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U MI MICROFILMED 199S INFORMATION TO USERS This manuscript has been reproduced frommicrofilm the master. UME films the text directly from the original or copy submitted. Thus, some thesis and dissertation copies are in typewriter face, while others may be from any type of computer printer. The quality of this reproduction Is dependent upon the quality of the copy submitted. Broken or indistinct print, colored or poor quality illustrations and photographs, print bleedthrough, substandard margin*, 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 of the 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. A Bell & Howell information Company 300 North Z eeb Road. Ann Arbor. Ml 48106-1346 USA 313.'761*4700 800/521-0600 NUTRIENT DYNAMICS AND RESORPTION IN FOUR UNDERSTORY WOODLAND PLANTS AND NOTES ON THE MYCORRHIZAL STATUS OF SOME TYPICALLY NONMYCORRHIZAL PLANTS DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy in the Graduate School of The Ohio State University By t Brent Gordon DeMars, B.A., M.S. ***** The Ohio State University 1995 Dissertation Committee: Approved by R. E. J. Boerner M. G. Cline P. S. Curtis \ \ J Advisor Department or Plant Biology E. K. Sutherland UMI Number: 9533957 UNI Microform 9533957 Copyright L995* by UMI Company. All 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 To Brittany ii ACKNOWLEDGEMENTS I would like to thank Jin Runkle at Wright state University for all his assistance in the past and present and Ralph Boerner for his guidance and support throughout this work. Thanks go to Jennifer Brinknan for perfoming all the Nitrogen analyses and the OSU Soil Ecology Lab for equipment sharing. I also thank Dr. Folly Penhale of the Division of Polar Biology and Medicine/NSF for field work in Antarctica. I also thank my family for their support and love throughout my time at Ohio State. iii VITA August 30, 1964 .................... Born Lakewood, Ohio 1986 ................................ B.A., Wright State University, Dayton, Ohio 1988 ................................ M.S., Wright State University, Dayton, Ohio 1991-1995 G.T.A. Department of Plant Biology, Ohio State University, Columbus, Ohio PUBLICATIONS DeMars, B.G. and J.R. Runkle. 1992. Groundlayer vegetation ordination and site-factor analysis of the Wright State University woods (Greene county, Ohio). Ohio Journal of Science 92: 98-106. DeMars, Brent G. 1994. Star-of-Bethlehem, Ornithogalum umbellatum L. (Liliaceae): an invasive, naturalized plant in woodlands of Ohio. Natural Areas Journal 14: 308-309. DeMars, B. and R. E. J. Boerner. 1994. Vesicular- arbuscular mycorrhizal fungi colonization in Capsella bursa-pastoris (Brassicaceae). American Midland Naturalist 132: 377-380. FIELD OF STUDY Major Field: Plant Biology iv TABLE OF CONTENTS DEDICATION ............................................ ii ACKNOWLEDGMENTS ...................................... iii VITA .................................................. iv LIST OF TABLES ...................................... viii LIST OF FIGURES............................. X CHAPTER PAGE I. INTRODUCTION..... ............................ 1 II. INTRASTAND VARIATIONS IN FOLIAR NUTRIENT DYNAMICS AND RESORPTION IN NATURALIZED LONICERA MAACKII (CAPRIFOLIACEAE) POPULATIONS IN OHIO, USA Introduction..... ............................ 6 Materials and Methods ........................ 7 Results.................................... 13 Discussion................................. 34 III. NUTRIENT RESORPTION IN LONICERA MAACKII (CAPRIFOLIACEAE) ALONG A SOUTH-WESTERN OHIO FOREST CHRONOSEQUENCE Introduction..... ........................... 41 Materials and Methods ........ 42 Results.................................... 47 Discussion................................. 55 v IV. FOLIAR PHOSPHORUS AND NITROGEN RESORPTION IN THREE WOODLAND HERBS OF CONTRASTING PHENOLOGY Introduction...... ......................... 58 Materials and Methods ..................... 60 Results ............... 67 Discussion ................................ 77 V. ARBUSCULAR MYCORRHIZAL DYNAMICS OF THREE WOODLAND HERBS OF CONTRASTING PHENOLOGY Introduction...... ......................... 83 Materials and Methods ..................... 84 Results ................................... 90 Discussion ............................... 