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Experimental Induction of Tropic Responses in Trifolium Fragiferum L

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Experimental Induction of Tropic Responses in Trifolium Fragiferum L This dissertation has been microfilmed exactly as received 69-22,139 HANSEN, Dale J., 1939- EXPERIMENTAL INDUCTION OF TROPIC RESPONSES IN TRIFOLIUM FRAGIFERUM L. STOLONS. The Ohio State University, Ph.D., 1969 Agronomy University Microfilms, Inc., Ann Arbor, Michigan EXPERIMENTAL INDUCTION OP TROPIC RESPONSES IN TRIFOLIUM FRAOIFERUM L. STOLONS DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University By Dale J. Hansen, B.S., M.S. ****** The Ohio State University 1969 Approved by Adviser Department of Agronomy ACKNOWLEDGMENTS I wish to thank Dr. Leo E. Bendixen for the help he has given me as my adviser; especially for the guidance given concerning my research and dissertation. This work was made possible by the teaching and research associate positions granted to me by the faculties of the Botany and Agronomy Departments. All assiBtanoe was sincerely appre­ ciated. I am indebted to my wife, Ruth, for the many hours spent in supporting my graduate studieB and in typing this dissertation. Dr. Richard Fopham kindly provided assistance and materials for the anatomical studies presented in this dissertation. ii VITA Sept. 3, 1939 • * • Born — Idaho Falls, Idaho June, 1963........ B.S., University of Idaho Mosoow, Idaho 1963-1966 ........ Teaching Assistant, Department of Biological Sciences, University of Idaho, Moscow, Idaho 1966* ••••••• M.S., University of Idaho Moscow, Idaho 1966-1967 ..... Teaching Assistant, Department of Botany, The Ohio State University, Columbus, Ohio 1967-1969 ........ Research Associate, Department of Agronomy, The Ohio State University, ColumbuB, Ohio FIELD OF STUDY Plant Physiology iii TABLE OF* CONTENTS Page ACKNOWLEDGMENTS ii VITA iii LIST OP TABLES VI LIST OP FIGURES vii INTRODUCTION 1 Chapter I. LITERATURE REVIEW 4 Tropisma— A Definition The Test Plant Effect of Light on TropismB Fhytoohrome Gravity Geoelectrio Effect Oxygen and Carbon Dioxide Requirements Exogenously Applied Growth-Regulating Chemicals Gibberellins Auxins Ethylene Cobalt Morphaotins II. MATERIALS AND METHODS 28 General Growing Conditions Application of Exogenously Applied Materials Liquids Gases Light Studies Red— far-red Photoperiod Light intensity Assays of Endogenous Materials Ethylene Oibberellin-like substances Anatoraioal Studies iv TABLE CP CONTENTS— Continued Page III. RESULTS 37 Erogenously Applied Growth-Regulating Compounds General observations Gibberellic acid plus cobalt Oibberellic acid plus Morphaotin IT 3233 Ethrel and ethylene-producing substances DC MU Altering atmospherio components Light Studies Red— far-red Photoperiod Light intensity Assays of Endogenous Materials Ethylene Gibberellin-like substances Anatomy of Strawberry Clover Stems IV. DISCUSSION 63 V. SUMMARY 78 APPENDIX 81 LITERATURE CITED 82 v LIST OF TABLES TABLE PAGE 1. Some Aspects of Plant Growth and Development Which Are Affected by Red and Far-Red L i g h t ..................... 12 2. Concentrations of CheraicalB Used on Prostrate Stolons. 30 3. Shading Effect Produced by Various Layers of Cheese C l o t h .................... 33 4. Release of Ethylene from Bending Stolons .......... 60 vi LIST OF FIGURES FIGURE PAGE 1. The typioal growth habit of Trifolium fragiferuin L. f variety Salina. ........................................ 2 2. A skeletal presentation of porphyrin, some common bile pigments, and phytochrome.............................. 13 3. An early bending stage of a prostrate clover stem induced by GA3................................................... 38 4. A later stage of curvature .............................. 39 5. A typical 90° bend promoted by GA^ ............. 40 6. A tropic effect other than stem-tip curvature.............. 41 7. The effect of GA^ (1 g/l) on stolons...................... 43 8. Bending of clover BtolonB as a result of movement. ..... 44 9. Stem-tip elevations of stolonB treated with 1 g/l QA^ and 1 g/l GA^ plus cobalt chloride...................... 45 10. A comparison of tropic curvatures following treatment with -2 50 ppm ethylene or 3 x 10 M Ethrel treatments. ..... 47 11. Tropic curvature following IAA (1 x 10 ^M) application . 49 12. Tropic curvatures of stolons in the absence of Og, CO2, or treatments with 10 DCMU.............................. 50 13. The path of least resistance .................... 52 LIST OP FIGURES— Continued FIGURE PAGE 14» The Lending of stolons treated with red or far-red light. 53 15. Effect of photoperiod on erect type of strawberry clover. 54 16. Effect of photoperiod on the prostrate type of strawberry clover................. • ......................... • 56 17. A comparison of erect and proBtrate-type strawberry clover plants under 24-hour photoperiod ...................... 