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Jthrrm. Trappe AN ABSTRACT OF THE DISSERTATION OF An Jumpponen for the degree of Doctor of Philosophy in Forest Science presented on March 16, 1998. Title: Primary Succession on a Receding Glacier Forefront - with Special Emphasis on Root-Colonizing Fungi. Signature redacted for privacy. Abstract approved: JthrrM. Trappe Plant succession is among the fundamental concepts in ecology. Studies addressing plant recruitment, successional mechanisms, and the role of root-colonizing fungi with focus on dark-septate endophytes (DSE) were conducted in the laboratory and on the forefront of receding Lyman Glacier (North Cascade Range, Washington, U.S.A.). Primary successional studies primarily focus on biotic interactions although initial recruitment is controlled abiotically. Ina field study, we found that plant distribution is not random but determined by sites that allow seed trapping and protect germinating seeds from desiccation. An additional hypothesis that early successional plant individuals form centers of establishment for subsequent colonizerswas tested by assaying the effect of willow shrubs on invading plants. Results suggested that while the canopy had either neutral or inhibitory effectson new seedling establishment, the soil beneath the canopies favors establishment. The role of fungi in succession is poorly known. We present results of eight years of floristic surveys of the ectomycorrhizal macrofungi at the site. The data show that the communities of ectomycorrhizal fungi differ between primary and secondary successional sites. Plants on the glacier forefront were frequently colonized by nonmycorrhizal DSE. We reviewed the literature on DSE and listed reports of about 600 colonized plant species from the tropics to arctic and alpine habitats. Although they appear to be ubiquitous, the ecological role of DSE is poorly understood: results of past experiments are inconsistent and do not explain the role of DSE in their natural habitats. We further investigated the host-DSE association and its relevance to plant fitnesson the glacier forefront in laboratory and field studies. We conclude that some DSEmay function as typical mycorrhizal associates but that the research tools for studying these associations are inadequate and biased. A field study on the distribution of discrete RAPD-phenotypes (individuals) showed that DSE colonize several plant species under natural conditions. A single fungal individualcan colonize adjacent plants, potentially providinga mycelial pathway for resource flow as hypothesized for ectomycorrhizal systems. ©Copyright by An Jumpponen March 16, 1998 All Rights Reserved Primary Succession ona Receding Glacier Forefront - with Special Emphasis on Root-Colonizing Fungi An Jumpponen A DISSERTATION submitted to Oregon State University in partial fulfillment of the requirements for the degree of Doctor of Philosophy Presented March 16, 1998 Commencement June 1998 ACKNOWLEDGMENT Completion of this program is by no means by a single person's effort. Volunteers joined in the field expeditions, sharing the weight and loadon the trail and hardships atcamp, including snowbunnies and girl scouts. Personnel at the Forestry Sciences Laboratory provided thesupport needed to get 'things' done. Coauthors substantially contributed to the writing effort and reviewers, anonymous and those known byus, improved the final product, or at least confused the senior author. I hope I have all these contributions acknowledged in their respective places within this dissertation. I reserve the following few words for thanking all those who merit spcial notice. This program was supported by Papa Jumpponen'sgenerous donations, U.S. National Science Foundation Grant DEB-931OOO6, Emil Aaltonen's Foundation, The Mazarnas in Portland, and the U.S. Forest Service PNW Research Station. Personnel at the U.S. Forest Service, Wenatchee National Forest, Chelan Ranger District, WA (especially Ken Dull and Al Murphy) provided the logistical support. Horsepackers Cragg Courtney and Sid Bums transported supplies and samples and savedmy back. I owe to Lisa Ganio, Manuela Huso, and Tom Sabin for thrilling discussions about experimental design and statistical analyses. Sandy Lewis, Donaray McKay, and Glenda Serpa generously and liberally accepted responsibilitiesas three mothers. Caryn Davis not only suffered through almost all the chapters but she kindly provided thepoem that you find as the preface of this dissertation. Kim Mattson weeded all the creativity out ofmy boring science writing, teaching me to recognize all that is necessary for science chat. My committee was most helpful. Steve Strauss made his lab and expertise available for the molecular work. Jeff Stone (and Randy Currah) introduced me to the untamed world of miscellaneous