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248 11. CONCLUSIONS ABOUT EVOLUTION... Apart from L. multiflorum and L. perenne the present molecular studies based on enormous number of 2984 molecular markers representing various methodologies have prompted major revisions of thoughts concerning evolutionary relationships within the genus Lolium. It is now clear that the progenitor of the genus was common to Lolium and F. pratensis, had the closest affinity to Schedonorus taxa and gave rise to two lineages, of which diversification began 2.7 MYA. Today they are represented by the Temulentum clade consisted from autogamous annual species, L. persicum, L. remotum and the best known member of this clade - L. temulentum. L. persicum was the first species that differentiated about 1.3 MYA, then the remaining two species were split about 0.95 MYA. Nevertheless, as weeds of crops, all of them might have been domesticated unconsciously in days of the primitive agriculture, circa about 10 000 years ago. The second lineage is represented by the Perenne clade that involves allogamous species L. perenne with two subspecies, annual or biennial L. perenne ssp. multiflorum, perennial L. perenne ssp. perenne, annual L. rigidum as well as autogamous L. loliaceum. This clade diverged from the common ancestor with F. pratensis about 2.35 MYA. The earliest diverged species was L. rigidum that split from the others about 1.4 MYA. L. loliaceum evolved through several point mutations related with the transition from allogamy to autogamy and its divergence from L. perenne can be dated to 1.2 MYA. The populations of L. perenne started to diversify during the Tertiary and Quaternary glaciations. It is likely that the perennial forms came into existence 167 000 years ago as a response to more harsh environment. Further diversification has been prompted by the domestication process that started before twelfth century in Italy, of which we are eyewitnesses today. A spectacular discover is that transposons are the most important driving forces of evolutionary processes observed in multiflorum and perenne. It is elegantly shown by their higher mobility in L. perenne ssp. multiflorum observed in genetic diversity studies, strong distortions from the expected Mendelian segregation ratios and their association with the QTLs noticed in mapping studies. From these wide molecular surveys one the most important notion emerges that the evolutionary properties of any given group of organisms could be characterized by a number of methods addressing to different levels of the same phenomena. Only then the reasonably clear picture of genetic diversity can be obtained taking into account the present development of technologies. Among many technologies used in the studies on Lolium, it is worth mentioning markers derived from bacterial sequences (B-SAP). It was a great surprise to be able to amplify plant DNA using primers specific to so distant organism as M. tuberculosis. The astonishment was even greater when it was possible to map markers derived from catalase-peroxidase of M. tuberculosis in close proximity to peroxidase loci and the other enzymatic loci. This example demonstrates how fascinating and strange can be the evolutionary roads. And for the first time, we have tools which, if coupled with advance statistics and traditional morphology have potential to reveal secrets of evolutionary changes and help us to understand and protect enormous diversity of living creatures. And last but not least, new methods that will emerge in the nearest future will certainly add a new look at the genus Lolium, at any other taxa and at the evolution of life on the Earth.
12. LITERATURE CITED Aguirre-Planter E, Furnier GR, Equiarte LE. 2000. Low levels of genetic variation within and high levels of genetic differentiation among populations of species of Abies from southern Mexico and Guatemala. Am J Bot 87:362-5371. Application of DNA based marker mutations for improvement of cereals and other sexually reproduced crop plants. IAEA-TECDOC-1010, IAEA Vienna (1998). Arcioni S, Mariotti D. 1983. Selfing and interspecific hybridization in Lolium perenne L. and L. multiflorum Lam. evaluated by phosphoglucoisomerase as izozyme marker. Euphytica 32:33-40. Armstead IP, Turner LB, King IP, Cairns AJ, Humphreys MO. 2002. Comparison and integration of genetic maps generated from F 2 and BC 1 -type mapping populations in perennial ryegrass. Plant Breeding 121:501-507. Avise JC. 2004. Molecular markers, natural history, and evolution. 2 nd Edition. Sunderland, Massachusetts: Sinauer Associates, Inc. Publishers. 684 p. Axelrod B, Belzile JR. 1958. Isolation of an alkaloid, annuloline, from the roots of Lolium multiflorum. J Org Chem 23:919-920. Babu R, Nair SK, Prasanna BM, Gupta HS. 2004. Integrated marker-assisted selection in crop breeding - prospects and challenges. Curr Sci India 87:607-618. Bachman K. 2000. Molecules, morphology and maps: new directions in evolutionary genetics. Plant Spec Biol 15:197-210. Balfourier F, Charmet G, Ravel C. 1998. Genetic differentiation within and between natural populations of perennial and annual ryegrass (Lolium perenne and L. rigidum). Heredity 81:100- 110. Balfourier F, Imbert C, Charmet G. 2000. Evidence for phylogenetic structure in Lolium species related to the spread of agriculture in Europe. A cpDNA study. Theor Appl Genet 101:131- 138. Barbash DA, Siino DF, Tarone AT, Roote J. 2003. A rapidly evolving MYB-related protein causes species isolation in Drosophila. Proc Natl Acad Sci USA 29:5302-5307. Barker RR, Warnke SE.1999. Genetic separation of annual from perennial ryegrass. J Seed Tech- J Seed Technol 16:74-75.