100 VI. ARBUSCULAR MYCORRHIZAL FUNGI COLONIZATION IN CAPSELLA BURSA-PASTORIS (BRASSICACEAE) Introduction...... ........................ 106 Materials and Methods .................... 107 Results .................................. 109 Discussion ............................... 112 VII. ARBUSCULAR MYCORRHIZAL DEVELOPMENT IN THE BRASSICACEAE IN RELATION TO PLANT LIFE SPAN Introduction ........................ 115 Materials and Methods .................... 120 Results .................................. 122 Discussion ............................... 142 VIII. ARBUSCULAR MYCORRHIZAL DEVELOPMENT IN AN ANNUAL, BIENNIAL, AND PERENNIAL CRUCIFER Introduction ........................ 145 Materials and Methods ................... 146 Results .................................. 149 Discussion ............................... 157 vi IX. MYCORRHIZAL STATUS OP DESCHAMP5IA ANTARCTICA DESV. IN THE PALMER STATION AREA, ANTARCTICA Introduction.. ............................ 162 Materials and Methods ...................... 163 Results ................................... 165 Discussion................................ 165 . X. SUMMARY OF MAJOR RESULTS AND CONCLUSIONS .. 168 LIST OF REFERENCES ............................... 176 vii LIST OF TABLES TABLE PAGE 1. Soil characteristics of study sites by topographic position ........................ 11 2. Mean summer foliar nutrient concentrations in Lonicera maackii by year by site by topographic position .................................... 24 3. Analysis of variance of pooled summer (July September) foliar nutrient concentrations ... 25 4. Comparison of foliar nutrient enrichment (demand:availability quotients) ............ 27 5. Comparison of foliar nutrient resorption in Lonicera maackii during 1992-1994 .......... 31 6. Univariate repeated measures analysis of variance for absolute and proportional P and N resorption for YEAR-repeated measures ...... 32 7. P and N foliar proportional resorption reported for temperate deciduous shrubs of forests ... 40 8. Characteristics of the four study stands in the Wright State University woods (Greene County, Ohio) .................. 44 9. Comparison of summer (July-September) mean foliar nutrient concentrations in Lonicera maackii by successional age per y e a r ........ 51 10. Comparison of foliar nutrient resorption in Lonicera maackii for 1992-1994 within the Wright State University wood's chronosequence .... 52 11. Univariate repeated measures analysis of variance of absolute and proportional foliar P and N resorption for YEAR-repeated measures 53 viii 12. Soil characteristics of study sites by topographic position ....................... 65 13. Mean foliar P concentrations determined from presenescent sampling periods when leaves were at maximum leaf area ....................... 71 14. Overall analysis of variance for absolute and proportional foliar P resorption............ 73 15. Mean foliar N concentrations determined from presenescent sampling periods when leaves were at maximum leaf area ....................... 74 16. Overall analysis of variance for absolute and proportional foliar N resorption............ 76 17. Soil characteristic of study sites by topographic position ....................... 89 18. Overall ANOVA for percent of root length colonized by AM fungi in Trillium flexipes and Smilacina racemosa ......................... 98 19. Percent AM fungi colonization in Caosella bursa- pastoris .................................. 110 20. ANOVA of percent root length colonized by AM fungi in Caosella bursa-oastoris ........... ill 21. Crucifer taxa examined for AM development as reported in the literature................. 117 22. The occurrence of AM development in greenhouse- inoculated crucifers ...................... 125 23. Brassicaceae taxa not exhibiting AM fungi colonization following experimental inoculation ............................... 132 24. Number of plants colonized by mycorrhizal fungi by harvest date ........................... 154 25. ANOVA of mean root length colonized by mycorrhizal fungi in 3 crucifer species .... 155 26. Number of plants colonized by mycorrhizal fungi by harvest d a t e ........................... 156 27. Summary of proportional P and N resorption . 170 ix LIST OF FIGURES FIGURE PAGE 1. Seasonal patterns of Lonicera maackii foliar P within the Wright State University woods during.1992-1994 .........................