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INFORMATION TO USERS This manuscript has been reproduced from the microfilm master. UMI films the text directly from the original or copy submitted. Thus, some thesis and dissertation copies are in typewriter face, wrule ot.~~rs may be from any type ofcomputer printer. The quality ofthis reproduction is dependent upon the quality ofthe copy submitted. Broken or indistinct print, colored or poor quality illustrations and photographs, print bleedthrough, substandard margins, and improper alignment can adversely affect reproduction. In the unlikely event that the author did not send UMI a complete manuscript and there are missing pages, these will be noted. Also, if unauthorized copyright material had to be removed, a note will indicate the deletion. Oversize materials (e.g., maps, drawings, charts) are reproduced by sectioning the original, beginning at the upper left-hand comer and continuing from left to right in equal sections with small overlaps. Each original is also photographed in one exposure and is included in reduced form at the back ofthe book. Photographs included in the original manuscript have been reproduced xerographically in this copy. Higher quality 6" x 9" black and white photographic prints are available for any photographs or illustrations appearing in this copy for an additional charge. Contact UMI directly to order. UMI A Bell & Howell Information Company 300 North Zeeb Road, Ann Arbor MI 48106-1346 USA 313n61-4700 800/521-0600 --- - --_._-- . - .... -=,.- PHOTOADAPTIVE STRATEGIES OF HAWAllAN MACROALGAE A DISSERTATION SUBMITfED TO THE GRADUATE DMSION OF THE UNIVERSITY OF HAWAll IN PARTIAL FULFllLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY IN BOTANICAL SCIENCES (BOTANY) MAY 1996 by Kevin Scott Beach DISSERTATION COMMI'ITEE: Celia M. Smith, Chairperson Isabella A. Abbott Guillermo H. Goldstein Wayne T. Iwaoka Clifford W. Morden OMI Number: 9629809 UMI Microform 9629809 Copyright 19%. by UMI Company. Ail rights reserved. This microform edition is protected against unauthorized copying under Title 17, United States Code. UMI 300 North Zeeb Road Ann Arbor, MI 48103 © Copyright 1996 by Kevin Scott Beach iii ACKNOWLEDGMENTS I would like to recognize the invaluable field assistance and moral support of1ason Adolf, Heidi Borgeas, Janet Good, Teena Michael, Tim Motley, Nathan Nishimura and Cri Preus. Richard Zimmerman and Randall Alberte. each provided advice and technical training without which this work could not have been completed. I greatly appreciate the support ofmy Dissertation Committee. the Department ofBotany at University ofHawai'i, especially the help ofHarriet Matsumoto and Gerry Ochikubo. the Phycological Society of America and the Research Corporation oftile University ofHawai'i while conducting this research. iv ABSTRACT Stresses ofthe marine iITadiance environment have resulted in a diversity of photoadaptative traits in macroalgae. As judged by in Iim absorption spectra, pigment identities in tropical macroalgae reflected the conservation ofpigments within taxonomic groups. However, pigment concentrations and photosynthetic performance oftropical macroalgae were altered over very precise scales in response to imdiance stress. Specific ioter- and intra-life cycle phase modifications ofphotosyntbetic cbaractcristics appear to contribute to the ecological success ofboth pioneer and perennial macroaIgae. For ~ fm;ciQ and Enteromoazha flexuosa. piooeerHawaiian macroalgae, rates ofphotosynthesis ofreproductive unicells can be equivalent to that ofvegetative adults. Reproductive unicells of!I. fascjata are low-light adapted compared to fa. flexuosa. Concomitant with photosynthetic differences is pigment differentiation found in reproductive unicells oflL. fasciati but not in cells ofa tlexuosa. Spectral analyses indicated either fonnation ofa novel-carotenoid or alteration of existing carotenoid ratios. Maintaining substantial rates ofphotosynthesis by achieving uet-fixation at low irradiances may increase the likelihood that uniceUs will successfully colonize. AhnteltiQpsis concinna and I .anmncia TJJr!Jmpjdjs, perennial Hawaiian macroalgae. exhibit marked pigment differences between canopy and understory tissues of the same thallus separcued by <10 em. Although both species decrease phycobilin concentrations and increase photoprotective pigment levels with increased iITadiance, the specific phycobilins adjusted differ. Photoacclimation in A.. concinna responded precisely to iITadiance quantity but not quality. Concomitant with pigment adjustments was acclimation ofphotosynthetic performance. I ann;ncia mcdermidiae and A. concinua exhibited a sun- to shade-type acclimation over an extremely responsive scale. Understory tissues of A.. concinoa had