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XEROTOLERANCE in FRANKIA by JANA COLLINS WILLIAMS

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XEROTOLERANCE in FRANKIA by JANA COLLINS WILLIAMS XEROTOLERANCE IN FRANKIA By JANA COLLINS WILLIAMS 11 Bachelor of Science Oklahoma State University Stillwater, Oklahoma 1982 Submitted to the Faculty of the Graduate College of the Oklahoma State University in partial fulfillment of the requirements for the Degree of MASTER OF SCIENCE July, 1986 --nle.~I~ 1'1Slo w'1d,.4x. · ~~~ XEROTULRRANCE IN ~ R A N KIA T hesis A pproved ~ _£/~~---~ Thesi: ~isee/ ~. rr , - - - --~ -- --~--~­ __ fi!._'4~~-- ~ _-2Z~zz . D~ Dean o f the Graouate Co l l e ge ---- 125~~82 ACKNOWLEDGEMENTS I wish to express my gratitude to the people who have made the completion of this degree possible. I especially thank my major professor, Dr. Helen S. Vishniac, for her patience, thorough training, and excellent guidance during my masters program. I also thank Dr. Thomas c. Hennessey for serving as a member on my graduate committee, for as- sisting in collecting the wild Alnus maritima, and for allow- ing the use of his facilities. And I thank Dr. Mark R. Sanborn for his advice and service on my committee. A special thanks is due to Ed Lorenzi and Malinda Spatz, , the graduate students and employees of the Department of Forestry involved in this project, for their friendship and assistance in plant inoculation. I thank my husband Patrick for his love and encourage- ment. Finally, I express my love and appreciation for my parents, without whose support, both financial and moral, I would never have succeeded. This work was funded by a grant from the National Science Foundation, grant number NSF-PCM 8213804. Finan- cial support was also provided by the Department of Forestry, and by Department of Botany and Microbiology teaching assistantships. iii TABLE OF CONTENTS Chapter Page I • INTRODUCTION . 1 II. REVIEW OF LITERATURE . 3 Soil Nitrogen and Nitrogen Fixation . 3 Actinorhizal Plants • • . • . • 6 Frankia Taxonomy ...••••. 7 Frankia Isolation and Cultivation . 14 Media • • • • . 21 Other Conditions of Culture 28 Xerotolerance . • • • 29 Host Plant Inoculations 33 Conclusions . • 35 III. MATERIALS AND METHODS 36 Frankia Strains .••••• 36 Chemicals and Media • . • . 39 Strain Maintenance 40 Conditions of Culture 40 Homogenization Procedure 48 Visual Growth Evaluations . 50 Host Alnus Clones . 53 Longevity of Homogenates 53 Medium Development 55 Substrate Utilization Experiments • . 63 Isolations . • . • 65 Measuring Osmolalities of Media . 69 Preliminary Osmototolerance Tests •. 69 Osmotolerance Screens . • . 70 Quantitatives Osmotolerance Measurements . 71 Inoculation of Host Plants 72 IV. RESULTS 74 Longevity of Homogenates 74 Medium Development 74 Substrate Utiliiation Experiments . 82 Isolations 85 Measuring Osmotic Potentials of Media . 88 jv Chapter Page Preliminary Osmototolerance Tests .. 91 Quantitatives Osmotolerance Measurements . 95 Inoculation of Host Plants 106 V. DISCUSSION •.• • 108 Introduction • • • • • . • . 108 Longevity of Homogenates • • 108 Medium Development . 109 Substrate Utilization Experiment • 110 Isolations • . .•. 111 Xerotolerance in Frankia . • • • 111 Inoculation of Host Plants . • • • 115 VI. SUMMARY, CONCLUSIONS, AND SUGGESTIONS FOR FURTHER STUDY ..• • .•.• 116 LITERATURE CITED . 119 v LIST OF TABLES Table Page I. Known Actinorhizal Plant Genera 8 II. Strain Information for Standardized Catalog • . • . • . 15 III. M-3 Medium . 19 IV. Supplemented BD Medium . 20 v. QMOD B . • . 22 VI. Trace Mineral Solution for QMOD B 23 VII. Frankia Broth (YD Medium) 24 VIII. Blom's Medium 25 IX. Substrates Used by Some Frankia Strains . • . • . 27 X. Shipton and Burggraaf's Medium . 34 XI. Strain Acronyms 37 XII. 3AP/2 41 XIII. Frankia Vitamin Mix 42 XIV. Modified Lalonde Trace Mineral Solution . • . 43 xv. Modified QMOD B 44 XVI. Modified Frankia Broth 45 XVII. Phosphate Buffered Saline (PBS) 46 XVIII. Visual Evaluation Grade System for Ten Milliliter Cultures 52 XIX. Alnus Clones 54 XX. Base Experiment Media: Base "A" 57 vi Table Page XXI. Base Experiment Media: Base "B" 58 XXII. Modified Wickerham's Vitamin Mix . 59 XXIII. Vishniac and Santer Trace Mineral Solution 60 XXIV. 3AP 62 XXV. Purshia Strain Base 64 XXVI. Viability of Stored Frankia Homogenates • . • • 75 XXVII. Growth of MPl in Experimental Base Medi a • • • • • • • • . 80 XVIII. Growth of MPl in Lipid/Caco3 Test Media • • • • • • • 81 XXIX. Growth of Five Strains at Varied pH 83 XXX. Ptil Substrate Utilization in Experimental Bases 84 XXXI. MPl Substrate Utilization 86 XXXII. Carbon Source Utilization by Frankia Strains • • . • . 87 XXXIII. Strain Information for OSU 01180101 89 XXXIV. Measured Osmotic Potentials of Media • 90 xxxv. Growth .of MPl in Osmotic Media . 