydrocotyle bonariensis and Iva imbricata in the sand dune habitat……………………………………………..129 Table V – 2: Comparison of leaf structure, leaf temperature, and leaf photosynthetic gas exchange between Hydrocotyle bonariensis and Iva imbricata…………………………………..130 Table V – 3: Best fit models for predicting leaf temperature and leaf photosynthetic gas exchange from leaf structure for Hydrocotyle bonariensis and Iva imbricata……………………131 Table VI – 1: Mean incident sunlight, air temperature, vapor pressure deficit, and sand temperature in 2009 for Topsail Island, NC………………………………...………….165 Table VI – 2: Comparison of leaf structure and orientation between Hydrocotyle bonariensis and Iva imbricata……………………………………………………………………………166 Table VI – 3: Comparison of incident sunlight on adaxial and abaxial leaf surfaces, leaf temperature, leaf photosynthetic gas exchange, and photosystem II efficiency between Hydrocotyle bonariensis and Iva imbricata…………………………………………….167 Table VI – 4: Effect of time and month on incident sunlight on adaxial and abaxial leaf surfaces and photosystem II efficiency for Hydrocotyle bonariensis and Iva imbricata………..168 v Table VI – 5: Effect of time and month on leaf temperature and photosynthetic gas exchange for Hydrocotyle bonariensis and Iva imbricata…………………………………………….169 Table VI – 6: Best fit models for predicting leaf temperature, leaf photosynthetic gas exchange, and photosystem II efficiency from leaf structure and abiotic environment for Hydrocotyle bonariensis and Iva imbricata…………………………………………….170 vi LIST OF FIGURES Figure I – 1: Developing sand dunes on Topsail Island, NC………………………………….....25 Figure I – 2: Effect extreme episodic storm events on sand dune communities...……………….26 Figure I – 3: Characteristic vegetation zones of mixed energy mesotidal barrier islands………..27 Figure I – 4: Negative impacts of seawall at Galveston Island, TX……………………………...28 Figure I – 5: Vegetated sand dunes in front of beach homes at Topsail Island, NC before and after a coastal storm…………………………………………………………………………...29 Figure II – 1: Location of study site at Topsail Island, NC………………………………………44 Figure II – 2: Species composition of the sand dune vegetation zones at Topsail Island, NC…...45 Figure II – 3: Mean species diversity and species richness of sand dune vegetation zones at Topsail Island, NC…………………………………………………………………….…46 Figure II – 4: Growth habit of Hydrocotyle bonariensis in the sand dune system at Topsail Island, NC………………………………………………………………………………………..47 Figure II – 5: Fresh leaf cross-sections of Hydrocotyle bonariensis and Iva imbricata………….48 Figure II – 6: Stomata impressions of abaxial and adaxial leaf surfaces of Hydrocotyle bonariensis and Iva imbricata…………………………………………………………...49 Figure II – 7: Growth habit of Iva imbricata in the sand dune system at Topsail Island, NC…...50 Figure II – 8: Scanning electron microscope image of surface topography and trichomes in Iva imbricata…………………………………………………………………………………51 Figure III – 1: Growth habit of Hydrocotyle bonariensis in the sand dune system at Topsail Island, NC………………………………………………………………………………………..78 Figure III – 2: Illustration of methodology of measuring leaf angle and leaf azimuth…………...79 Figure III – 3: Diurnal pattern of leaf angle and leaf azimuth in Hydrocotyle bonariensis………80 Figure III – 4: Seasonal pattern of leaf angle and leaf azimuth in Hydrocotyle bonariensis……..81 vii Figure III – 5: Diurnal and seasonal pattern of sunlight incidence on adaxial and abaxial leaf surfaces in Hydrocotyle bonariensis……………………………………………….…….82 Figure III – 6: Fresh leaf cross-section and stomata impression of Hydrocotyle bonariensis…....83 Figure III – 7: Illustration of seasonal leaf orientation of Hydrocotyle bonariensis in relation to diurnal sun path…………………………………………………………………………..84 Figure IV – 1: Experimental setup of forced horizontal leaves and insertion of leaf thermocouples in Hydrocotyle bonariensis……………………………………………………………..102 Figure IV – 2: Diurnal leaf temperature of inclined control and experimental horizontal Hydrocotyle bonariensis leaves………………………………………………………...103 Figure IV – 3: Difference between leaf temperature and air temperature for inclined control and experimental horizontal Hydrocotyle bonariensis leaves………………………………104 Figure IV – 4: Mean midday photosynthetic gas exchange for inclined control and experimental horizontal Hydrocotyle bonariensis leaves……………………………………………..105 Figure IV – 5: Mean midday leaf
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