higher Pmax than canopy tissues. This and other evidence indicates that canopy tissues ofA.. concinna were chronically photoinhibited. v Ahnfeltio.psjs cODciooa is exposed to extreme levels ofemersion stresses. UnJ..ike other intertidal macroalgae, ~ concipna did not exhibit increased temperature tolerance while desiccated. Temperature and osmolarity photosynthetic response curves demonstrated a broad range oftolerance, indicative ofadaptation to high tidal elevatioDS. Although canopy tissues of~ r.ondnoa are chronically pbotoinhibited, they have the physiological capacity to recover photosynthetic activity from emersion stresses more efficiently than do understory tissues from the same thallus. vi TABLE OF CONTENTS ACKNOWLEDGMENTS .iv ABSTRACT v LIST OF TABLES xv LIST OF FIGURES xvii LIST OF ABBREVIATIONS xx PART I. LITERATURE REVIEW AND DISSERTATION PROPOSAL 1 Introduction 2 Ruderal strategists 3 Stress tolerance strategists 4 Photosynthetic pigments in marine macroalgae 6 Chlorophylls 7 Phycobiliproteins 9 Carotenoids 11 Photoacclimation and photoadaptation 14 Environmental factors affecting intertidal algae 16 Irradiance 16 Emersion 18 Organisms of primary interest. 19 Research Objectives 21 vii Hypothesis 23 Literature cited 24 PART n. lli YIYQ. ABSORBANCE SPECTRA AND UV-ABSORBING COMPOUNDS OF TROPICAL REEF MACROALGAE 31 Abstract , 32 Introduction 33 Materials and methods 34 Sample collection and handling 34 In ~ absorption spectral analyses 35 UVACs extraction 35 Results 35 111 rimPAR absorption and fourth derivative spectra in macroalgae 35 Chlorophyta 36 Phaeophyta 36 Rhodophyta 37 Characterization of UV-absorption in tropical macrophytes 38 Correspondence ofillnm absorption and extracts of UVACs 39 Discussion 39 In Yim absorbance spectra in PAR region of irradiance spectrum. 39 In riYsl absorbance spectra in UV region of irradiance spectrum 41 Conclusions ,..42 Literature cited 44 Figures 48 viii ----_._-- PART m. PHOTOSYNTHESIS IN REPRODUcrIVE UNICELLS OF ULVA EASCIAIA DEI,n E AND ENlEROMORPHA FLEXUOSA (WULFEN) I. AGARDH: IMPLICATION FOR ECOLOGICAL SUCCESS 72 Abstract. 73 Introduction 74 Materials and methods 76 lIllim absorption and fourth derivative spectra analyses 76 Release and handling ofmotile cells for physiological measurements 77 Physiological measurements of adult thalli..'" 78 Photosynthesis and respiration measurements 78 Statistical analyses 79 Results 79 Gametophyte and gamete physiological features 79 Sporophyte and zoospore physiological features 79 Sporophyte and recently settled zoospore physiological features 80 In lim absorption features and fourth derivative spectral analysis :.81 Discussion 82 Conclusions 85 Literature cited 87 Tables 90 Figures 93 PART IV. MICROSCALE ACCLIMATION IN PIGMENTATION 99 Abstract. 100 Introduction 101 Materials and methods 104 ix Microsite definitions and comparison scheme l04 PPFD at microsites 105 Sample collection and handling 104 Photosynthetic pigment quantification 106 Analyses ofin lim absorption spectra 106 Extraction of UV-absorbing compounds l07 Statistical analyses 108 Results 108 Microsite characterization l 08 Comparison of photosynthetic pigment at microsites 109 Microsite variation for in Yix2 440 to 750 nm absorbance 109 111 Yim UV-absorption and fourth-derivative spectra 110 Microsite variation as revealed by illnm UV-absorbance 111 Correspondence ofill ~ absorption and extracts ofUV-absorbing compounds 111 Discussion 112 Conclusions 118 Literature cited 120 Tables 127 Figures 128 PART V. MICROSCALE ACCLIMATION IN PHOTOSYNTHESIS I44 Abstract 145 Introduction 146 Methods and materials 148 Sites and sampling ,,.148 x -----~ _..._--_ ...- Photosynthesis and respiration measurements 149 Photosystem unit characteristics 149 Fluorescence emission spectra 150 Relationship ofin vi vo absorption spectra and P vs. I curve parameters 150 Statistical comparisons 151 Results 151 Intertidal distribution 151 Photosynthesis and respiration for canopy and understory tissues 152 Photosynthesis and respiration in Ahnfeltiopsis concinna peripheral and central tissues 153 Photosynthesis and respiration in Abnfeltiopsis concinna HPFD and LPFD tissues 153 Evaluation ofphotosynthetic parameters over vertical distribution 153 Relationship ofilllimabsorption spectra characteristics to photosynthetic performance 154 Discussion 154 Scale ofphotoacclimative and constraints on marine primary producers.154 Diversity in acclimation strategies: a comparison of tropical turfs 156 Conclusions 160 Literature cited 162 Tables 168 Figures 168 PART VI EXPERIMENTAL ANALYSIS OF PHOTOACCLIMATION POTENTIAL 177 xi Abstract.......... 178 Introduction 179 Methods and materials 181 Assessment of photoacclimation rates 181 Statistical analysis 183 Results 184 Day 0 in vivo absorbance characteristics 184