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Acknowledgements I would like to th
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8 CONTENTS 4.4.4. Role of transposo
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10 CONTENTS 9.3.2. Similarity of ge
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12 1. INTRODUCTION
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14 1. INTRODUCTION occasionally use
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16 1. INTRODUCTION On the basis of
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18 1. INTRODUCTION The evidences fr
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20 1. INTRODUCTION overlap between
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22 1. INTRODUCTION et al. 2000; Cat
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24 1. INTRODUCTION According to the
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26 1. INTRODUCTION 1.6. APPLICATION
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28 1. INTRODUCTION of evolution wit
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30 1. INTRODUCTION are continuously
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2. GOALS The aim of this research w
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3. MORPHOLOGICAL VARIATION OF L. MU
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38 3. MORPHOLOGICAL VARIATION...
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44 3. MORPHOLOGICAL VARIATION... 3.
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52 3. MORPHOLOGICAL VARIATION... di
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4. GENETIC DIVERSITY OF L. MULTIFLO
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56 4. GENETIC DIVERSITY... deed con
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58 4. GENETIC DIVERSITY... 4.2.2. D
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60 4. GENETIC DIVERSITY...
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64 4. GENETIC DIVERSITY... RAPD and
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66 4. GENETIC DIVERSITY... SSAP pol
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82 4. GENETIC DIVERSITY... power of
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84 4. GENETIC DIVERSITY... Hamrick
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86 4. GENETIC DIVERSITY... deviatio
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90 4. GENETIC DIVERSITY... (inversi
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92 4. GENETIC DIVERSITY... related
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94 4. GENETIC DIVERSITY... cies and
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96 4. GENETIC DIVERSITY... their sp
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98 4. GENETIC DIVERSITY... 4.5. CON
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100 5. ORIGIN OF SEEDLING... Applic
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102 5. ORIGIN OF SEEDLING... 5.2.2.
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104 5. ORIGIN OF SEEDLING... 5.3. R
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106 5. ORIGIN OF SEEDLING... very l
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108 5. ORIGIN OF SEEDLING... 5.3.4.
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110 5. ORIGIN OF SEEDLING... 5.4. D
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112 5. ORIGIN OF SEEDLING... A seco
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114 5. ORIGIN OF SEEDLING... Perhap
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6. DO ANY SPECIES BOUNDARIES EXIST
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118 6. DO ANY SPECIES BOUNDARIES...
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7. ROLE OF QTL s IN THE EARLY EVOLU
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196 8. MOLECULAR PHYLOGENY OF THE G
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Page 250 and 251: 250 12, LITERATURE CITED Bączkiewi
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298 13. SUPPLEMENTARY MATERIALS ANN
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300 13. SUPPLEMENTARY MATERIALS ANN
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302 13. SUPPLEMENTARY MATERIALS Ann
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The average gene diversity between
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306 13. SUPPLEMENTARY MATERIALS Unb
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308 14. ABBREVIATIONS Lolcopia1 - L
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310 14. ABBREVIATIONS Drosophila wi
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312 14. ABBREVIATIONS Saccharomyces
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314 15. SUMMARY At different stages
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316 15. SUMMARY L. multiflorum mito
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REVIEWERS’ COMMENTS With a great
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