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Cytotaxonomic Studies in the Artemisia Ludoviciana Polyploid Complex of the Pacific Northwest

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Cytotaxonomic Studies in the Artemisia Ludoviciana Polyploid Complex of the Pacific Northwest AN ABSTRACT OF THE THESIS OF James Russell Estes for the Ph. D. (Name) (Degree) in Botany (Systematic Botany) presented on I' i..,. ( . /0 (î; / (Major) (Date) Title: CYTOTAXONOMIC STUDIES IN THE ARTEMISIA LUDOVICIANA POLYPLOID COMPLEX OF THE PACIFIC NORTHWEST Abstract approved: Kenton Lee Chambers The Artemisia ludoviciana polyploid complex is a highly poly- morphic assembledge of eight taxa in the Pacific Northwest, with a chromosome base number of x =9. A cytogenetic analysis of this complex was carried out to determine the sources of its variation, the types of polyploidy present, and the genetic processes active in its evolution. Chromosome counts in the complex disclosed two diploid taxa, A. michauxiana and A. cavaticaulis, in addition to the previously counted diploids, A. suksdorfii and A. lindleyana. Results of meiotic analysis and artificial hybridization experi- ments strongly suggested the autoploid origin of the polyploid members of the complex. These results included the constancy of multivalent formation among the polyploids, the maximum number of multivalents formed in each taxon, the pairing relationships of the hexaploid and trisomic hexaploid A. douglasiana, the degree of homology between the included genomes of the artificial hybrids, and the presence of heteroploidy in at least five of the included taxa. The attributes of the complex that might have contributed to the origin and success of autoploidy were also examined. Observations indicate that the diploids produce a low but constant number of un- reduced pollen grains. Introgression from the diploid to the tetra - ploid level, through tetraploid progeny, provides a source of variation for the newly formed autoploids. The complex has a broad physio- logical base. Mechanisms are present in the meiotic apparatus that could allow for an increase in fertility of autoploid individuals. The phylogenetic and taxonomic relationships among several of the taxa are discussed. CYTOTAXONOMIC STUDIES IN THE ARTEMISIA LUDOVICIANA POLYPLOID COMPLEX OF THE PACIFIC NORTHWEST by James Russell Estes A THESIS submitted to Oregon State University in partial fulfillment of the requirements for the degree of Doctor of Philosophy June 1968 APPROVED: Professor of Botany in charge of major Hea of Dep tment of Bony and Plant Pathology Dean of Graduate School Date thesis is presented .) al( L i 1 Typed by Kay Smith for James Russell Estes ACKNOWLEDGMENT The author wishes to express his appreciation to Dr. Kenton L. Chambers for his guidance and technical assistance during the course of this study. Mrs. LaRea Dennis Johnston and Mr. Ronald J. Tyrl also provided many helpful suggestions and criticisms which the author gratefully acknowledges. The author also wishes to thank the numerous other individuals who made valuable contributions toward the completion of this thesis in collecting and sending living specimens, often from great distances; these individuals are cited in Appendix I. A special note of thanks is due my wife, Nancy, not only for her assistance on collecting trips and the compilation of the thesis but also for her cheerful encouragement. Funds for this study were provided in part by the National Science Foundation in the form of a NSF Cooperative Graduate Fellowship and a NSF Summer Teaching Fellowship. TABLE OF CONTENTS Page I. INTRODUCTION 1 II. LITERATURE REVIEW 3 Classification of Polyploids 3 Hybridization and Polyploidy 5 Stages in the Evolution of Polyploid Complexes 6 Characteristics of Autopolyploids 7 Evolutionary Trends in Autopolyploids 12 The Species Concept in Relation to the Autopolyploid Complex 15 Origin of Autopolyploids 18 Previous Taxonomic and Cytological Studies in Artemisia 19 The Subgenus Seriphidium 20 The Subgenus Dracunculus 22 The Subgenus Artemisia 23 III. MATERIALS AND METHODS 34 Collections 34 Cytological Studies 36 Meiotic Analysis 36 Pollen Fertility 37 Pollen Diameter 38 Hybridization 38 IV. RESULTS 41 Cytology 41 Chromosome Counts 41 Pollen Diameter 43 Meiotic Analysis of the Diploids 43 Meiotic Analysis of the Tetraploids 55 Meiotic. Analysis of the Hexaploids 67 Hybrids 67 Cytology of the Hybrids 80 V. DISCUSSION 87 Page Autopolyploidy within the Complex 87 Indicators of Autopolyploidy 87 The Formation of Unreduced Pollen 94 The Introduction of Variation 96 Ecotypic Variation 97 Mechanisms for Fertility Improvement 99 Taxonomic Relationships within the Complex 103 Intertaxon Relationships 105 The origin of Artemisia douglasiana 105 The taxonomic position of Artemisia prescottiana 107 The relationship between Artemisia ludoviciana and Artemisia. lindleyana 108 Artemisia michauxiana, a diploid - tetraploid pair, and Artemisia ludoviciana ssp. incompta, 111 Artemisia suksdorfii and Artemisia cavaticaulis 112 Taxonomy of the Artemisia lu:doviciana- Complex 117 BIBLIOGRAPHY 118 APPENDICES 125 Appendix I. Site Data and Chromosome Numbers for Artemisia Populations 125 Appendix II. Leaf Outlines of Selected F1 Hybrids and Their Parents 140 Appendix III. Relevant Data for Specimens Used in Pollen Studies 156 LIST OF FIGURES Figure Page 1 Distribution of Artemisia vulgaris, sensu lato. 25 2 Comparative pollen grain sizes. 