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194 7. ROLE OF QTLs IN THE EARLY EVOLUTION... 4. In total 145 QTLs govern the L. multiflorum and L. perenne evolution however, the most important traits related to domestication are governed by 51 QTLs. These traits involve height at ear emergence, spike and spikelet length, spikelet number, floret number, green and dry weight of tillers, green and dry weight of vegetative parts and winter survival. 5. Domestication of L. multiflorum and diversification from L. perenne is at least partially connected with transposons. This finding confirms the role of mobile genetic elements in the early evolution of L. multiflorum and L. perenne. 6. Implementing the same philosophy as it is done for maize and teosinte, or indica and japonica forms in rice, it is proposed to classify L. multiflorum and L. perenne with a rank of subspecies i.e., L. perenne ssp. perenne for a wild form and L. perenne ssp. multiflorum for a domesticated form.
8. MOLECULAR PHYLOGENY OF THE GENUS LOLIUM 8.1. INTRODUCTION Once evolutionary relationships between L. multiflorum and L. perenne are resolved the phylogenesis of the whole genus Lolium can be reconstructed. First, it should pointed out that all data presented within chapters 2-7 demonstrate clearly that taxonomic rank of L. multiflorum and L. perenne should be lowered to the subspecies level. Thereby both taxa should be reclassified following the Integrated Taxonomic System of the USA (2007), in which they have names L. perenne ssp. multiflorum and L. perenne ssp. perenne. Consequently, this classification is used in all subsequent parts. Nevertheless, to make the text more readable and to avoid repeating the long subspecies names over and over again the abbreviations are used, multiflorum for L. perenne ssp. multiflorum and perenne for L. perenne ssp. perenne. Phylogenesis is the sequence of events involved in the evolutionary development of a taxonomic group. Methods of reconstruction of phylogenetic trees were developed independently in numerical taxonomy (phenetic approach) for morphological traits and in population genetics for allele frequency data. Until then years of studies had been necessary to accumulate experience and classify species into a hierarchical system. The principles guiding phenetic approach are classifications based on overall similarity (or distance) between taxa measured by strictly defined rules and using as many characters as possible. Evolutionary relationships are presented graphically as phylogenetic trees or networks. External nodes are extant units; and internal nodes are deduced ancestral units. Branch lengths reflect the number of evolutionary changes along each ancestral-descendant pathway. The major advantage of numerical taxonomy is that a potential danger of authoritative selections of characters is limited. An alternative philosophy has been proposed by the cladistic school, according to which phylogeny should be appraised not by overall similarity, but rather by a set of similarities attributable to some shared-derived traits (Nei and Kumar 2000; Avise 2004). The basic difference between phylogenetic and cladistic approach is that the former tends to focus on branch lengths whereas the latter on the branching component. Indeed, there is a little justification for the differentiation of both approaches, first, because cladistic methods come close to phenetic ones especially as applied to large datasets. The second, more serious difficulty in all phylogenetic methods, either phenetic or cladistic is the assumption that the characters analysed are “independent”. This assumption is critical for most mathematical algorithms but unfortunately, it is probably not valid for the majority of morphological characters. Thus, although phenograms and cladograms based on morphological traits have
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This research was done within the E
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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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36 3. MORPHOLOGICAL VARIATION... fo
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38 3. MORPHOLOGICAL VARIATION...
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40 3. MORPHOLOGICAL VARIATION... Wi
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42 3. MORPHOLOGICAL VARIATION... va
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44 3. MORPHOLOGICAL VARIATION... 3.
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46 3. MORPHOLOGICAL VARIATION...
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48 3. MORPHOLOGICAL VARIATION... in
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50 3. MORPHOLOGICAL VARIATION... pl
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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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68 4. GENETIC DIVERSITY... It can b
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72 4. GENETIC DIVERSITY... than wit
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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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88 4. GENETIC DIVERSITY... individu
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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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244 10. MARKER EFFICIENCY proportio
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246 10. MARKER EFFICIENCY 10.7. CON
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248 11. CONCLUSIONS ABOUT EVOLUTION
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250 12, LITERATURE CITED Bączkiewi
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252 12, LITERATURE CITED Clayton WD
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254 12, LITERATURE CITED Giddings G
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256 12, LITERATURE CITED Jing R, Kn
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258 12, LITERATURE CITED MacDonald
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260 12, LITERATURE CITED Payne RC,
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262 12, LITERATURE CITED Schiex T,
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264 12, LITERATURE CITED United Sta
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266 12, LITERATURE CITED Zielinski
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268 13. SUPPLEMENTARY MATERIALS pla
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270 13. SUPPLEMENTARY MATERIALS
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274 13. SUPPLEMENTARY MATERIALS ANN
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280 13. SUPPLEMENTARY MATERIALS Ann
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290 13. SUPPLEMENTARY MATERIALS •
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292 13. SUPPLEMENTARY MATERIALS The
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296 13. SUPPLEMENTARY MATERIALS fie
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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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