57 18. Upward curvature of stolons placed in darkness. ...... 58 19* Measurements of gibberellin-like substances from dark and light-grown stolons with the barley endosperm bioassay . 61 viii INTRODUCTION Gravity, light, and various chemical substances cause plant organs to bend. Isolation of the receptor mechanism in tropic responses and the identification of biochemical Btep3 leading to differential growth provide many challenging problems. Prostrate and erect stems of Trifolium fragiferum L. (straw­ berry clover), variety Salina (Pig. 1) were used in the present study on tropisms. Prostrate strawberry clover stems become erect when kept in dim light or darknesB, when kept in a nitrogen atmosphere, when covered with water, or when treated with gibberellic acid (OA^) (Bendixen, 1960). The present study was conducted to understand why the above phenomena occur and to explain why Borne strawberry clover stems are prostrate and others are erect. Three specific areas of research looked promising and were investigated* chemical-, atmos­ pheric-, and light-regulation of tropisms. Strawberry clover stolons become erect when treated with GA-^ but curve upward only slightly following an indoleacetic acid (IAA) application. It was desirable to know if tropic responses could be induced or inhibited in these prostrate stems with other growth regu­ lating ohemioals Buch as ethylene, kinetin, and Morphaotin. Ethylene evolution from bending Btolons was measured by gas chromatography. Gibberellin-like compounds were extracted from prostrate and bending stems and compared using the barley endosperm bioassay. Clover stolons 2 “ .. - Ti^ i .*■• .. t. jv' *■ ^V. * .rj £aua&m*a% ^ ^ i r . r - ^ W ft/ ■'■•■ ■ ■ s.tff*r,-.v’V:.,jsiiu*>*"fc- +* '••jnipw/TLdMl . iavaiUNWBL. ««l«V r maAtamwummmum^iA ■ m^avo»Min!nwtnu3t jffiraMBnmnsmw .. ’te^3XSjfte£A£lxa&'i£.^^ 1 Fig. 1.— The typical growth habit of Trifoliuin fragiferum L., variety Salina. The prostrate plant on the left demonstrates the growth form commonly associated with T. fragiferum. The erect plant on the right is a single—gene recessive mutant of the prostrate form. covered with water or kept in a nitrogen atmosphere curved upward. It was suspected that a limited oxygen or carbon dioxide supply was the indirect cause of the curvature. Experiments were conducted in which the atmospheric composition was varied to investigate this hypothesis. Light quantity, quality, and duration were Btudied to discover why strawberry clover stolons become erect when kept in darkness but are prostrate in bright light. A microscopic comparison of tissues and cell types of erect and prostrate strawberry clover stems was carried out. The first ten elongated internodes were examined in transection in an effort to determine why one type growB erect and the other grows prostrate. Sufficient information was obtained from the studies on chemical-, atmospheric-, and light-regulation of tropisms to postulate a mechanism of bending. I. LITERATURE REVIEW Tropisms— A Definition Plant movements have been of interest to scientists Bince the early 19th Century (Knight, 1806). The movements of higher plant organs (in-..which the direction of movement is not determined by the structure) have been categorized using the suffix "tropic" with a prefix denoting the source of the stimulus. ThUB, the adjectives phototropic, geotropic, hydrotropic, thigmotropic, chemotropic, magno- tropio, etc. characterize the movements of plant organs. Further complicating the study of bending effects, workers have categorized types of a given stimulus. Words such as diageotropic, orthogeotropic, piagiogeotropic, ageotropio, positively geotropic, and negatively geotropic have been used. A tropic response is the curvature developed by a plant organ resulting from a varied environmental factor (Wilkins, 1966). Tropic movements, unlike nastic movements, are not predetermined by the structural characteristics of the plant part involved. Two prefixes are often coupled with the word tropic. The first prefix describes the direction of growth of the plant part while the second prefix describes the cause. Thus, the term diageotropic, describes an organ growing perpendicular to the force of gravity, while orthogeotropic describes an organ growing parallel to the force of gravity. Plagiogeotropic denotes growth in some plane not parallel or perpendicular to the 5 force of gravity. Ageotropic is applied to plant organB not affected by gravity, while positively geotropic and negatively geotropic desig­ nate whether the curvature is toward or away from the force of gravity. The Test Plant Recent work on the bending mechaniBm of plant organa has centered around a few plant species. Tropic studies of stoloniferous plantB have been primarily limited
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