and mysterious sterile endophytes. Randy Molina provided his unconditional support and, furthermore, read and editedevery single word within these covers. Jim Trappe invited me to join his lab and allowedme to try out and experiment with several different approaches. He, like Randy, read and editedevery word within these covers. Hemay actually have written it altogether. Friends too numerous to mention here supportedme through the years of struggle. Myself and the cat, we love you! Wes Colgan and Ronald Rhatigan introduced me to all that is worthwhile: Ren and Stirnpy, kegorators, Joe Bob Briggs, and PPB. Ashley Schultz valiently workedat keeping me straight when I tended to stray, unfortunately, withpoor success. Kei Fujimura, Lisa Grubisha, Teresa Lebel, and Jamie Plattwere always available when bitchjn' about the world's unfairness seemed to be in need. Marjoriitta Möttönen contributed to the cyber-comiseratingover graduate school. Math Maltamo infused the fresh winds of Finnish alternativesto music. Papu ja pojat, Morphine, Jyrki Walli, A.W. Yrjana, Ismo Alanko, Trent Reznor, and Don Huonot joined Ankie Camacho, Rod Wayne and myself when deviation from the usual was desired. Last but not least I want to thankmy parents and the two girls in my life: Nancy-girl and Hackysack the Cat. I don't think this would have been possible without you. CONTRIBUTION OF AUTHORS Several co-authors contributed to the chapters. Dr. Efren Cázares collected most of the data for chapter 4 and assisted in field work. Drs Rauni Ohtonen and Henry Väre assisted in experimental design, field work, data entry, data analyses, and faithfully edited the manuscripts for chapters 2 and 3. Dr. Kim Mattson and Dr. James Trappe participated all stages of theprocess, including the initiation, brain-stroming, money-gathering, design, analysis, and editing. TABLE OF CONTENTS Page CHAPTER 1 1 INTRODUCTION 1 CHAPTER 2 8 CHARACTERIZATION OF "SAFE SITES" FOR PLANT RECRUITMENT ON A PRIMARY SUCCESSIONAL GLACIER FOREFRONT 8 ABSTRACT 9 INTRODUCTION 9 MATERIALS AND METHODS 11 Study Site 11 Sampling and Data Collection 12 Statistical Analyses 14 RESULTS 15 DISCUSSION 21 ACKNOWLEDGMENTS 25 CHAPTER 3 26 EFFECTS OF ESTABLISHED WILLOWS ON PRIMARY SUCCESSION ON LYMAN GLACIER FOREFRONT, NORTH CASCADE RANGE, WASHINGTON, U.S.A.: EVIDENCE FOR SIMULTANEOUS CANOPY INHIBITION AND SOIL FACILITATION 26 ABSTRACT 27 INTRODUCTION 27 TABLE OF CONTENTS (Continued) Page MATERIALS AND METHODS 31 Study Area 31 Experiment 1: Effect of Willow Shrubs on Frequencies of Indigenous Species 32 Sampling 32 Statistical Analysis 33 Experiment 2: Effects of Willow Canopies and Willow Soilon Emergence of Pin us con torta Germinants 34 Establishment of Seeding Treatments 34 Statistical Analysis 36 Characterization of Soil Nitrogen and Organic Matter Contents36 Sampling and Chemical Analysis 36 Statistical Analysis 37 RESULTS 38 Experiment 1: Effect of Wifiow Shrubs on Frequencies of Indigenous Species 38 Experiment 2: Effect of Wifiow Shrubs on Emergence of Pinus con torta Germinants. 38 Characterization of Soil Nitrogen and Organic Matter Contents42 DISCUSSION 45 ACKNOWLEDGMENTS 49 CHAPTER 4 51 ECTOMYCORRHIZAL FUNGI IN LYMAN LAKE BASIN: A COMPARISON BETWEEN PRIMARY AND SECONDARY SUCCESSIONAL SITES. 51 ABSTRACT 52 INTRODUCTION 53 MATERIALS AND METHODS 54 RESULTS 60 TABLE OF CONTENTS (Continued) Page DISCUSSION 64 Ectomycorrbizal Species Shared Between Sites 64 Ectomycorrhizal Species Unique to the Lyman Glacier Forefront Site 67 EctomycorrKizal Species Unique to the Glacier View Ridge Site 70 Ectomycorrhizal Species Unique to Lyman Lake Site 73 CONCLUSIONS 78 ACKNOWLEDGMENTS 82 CHAPTER 5 83 DARK-SEPTATE ENDOPHYTES: A REVIEW WITH SPECIAL REFERENCE TO FACTJLTATIVE BIOTROPHIC SYMBIOSIS 83 ABSTRACT 84 INTRODUCTION 84 EARLY OBSERVATIONS ON DSE 86 HOSTS AND GEOGRAPHICAL RANGES 88 MORPHOLOGY OF ROOTS COLONIZED BY DSE 95 TAXONOMIC AFFINITIES OF DSE 98 EFFECTS OF DSE ON THEIR HOSTS 105 ECOLOGY 108 DISCUSSION AND CONCLUSIONS 112 ACKNOWLEDGMENTS 115 TABLE OF CONTENTS (Continued) Page CHAPTER 6 116 MYCORRHIZAL FUNCTIONING OF PHIALOCEPHALA FORTINII WITH PINUS CONTORTA ON GLACIER FOREFRONT SOIL: INTERACTIONS WITH SOIL NITROGEN AND ORGANIC MATTER 116 ABSTRACT 117 INTRODUCTION 117 MATERIALS AND METHODS 119 RESULTS 121 Root Colonization by Phialocephala fortinii 121 Growth Response 122 Nutrient Accumulation 122 DISCUSSION 126 ACKNOWLEDGMENTS 128 CHAPTER 7 130 PERFORMANCE OF PINUS CONTORTA INOCULATED WITH TWO STRAINS OF ROOT ENDOPHYTIC FUNGUS, PHIALOCEPHALA FORTINII: EFFECTS OF SYNTHESIS SYSTEM AND GLUCOSE CONCENTRATION 130 ABSTRACT 131 INTRODUCTION 132 MATERIALS AND METHODS 133 Aseptic Synthesis 133 Open Pot Synthesis 137 TABLE OF CONTENTS (Continued) Page RESULTS 138 Aseptic Synthesis 138 Biomass 141 Nutrient Analyses
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