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Below Ground Biology of Botrychium Pumicola

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Below Ground Biology of Botrychium Pumicola An Abstract of the Thesis of Francisco 1. Camacho for the degree ofDoctor ofPhilosophy in Botany and Plant Pathology presented on 22 February 1999. Title: Below Ground Biology ofBotrychium pumicola (Ophioglossaceae). Redacted for Privacy Abstract Approved: __ James M. Trappe Botrychium pumicola Coville is a rare fern, with extant populations in Klamath, Lake and Deschutes counties, ofcentral Oregon. It grows on subalpine pwnice ridges and lower montane lodgepole pine forest openings on pwnice-rich soils. The goal ofthis research was to better understand the below ground biology ofB. pumicola. Detailed examination ofthe subterranean structures ofB. pumicola revealed sporophytic gemmae attached to the stem. Developing gemmae are a non-photosynthesizing stage in the life cycle ofthis plant and are presumed to depend on mycorrhzial fungi for their nutrition. Population genetic analysis ofB. pumicola using inter-simple sequence repeats (ISSR) suggests that the gemmae do not disperse far from the parent plant. Examination ofthe endophytic fungal structures in the roots ofB. pumicola reveal arbuscular mycorrhziae and a high abundance ofseptate hyphae. To better characterize the root fungi, the internal transcribed spacer region offungal ribosomal DNA (ITS) was amplified from root DNA by the polymerase chain reaction (PCR). The ITS amplicon was cloned and sequenced in order to characterize the different fungi in a root segment. Arbuscular mycorrhizal-like Below Ground Biology of Botrych;um pum;cola (Ophioglossaceae) by Francisco J. Camacho A THESIS submitted to Oregon State University in partial fulfilhnent of the requirements for the degree of Doctor ofPhilosophy Completed February 22, 1999 Commencement June 1999 Doctor ofPhilosophy thesis ofFrancisco J. Camacho presented on February 22. 1999 APPROVED: Redacted for Privacy Major Botany and Plant Pathology Redacted for Privacy Chair ofDepartment ofBotany and Plant pathology Redacted for Privacy Dean ofGrad+hool I understand that my thesis will become part ofthe permanent collection ofOregon State University libraries. My signature below authorizes release ofmy thesis to any reader upon request. Redacted for Privacy ~rancisco J. Camacho, Author CONTRIBUTIONS OF AUTHORS Chapter three was coauthored with Aaron Liston, who advised in the design and analysis, provided laboratory resources, and reviewed several drafts. TABLE OF CONTENTS Page CHAPTER 1 Introduction..........................................................................................1 1.1 Abstract......................................................................·························· 1 1.2 Introduction.......................................................................................... 1 1.3 Review ofBotrychium pumicola ...........................................................3 1.4 Other Species ofBotrychium.................................................................6 1.5 Conclusion............................................................................................ 11 CHAPTER 2 Below Ground Structures ofBotrychium pumicola, Roots, Stem, and Gerrunae................................................................................................ 12 2.1 Abstract................................................................................................ 12 2.2 Introduction.......................................................................................... 13 2.3 Methods............................................................................................... 15 2.4 Results................................................................................................. 15 2.5 Discussion............................................................................................22 2.6 Acknowledgments................................................................................23 CHAPTER 3 Population Structure and Genetic Diversity ofBotrychium pumicola (Ophioglossaceae) Based on Inter-Simple Sequence Repeats (ISSR) .......25 3.1 Abstract...............................................................................................25 3.2 Introduction.........................................................................................26 3.3 Methods..............................................................................................28 3.4 Results.................................................................................................30 3.5 Discussion...........................................................................................38 TABLE OF CONTENTS (cont.) 3.6 Acknowledgments................................................................................43 CHAPTER 4 The Mycorrhizae ofBotrychium pumicola (Ophioglossaceae).............. .44 4.1 Abstract................................................................................................44 4.2 Introduction..........................................................................................45 4.3 Methods...............................................................................................48 4.4 Results.................................................................................................50 4.5 Discussion............................................................................................54 4.6 Acknowledgments................................................................................55 CHAPTER 5 DNA Examination ofthe Root Fungal Community ofPumice Grape Fern, Botrychium pumicola (Ophioglossaceae)..............................................57 5.1 Abstract................................................................................................57 5.2 Introduction.........................................................................................58 5.3 Methods...............................................................................................60 5.4 Results.................................................................................................66 5.5 Discussion...........................................................................................77 5.6 Acknowledgments...............................................................................80 CHAPTER 6 Conclusion...........................................................................................82 6.1 Abstract...............................................................................................82 6.2 Introduction.........................................................................................83 6.3 Discussion............................................................................................84 BIBLIOGRAPHY.......................................................................................................87 APPENDICES............................................................................................................98 TABLE OF CONTENTS (cont.) Appendix A ISSR Data Set.. ........................................................................ 99 Appendix B DNA Alignment ofthe Complete ITS nrDNA Region ................ 110 Appendix C DNA Alignment ofthe 5.8S nrDNA. ....................................... 127 LIST OF FIGURES Figure Page 2.1 Class 1 Botrychium pumicola plant, stem with emergent leaL ......................... l 7 2.2 Class 3 Botrychium pumicola plant.. ................................................................ 17 2.3 Class 3 Botrychium pumicola plant.................................................................. 17 2.4 Class 3 Botrychium pumicola plant.................................................................. 17 2.5 Class 3 Botrychium pumicola plant.................................................................. 17 2.6 Class 3 Botrychium pumicola plant.. ................................................................ 18 2.7 Class 5 germinated gemmae ofBotrychium pumicola .......................................18 2.8 The same class 5 germinated gemmae on Botrychium pumicola ....................... 18 2.9 The same class 5 germinated gemmae on Botrychium pumicola ....................... 18 2.10 Class 6, gemmae on Botrychium pumicola ....................................................... 19 2.11 Class 6, gemmae on Botrychium pumicola ....................................................... 19 2.12 Class 3, developing plants attached to a mature sporophyte .............................. 19 2.13 Class 3, developing plants attached to a mature sporophyte .............................. 19 2.14 Several plants ofBotrychium pumicola growing entwined ................................20 2.15 Horizontally developed underground stem ofBotrychium pumicola .................20 2.16 Root scar on the underground stem ofBotrychium pumicola ...........................20 2.17 Leafbud ofBotrychium pumicola.................................................................... 20 2.18 Unusually enlarged area ofa Botrychium pumicola root.. ................................. 20 3.1 Paulina view population ofsampled specimens of Botrychium pumicola .......... .32 3.2 Broken Top population ofsampled specimens ofBotrychium pumicola ............33 LIST OF FIGURES (cont.) Figure ~ 3.3 Katati-2 population of sampled specimens ofBotrychium pumicola ................. .34 4.1 Longitudinal section ofBotrychium pumicola root stained with trypan blue ......51 4.2 Longitudinal section ofBotrychium pumicola root stained with trypan blue ......51 4.3 Arbuscules and aseptate intercellular hyphae in a root ofBotrychium pumicola.51 4.4 Arbuscules
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