44 3 Distribution of Artemisia michauxiana and Artemisia ludoviciana ssp. incompta. 45 4 Meiosis in the Artemisia ludoviciana- complex. 54 5 First meiotic division of the desynaptic, tetraploid Artemisia douglasiana. 65 6 Metaphase I configuration in the trisomic hexaploid Artemisia douglasiana. 69 7 Abnormal hybrid seedlings. 77 8 Hooded .. condition in the S generation of Artemisia douglasiana (4x and 6x). 1 78 9 Genomic relationships in the Artemisia ludoviciana- complex. 91 10 Rhizomes of Artemisia cavaticaulis and Artemisia suksdorfii X Artemisia cavaticaulis F1 hybrids. 116 1 LIST OF TABLES Table Page 1 The reported frequency of quadrivalents in some autotetraploids (4x =28) in Gramineae. 10 2 Total configurations in the tetraploid Artemisia vulgaris at metaphase I. 26 3 Frequencies of diakinesis and metaphase I configurations in the diploids. 46 4 Frequency distribution of numbers of bivalents per cell in the diploids. 48 5 Chiasma .. frequency at metaphase I in the diploids. 49 6 Pollen viabilities of the different taxa and ploidy levels. 50 7 Frequencies of the types of sporads of microspores. 52 8 Frequencies of metaphase I configurations in the tetraploids. 56 9 Frequencies of metaphase I multivalent configurations in the tetraploids. 59 10 Maximum number of multivalents in the polyploids. 61 11 Chiasma frequency at metaphase I in the tetraploids. 62 12 Disjunctional ratios at anaphase I in a desynaptic tetraploid of Artemisia douglasiana. 66 13 Frequencies of metaphase 'I configurations in hexaploid Artemisia douglasiana. 68 14 Frequencies of metaphase I multivalent configurations in hexaploid Artemisia douglasiana. 68 15 Artificial hybrids. 71 Table Page 16 Numbers of dwarf and semi -dwarf progeny. 79 17 Cytology and pollen fertility of selected F1 hybrids. 81 18 A morphological comparison of the diploid and tetraploid races of Artemisia lindleyana. 110 19 A morphological comparison of the diploids, Artemisia suksdorfii and Artemisia cavaticaulis. 113 CYTOTAXONOMIC STUDIES IN THE ARTEMISIA LUDOVICIANA POLYPLOID COMPLEX OF THE PACIFIC NORTHWEST I. INTRODUCTION The Artemisia ludoviciana - complex is a polymorphic assem- blage of closely related taxa in the Artemisia vulgaris- aggregation. The complex occurs throughout western North America, and the included variants form a polyploid series with three known chromo- somal levels -- diploid, tetraploid, and hexaploid. The more recent taxonomic treatments of the complex (Keck, 1946; Cronquist, 1955; Ward, 1960) recognize several species, defined as series of popula- tions having correlated morphological characters, similar eco- geographic distribution, and a common chromosome number, if known. However, this taxonomic technique has proven to be difficult to apply in practice because of evident morphological continuity be- tween the taxa. The objective of this investigation is to apply cytogenetic methods to the taxonomic study of the members of this complex in northwestern North America, in order to determine the sources of its variation, the types of polyploidy present, and the genetic pro- cesses that have been active in the evolution of the complex. The results may lead to a better understanding of the systematic relation- ships of the taxa involved. The methods employed include 2 morphological and cytological observations of selected natural populations from the Pacific Northwest, transplant experiments with population samples, and cross -pollination of selected indivi- duals, followed by analysis of the resulting progeny. 3 II. LITERATURE REVIEW Classification of Polyploids The degree of homology between the contributed genomes is the basis for the classification of polyploid individuals. There is a range from autopolyploids with completely homologous genomes, to allo- polyploids with completely dissimilar genomes. Stebbins (1947) suggested the term "segmental allopolyploid" for those organisms intermediate between the two extremes, that is, similar in certain portions of the genome but dissimilar in others. Pairing relationships of the chromosomes during the first meiotic prophase and metaphase are the criteria most easily utilized to establish the degree of homology between the genomes of a poly - ploid. Lack of multivalent formation, however, is not necessarily evidence against chromosomal homology. For example, Riley (1960) investigated the genetic control
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