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MODE of REPRODUCTION and EVOLUTION in the COMPILOSPECIES DICHANTHIUM INTERMEDIUM (Ro Bro) DE WET ET HARLAN

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MODE of REPRODUCTION and EVOLUTION in the COMPILOSPECIES DICHANTHIUM INTERMEDIUM (Ro Bro) DE WET ET HARLAN MODE OF REPRODUCTION AND EVOLUTION IN THE COMPILOSPECIES DICHANTHIUM INTERMEDIUM (Ro Bro) DE WET ET HARLAN By SUNIL y,.ARAN Bachelor of Science Allahabad University · Allahabad, India 1954 Master of Science Patna University Patna, India 1956 Submitted to the faculty of the Graduate College of the Oklahoma State University in partial fulfillment of· the requirements for the degree of DOCTOR OF PHILOSOPHY May, 1968 OKLAHOMA STATE UNIVERSITY LIBRA R Y OCT 4'/ 1968 'Iii·~~. ·.p. , . ... '·: . .,. .......~ . ·• ,.· · ..•. ·,·-:·,·•. .,-,.,, •·. MODE OF REPRODUCTION AND EVOLUTION IN THE COMPILOSPECIES DICHANTHIUM INTERlYIEDIUM (R. Br.) DE WET ET HARLAN Thesis Approved: . Thesis Adyiser Iv . Deanl'Zc,f ~~ College ii PREFACE Each chapter of this dissertation is written, with minor modifications, in the form and style of the biological journal to which it will be presented for publication. It is believed that this method of presentation will allow for more accurate and comprehensive interpretation of the material o I wish to express my sincere gratitude to Dr. J.M. J. d.e Wet, Professor of Botany and Plant Pathology, for his constant guidance and Bncouragement throughout the course of this study. I am also indebted to Dr. de Wet for pro­ viding f i nancial support through a Research Assistantship without which my stay at Oklahoma State could not have been possible. I also wish to acknowledge my gratitude to the late Dr. w. w. Hansen f or the affectionate encouragement I received from him. I owe a debt of gratitude to Dr. Jack R. Harlan, Professor of Genetics, University of Illinois, Urbana, for his many helpful ideas and for critically reading the final manuscript. My appreciati ons are due to the members of my graduate committ ee: Dr. John E. Thoma s, Head of the Botany and Plant Pathology Department; Dr. Glenn w. Todd, Professor of Botany iii and Plant Pathology; Dr. Margaret H. Brooks, Associate Professor of Botl:Lhy and Plant Pathology, and Director, University Hqnors Program; Dr. Eddie Basler, Professor ,of Botany, for their help and encouragement in my graduate program. I also acknowledge my indebtedness to Patna University for granting me study leave and the United State Educational Foundation in India for providing a Fulbright travel granto I extend my appreciation to Mrs. Margaret Estes and Miss Velda Davis for their expert typing of the thesis. My final word of thanks to Mr. Brij Raj Kishore, Mr. Kanti Rawal, Mr. Gilbert Fletcher, Mr. s. P. Sen Gupta and Mrs. Cindy Kelln for their kind assistance and cooperation .. iv TABLE OF CONTENTS Chapter Page I .. INTRODUCTION • • 0 • • • • • • • • • • • • • 1 IIo THE EFFECTS OF ENVIRONMENT ON REPRODUCTION IN FLOWERING PLANTS •••• o ••••• o 8 I:Iistory .. • • • • • • • • • • .. • • • • 9 Temperature and Flowering. • • • • • • 10 Thermoperiodism ., • • • • • • . • • • • • 15 Photoperiodism· and Photoreactions·. • • 17 Endogenous Rhythms • • • • .. • • • • • 24 Growth Substances, Gibberellins and Flowering. • • • • • • • • • • • • • 25 Floral Initiation in Total Darkness. • 29 Floral Differentiation • • • • • • • • 29 Genetic Control of Environmental Responses. • • • • • • • • • • • • • 31 Environment and Reproduction in Grasses. • • • • • • • • • • • • • • 32 Environment and Apom~xis in Plants • • 33 Conclusion • • • • • • • • • • • • • • 37 III. A RAPID SQUASH METHOD FOR GRASS EMBRYOLOGY. 39 Materials and Methods~ • • • • • • • • 40 Results and Discussion Q • • • • • • • 41 IV. REPRODUCTION IN DICHANTHIUM INTERMEDIUM . .. 44 Materials and Methods. • • • • • • • • 45 Results • .. • • • • • .. • • • .. ., • . • • 46 Discussion • • • • • • • • • • • •. • • 56 Va A STRUCTURAL PECULIARITY OBSERVED IN THE SEXUAL EMBRYO SACS OF DICHANTHIUlVl INTERlVIEDIUM .. • • • • • .. • • • • • • • • 61 Materials and Methods. • • • • • • • • 62 Results and Discussion • • • • • • • • 63 v Chapter Page VIo ENVIRONMENTAL CONTROL OF THE MODE OF REPRODUCTION IN DICHANTHIUM INTERMEDIUlVI • 68 Materials and Methods ••• e o ft • . • 69 -Results • • • • • • • • • • • • 0 • • • 71 Discussion • o •••• o • • • • • • • 79 VII. SOME NEW DATA ON INTERRELATIONSHIPS OF DICHANTHIUM INTERlVIEDIUM AND DICHANTHIUM FECUNDUM • • • • • • • • • • • • • • • • • 83 Materials and Methods • • • • • • Q, • • 84 Results • • • • • • • • • • • • • • • • 85 Discussion ••• • o e • • • • • • • • 96 VIII .. EVOLUTIONARY TRENDS IN DICHANTHIUM INTERMEDIUM •••••••••• • • • • • 101 Materials and Methods. • • • • • • • • 102 Results and Discussion • • • • • • • • 103 Phylogenetic Survey of the Genus Dichanthium and the Status of the Compilospecies Dichanthium inter~ medium • • • • • • • • • • • • • • • 154 IX. SUMMARY e • • • • o • • • • • • • • • • • • 164 BIBLIOGRAPHY 0 • • • • • •. • • • • • • • • • • • • 167 vi LIST OF TABLES Table Page Embryological and Reproductive Behavior of Dichanthium intermedium o o • o o o 0 0 47 IIo Data on Apomictic and Sexual Embryo Sac Development in Do intermedium X-750 under Different Photoper1ods o o o o o •• o o • 72 II Io Data on Apomictic Seed Set in Do inter­ medium X-750 under Different~Photo- periods • o o o o o o o o o o • o ... 