92 XXXVI. Growth of Ptil in Osmotic Media 93 XXXVII. Growth of Subcultured Ptil in Osmotic Media 94 XXXVIII. Visual Evaluations for Osmotolerance Screen of Six Frankia Strains 96 XXXIX. Mycelial Dry Weight Yields of Four Strains in Osmotic Media 97 XL. Infectivity of Frankia Strains in Alnus Clones . • 107 vii LIST OF FIGURES Figure Page 1. The Pommer Diagram . 11 2. Viability of Stored Homogenates of Ptil . 77 3. Viability of Stored Homogenates of MPl . 79 4. Growth Response of Ptil to Decreasing Water Potential . • . 99 5. Growth Response of ARgP5 AG to Decreasing Water Potential . • • .~~ • • • • • • • 101 6. Growth Response of ARbN4Si to Decreasing Water Potential • • • • • • • • • • • 103 7. Growth Response of AVP3f to Decreasing Water Potential • • • • • • • • • • • 105 viii CHAPTER I INTRODUCTION The experiments described herein contribute to the development of a model system for studying resistance to water limitation stress in dinitrogen fixation by actino­ rhizal plants. Actinorhizal plants are certain dicotyle­ donous plants which can form root nodules with bacteria of . the genus Frankia. These symbiotic root nodules are the site of dinitrogen fixation, the biplogical process by which atmospheric molecular nitrogen is combined with hydrogen to form ammonium ions (Brill, 1975; Burns, 1979; Postgate, 1982). An experimental model for quantitative evaluation of the contributions of host, endosymbiont, and nodule physiol­ ogy to xerotolerance (the ability to withstand drying) in ac­ tinorhizal symbiotic dinitrogen-fixing systems (Alnus plus Frankia) is being sou~ht cooperatively by the Department of Forestry, and the Department of Botany and Microbiology. This model could be valuable in the development of plants for use in the many areas where dry, nitrogen-poor soils are prevalent, for plant development research is generally an expensive, long-term project. Under the direction of Thomas C. Hennessey, water 1 2 stress effects on unnodulated cloned host plants have been studied (Hennessey et al., 1985). They compared the effects of water stress on the growth of cloned hosts possessing contrasting xerotolerances which were either nodulated, un­ nodulated, or nitrogen fertilized. The effects of nodulation with Frankia of high or low xerotolerance will later be compared using hosts which have comparatively low or high water stress resistance. The actinorhizal plants produced using water stress re­ sistance model may be able to grow in soils which have lim­ ited nitrogen and moisture availability better than most other plants. They may aid in the development of these mar­ ginal soils and the reclamation of damaged ones into produc­ tive agricultural or silvicultural areas in order that the ever-growing demands for food, fiber, and timber can be sat­ isfied. Such plants have obvious advantages over others. They may be able to grow with minimal or no irrigation, and they may have no need for additions of expensive chemical fertilizers to supply nitrogen. The research presented here is limited to work concern­ ing the microbiological aspects of this shared project: Frankia medium development, isolation of Frankia from Alnus nodules collected from experimentally inoculated or wild hosts, quantitative determination of the xerotolerances of selected Frankia strains, and development of methods for bacterial inoculations of Alnus cuttings. CHAPTER II REVIEW OF LITERATURE Soil Nitrogen and Nitrogen Fixation Nitrogen is needed in large quantities by plants, large­ ly for protein synthesis~ only carbon, oxygen, and hydrogen are more abundant in plants. Because the adequately watered plant can fix carbon from the atmosphere, nitrogen is often the limiting factor in its growth. Nitrogen is usually absorbed from the soil, mainly as salts of nitrate (No 3-) ions by root tissues. The nitrate must be reduced to ammon­ ium before it can be used in biosynthetic processes (Salis­ bury and Ross, 1985). The major forms of nitrogen in soils are organic nitro­ gen associated with humus, ammonium nitrogen adsorbed by cer­ tain clay minerals, and soluble organic ammonium and nitrate. Most soil nitrogen is associated with the organic matter~ this nitrogen is slowly released by microbial mineralization, and therefore not readily available to the plant. Clay-ad­ sorbed ammonia is also slowly released. Soluble ammonia and nitrate in unfertilized soils are rarely more than one to two percent of the total nitrogen present. These soluble com­ pounds are subject to rapid loss by leaching and volatiliza­ tion, and they are readily removed by plants and soil 3 4 microbes (Brady, 1974). The main commercial source for replacement or supple- mentary nitrogen necessary for intensive agricultural methods is ammonia in chemical fertilizer. The ammonia is produced by the Haber-Bosch process by which gaseous dinitrogen and dihydrogen molecules are catalytically reacted at elevated temperature and pressure
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