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Occurring Δ2h and Δ13c Variation in a Marine Predator, The Fall 08 UNDERSTANDING EXPERIMENTALLY-INDUCED AND NATURALLY- OCCURRING δ2H AND δ13C VARIATION IN A MARINE PREDATOR, THE BROWN BOOBY A Thesis Presented to The Graduate Faculty of the University of Akron In Partial Fulfillment of the Requirements for the Degree Master of Biology Madelyn Jacobs May, 2020 Fall 08 UNDERSTANDING EXPERIMENTALLY-INDUCED AND NATURALLY- OCCURRING δ2H AND δ13C VARIATION IN A MARINE PREDATOR, THE BROWN BOOBY Madelyn Jacobs Thesis Approved: Accepted: Advisor Department Chair Dr. Anne Wiley Dr. Steve Weeks Committee Member Dean of the College Dr. Brian Bagatto Dr. Linda Subich Committee Member Acting Dean of the Graduate School Dr. Randall Mitchell Dr. Marnie Saunders Date ii ABSTRACT Refining our understanding of the sources of δ2H variation in terrestrial and marine organisms is a necessary step to expand the usefulness of stable hydrogen isotope values in ecological studies. Marine consumers in particular provide a simplified hydrogen isotope study system in which to test hypotheses concerning non-geographical sources of δ2H variation. The primary goal of our study was to test hypotheses about the source of hydrogen isotope variation in seabirds using manipulative experiments in brown booby chicks and naturally occurring variation in δ2H along a continuum of ages in a colony of brown booby (Sula leucogaster). First, we hypothesized that increased salt ingestion will increase salt gland activity and proportionately more protium will be expelled in salt gland excretions, causing the δ2H of seabird tissues to increase. Second, we hypothesized that increased lipid ingestion will result in proportionately more incorporations of dietary-lipid derived hydrogen into seabird tissues, resulting in lower δ2H values. A third, related hypothesis regarding carbon isotope variation was tested simultaneously: increased lipid ingestion will result in proportionately more incorporation of dietary-lipid derived carbon into seabird tissues, resulting in lower δ13C values. To test these hypotheses, a population of brown booby (Sula leucogaster) chicks was tube-fed one of two treatments for 14 consecutive days: a high salt solution (3% NaCl), or a high lipid solution (50% salmon oil). Two groups of chicks were tube-fed normal saline solution (0.9% NaCl) as control groups alongside the treatment groups. We then measured the hydrogen, carbon, and nitrogen isotopic composition (δ2H, δ13C, and δ15N) of the plasma prior to and after the 14-day manipulation. The change in δ2H plasma iii values was significantly larger for chicks in the high salt ingestion treatment group compared to the salt control group (0=0.036). The change in plasma δ2H of chicks receiving the high lipid solution treatment did not differ statistically from the change in plasma δ2H observed in the control group (p=0.0702). High salt or lipid ingestion, however, had no apparent effect on δ13C or δ15N plasma values compared to the chicks in the control groups. Our study further aimed to understand whether δ2H values change along a continuum of ages in a seabird species and if amino acid-specific isotope analysis could illuminate the mechanism behind widely reported age-related shifts in δ2H. Therefore, we compared bulk plasma δ2H values from brown booby chicks to corresponding amino acid-specific δ2H analyses to understand whether some amino acids may dictate patterns in bulk δ2H. In addition, we explored how age affected bulk and amino acid-specific δ2H values within the brown booby colony. We found several lines of evidence that age is an important source of δ2H variation in the brown booby. Importantly, none of the amino acid δ2H values correlated significantly with chick age in our study. We propose that changes in the amino acid composition of plasma and newly synthesized tissues may cause bulk δ2H values to change with age. To our knowledge, this is the first reported investigation of avian amino acid-specific δ2H values. iv ACKNOWLEDGMENTS I would like to acknowledge my advisor, Dr. Anne Wiley, for her unwavering support, optimism, and guidance throughout my master’s program. Her wisdom and thoughtfulness on experimental design and data interpretation were invaluable. In addition, I would like to acknowledge my collaborators that made sample collection and analysis possible. This study was completed in collaboration with Dr. Roxana Torres and Gala Castro Mejias from the National Autonomous University of Mexico, and Dr. Kaycee Morra and Dr. Marilyn Fogel from the University of California, Riverside. Lastly, I would like to thank my lab mates at the University of Akron for their help in sample analysis and data interpretation: Nathan Michael and Ryan Trimbath (PhD candidates), Allison Carpenter and Erin Taylor (undergraduate students). v TABLE OF CONTENTS Page LIST OF FIGURES..........................................................................................................viii LIST OF TABLES..............................................................................................................ix CHAPTER I. UNDERSTANDING THE ISOTOPIC INFLUENCE OF SALT AND LIPID INGESTION IN BROWN BOOBY (SULA LEUCOGASTER) CHICKS ON ISLA MARIETAS, MEXICO....................................................1 INTRODUCTION..........................................................................................1 MATERIALS AND METHODS Experimental design....................................................................................8 Sample collection and analysis..................................................................11 Statistical analyses.....................................................................................12 RESULTS Change in chick plasma hydrogen isotope values.....................................13 Change in chick plasma carbon and nitrogen isotope values....................17 Change in chick body mass.......................................................................17 DISCUSSION The effect of salt ingestion on δ2H............................................................18 The effect of lipid ingestion on δ2H...........................................................20 The effect of lipid ingestion on δ13C..........................................................21 SUMMARY..................................................................................................23 vi II. EFFECTS OF AGE AND PATTERNS OF AMINO ACID-SPECIFIC δ2H VALUES IN A COLONY OF BROWN BOOBIES (SULA LEUCOGASTER)............................................................................................24 INTRODUCTION........................................................................................24 METHODS Plasma sample collection and preparation.................................................27 Prey sample collection and preparation.....................................................28 Bulk sample analysis..................................................................................29 Amino acid-specific analysis.....................................................................30 Reporting δ2H............................................................................................30 Statistical analyses.....................................................................................30 RESULTS Chick plasma bulk δ2H and chick age.......................................................31 Prey muscle δ2H.........................................................................................34 Amino acid-specific chick plasma.............................................................34 DISCUSSION δ2H and age................................................................................................37 Amino acid-specific δ2H............................................................................43 LITERATURE CITED......................................................................................................45 APPENDIX........................................................................................................................55 vii LIST OF FIGURES FIGURE PAGE CHAPTER 1 1. Tube-feeding 50% salmon oil solution to a chick in the lipid treatment group........................................................................................................................8 2. Mean change in isotopic composition of brown booby chick plasma post- manipulation...........................................................................................................15 CHAPTER 2 1. δ2H values of prey muscle and brown booby plasma and feather samples...........33 2 2. Brown booby chick plasma amino acid δ H values.................................................36 viii LIST OF TABLES CHAPTER 1 Page 1. Description of tube-feeding treatment solutions and sample sizes..........................9 2. Summary statistics of brown booby chick plasma isotopic change post- manipulation and supplementary sample δ2H values............................................16 CHAPTER 2 1. Mean and standard deviation of δ2H values for 13 amino acids from brown booby chick plasma...........................................................................................................35 ix CHAPTER 1 UNDERSTANDING THE ISOTOPIC INFLUENCE OF SALT AND LIPID INGESTION IN BROWN BOOBY (SULA LEUCOGASTER) CHICKS ON ISLA MARIETAS, MEXICO INTRODUCTION Changing climate and industrialized fishing are shifting marine food web dynamics and may be partially responsible for the alarming global decline of seabird populations (69% in 50 years, Paleczny et
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