0 0 '78. IVo Data on the Crossability of Dichanthium intermedium with 12• fecundum • .. • • .. .. .. BT. v. Cytology of Parents and Hybrids 0 0 O O O O 89 VIo Cytology of Dichanthium Species o • o o .. .. 106 VI Io Key to the Different Taxa Recognized in this Study o • • o • • • o ., • • • o • 0 0 108 VIII .. Cytology of Dichanthium intermedium var .. intermedium .. o .. o .. o • • • o o • • 0 0 115 IXo Cytology of Dichanthium intermedi:um varo australis O O O O O O Q O Q f> 0 0 _o O O O 120 x. Cytology of,Dichanthium intermedium var. glabrum • • • .. • • .. • • o .. • • o • 0 0 124 XIo Cytology of Dichanthium intermedium var .. g;r:aham.i:i e • 0 0 0 0 . 0 0 O O O O • 0 0 0 0 136 XII .. Cytology of Dichanthium intermedium var. ind·i cum • • • .. • .. • • • .. • .. • .. o .. 0 142 XIII. Cytology of Dichanthium intermedium var. montan.um o .. o .. ·• • .. .. • • • • • • • 0 0 148 vii LIST OF PLATES Plate Page Stages Showing Development of the Sexual Embryo Sac in Dichanthium intermedium 0 0 50 II. Stages in the Development of ~he Aposporous Embryo Sacs in Dichanthium intermedium .. .. o o .. o .. o .. .. .. o o 0 0 53 III. Four-Nucleate Sexual and Aposporous Embryo Sacs of Dichanthium intermedium Showing Structural Peculiarities ~ • • Q. • • o .. o 65 IV. Sexual and Aposporous Embryo Sac Deyelopment in Dichanthium intermedium X-750_Under Different Photoperiods o " .. o o .. .. .. o .. 74 V. Graph Showing Percentage of Sexual and Aposporous Embryo Sacs Under Day-Lengths of 14 and 12 Hours o o .. .. .. .. .. .. .. .. o .. 76 VI .. Cytology of the Hybrids Between Diehanthium intermedium and ]2.. fecundum .. .. .. .. .. .. o 93 VII. Cytology of the Hybrids Between D. inter- ··· medium and D.. fecundum .... o .-.. ... .. .. • .. 95 VIIIo Distribution of Dichanthium intermedium varo intermedium O O O O • 0 O e Q O O 0 0 114 IX. Distribution of Dichanthium intermedium var .. australis .... o .... o o o o ...... 0 0 119 x. Distribution of Dichanthium intermedium varo glabrum O Q O O • 0 0 0 0 • 0 .. 0 0 0 123 XIo Distribution of Dichanthium intermedium v.~ro grahamii O O " 0 0 0 .. 0 0 0 0 0 0 0 135 XII., Distribution of Dichanthium intermedium V'ar.. indicum .. .. · .. ·· .. · .. ., .. .i .. ., .. .. .. 0 0 141 XIII .. Distri but.ion of Dichanthium intermedium var .. montanum. .. .. .. .. .. .. .. .. .. .. .. .. O O 147 viii Plate Page XIVo Cytology of the Varieties of Dichanthi:um · intermedium o o o o o • • .. .. ., o .. o o e 151 Cytology of the Varieties of Dichanthium intermedium o a a o o o o o o o a a ., 0 0 153 XVIo Classification of the Contributing Species and Their Hybrids o • o o o •• o o •• o 156 ix CH.APTER I INTRODUCTION In order to interpret the evolutionary dynamics of a plant group, an understanding of its breeding system and reproductive. physiology is of utmost importance. The overall reproductive potentiality of any group controls the spread of genetic variability, and will reflect its evolutionary trends. ""Many plant groups display a variable degree of versatility in. their reproductive behavior which may be manifested by a side-by-side persistence of al te.;rn.ative pathways of seed production either via agamospermous apomixis, or through normal sexual reproduction. Such facultative apomicts provide excellent materials for the study of evolution as directed by mode of reproduction. Facultatively apomictic plants further present a unique opportunity for a study of mechanisms that control sexual and asexual systems of seed formation. In.the ovule of some facultatively apomictic angiosperms two developmental pathways, the for­ mation of a normal. sexual embryo sac with a cytologically reduced egg and the formation of a cytologically unreduced aposporous embryo sac, function at the same time. Finally, one prevails over the other in forming the seed. In the sexual embryo sac, embryo development starts after 1 2 fertilization of the egg by a male gamete, whereas in the aposporous embryo sac meiosis and usually also fertilization are circumvented, and the cytologically unreduced egg develops parthenogenetically to form the embryo and finally the seed is produced. Occasionally, a cytologioal.ly reduced egg may develop parthenogenetically to form a haploid embryo, and the cytologically unreduced egg in an aposporous sac may be fertilized. In facultative apomicts, factors determining the ulti­ mate success of one system of reproduction over the other remain largely elusive.
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