Chapter 6been detected with GISH and FISH, and later on a U-type reunion led to the formation <strong>of</strong>anaphase bridges and fragments (Chapter 4). F<strong>in</strong>ally, microspores with different <strong>chromosome</strong>numbers have also been detected after meiosis (Zhou et al. 2008a). In conclusion,<strong>in</strong>tersectional lily hybrids show a range <strong>of</strong> abnormalities dur<strong>in</strong>g different stages <strong>of</strong> meiosis.Some other k<strong>in</strong>ds <strong>of</strong> meiotic abnormalities <strong>in</strong> <strong>in</strong>terspecific lily hybrids have also beenreflected and emphasized by progeny <strong>analysis</strong>. The first evidence is the polyploidizedbackcross progenies. The resultant progenies from crosses <strong>in</strong>volv<strong>in</strong>g <strong>in</strong>terspecific lily hybridswere predom<strong>in</strong>ant triploids, <strong>in</strong>dicat<strong>in</strong>g the functional gametes were unreduced gametes andthe mechanism has been identified as first division restitution (FDR) and <strong>in</strong>determ<strong>in</strong>atemeiotic restitution (IMR) (Lim et al. 2001a). The second feature <strong>in</strong> backcross progenies <strong>of</strong> lilyis aneuploidy. When analyz<strong>in</strong>g the genomic composition <strong>of</strong> these triploid lily hybrids, a smallproportion <strong>of</strong> aneuploids has been found. The last character <strong>of</strong> the backcross progeny is thepresence <strong>of</strong> iso<strong>chromosome</strong>s. In a few genotypes, result<strong>in</strong>g from some <strong>in</strong>terspecific hybrids <strong>of</strong>LA lilies, iso<strong>chromosome</strong>s with different sizes were detected, and these newly-generatedsmall aberrant <strong>chromosome</strong>s were derived from the fusion <strong>of</strong> the two short arms <strong>of</strong> themiss<strong>in</strong>g <strong>chromosome</strong>s dur<strong>in</strong>g meiosis, respectively (Chapter 5).Cross<strong>in</strong>g over and <strong>in</strong>trogression breed<strong>in</strong>gThe role <strong>of</strong> cross<strong>in</strong>g over dur<strong>in</strong>g evolution and speciation has long been realized and studied<strong>in</strong> flower<strong>in</strong>g plants. Cross<strong>in</strong>g over, which is one <strong>of</strong> the key features that dist<strong>in</strong>guish meiosisfrom mitosis, not only facilitates the proper segregation <strong>of</strong> homologous <strong>chromosome</strong> <strong>in</strong> thefirst meiotic division, but also generates novel comb<strong>in</strong>ations <strong>of</strong> alleles via homologous<strong>chromosome</strong> exchanges. This process, <strong>in</strong> addition to ma<strong>in</strong>ta<strong>in</strong><strong>in</strong>g the ploidy level dur<strong>in</strong>gsexual reproduction, contributes to genetic diversity, which is essential for <strong>in</strong>trogressionbreed<strong>in</strong>g.Cross<strong>in</strong>g over between homoeologous chromatids has been proven to be less frequent ascompared with cross<strong>in</strong>g over between homologous non-sister chromatids. In monosomicadditions <strong>of</strong> tomato, a homologous bivalent (II) together with a univalent was the ma<strong>in</strong>meiotic configuration, GISH has revealed that the number <strong>of</strong> rod bivalents (stands for s<strong>in</strong>glecross<strong>in</strong>g over) was much higher compared with that <strong>of</strong> r<strong>in</strong>g bivalents (stands for other types <strong>of</strong>cross<strong>in</strong>g over which probably lead to <strong>chromosome</strong>s with two or more recomb<strong>in</strong>ant sites),<strong>in</strong>dicat<strong>in</strong>g s<strong>in</strong>gle cross<strong>in</strong>g over was the predom<strong>in</strong>ant type <strong>of</strong> exchange between homologous<strong>chromosome</strong>s. While <strong>in</strong> the substitution l<strong>in</strong>e <strong>of</strong> tomato SL-8, reduction <strong>of</strong> homoeologousrecomb<strong>in</strong>ation has been revealed by the considerable decrease <strong>of</strong> r<strong>in</strong>g bivalent formation (Jiand Chetelat 2003). Similarly, results from several studies <strong>of</strong> homeologous recomb<strong>in</strong>ationbetween <strong>chromosome</strong>s <strong>of</strong> wheat and related species have showed the absence <strong>of</strong> multiplecrossovers (Dubcovsky et al. 1995; Lukaszewski 1995, 2000; Luo et al. 1996; Luo et al. 2000).Homoeologous cross<strong>in</strong>g over has been proven to occur with different frequencies <strong>in</strong>different species hybrids. Although different types <strong>of</strong> cross<strong>in</strong>g over events have been checked80
General Discussiondur<strong>in</strong>g meiosis <strong>in</strong> the <strong>in</strong>terspecific hybrids <strong>of</strong> lily (Chapter 3), the number <strong>of</strong> <strong>chromosome</strong>s <strong>in</strong>the half-bivalents with two or more recomb<strong>in</strong>ant sites is low, compared with those with onerecomb<strong>in</strong>ant site. S<strong>in</strong>ce different crossover events have certa<strong>in</strong> segregation patterns ( such ass<strong>in</strong>gle crossover produces two recomb<strong>in</strong>ant <strong>chromosome</strong>s each with one recomb<strong>in</strong>ant site,with the exception <strong>of</strong> multiple crossover), the 637 pairs <strong>of</strong> half-bivalents <strong>in</strong> pollen mothercells <strong>in</strong> Chapter 3 showed 1191 recomb<strong>in</strong>ant <strong>chromosome</strong>s <strong>in</strong> total. 1102 <strong>chromosome</strong>s,which occupied 92.5%, possessed one recomb<strong>in</strong>ant site and 89 <strong>chromosome</strong>s (7.5%) with tworecomb<strong>in</strong>ant sites. Although <strong>chromosome</strong>s with more than two recomb<strong>in</strong>ant sites did occurdur<strong>in</strong>g meiosis, the frequency is relatively low compared with other species hybrids. Inpolyploid cotton (Gossypium), the frequency <strong>of</strong> <strong>in</strong>tergenomic recomb<strong>in</strong>ation events possessedone, two, three or more recomb<strong>in</strong>ant sites were 70.3%, 20.6% and 9.1% respectively (Salmonet al. 2010). Similarly, <strong>in</strong> an alien substitution l<strong>in</strong>e <strong>of</strong> tomato, <strong>in</strong> which <strong>chromosome</strong> with twobreakpo<strong>in</strong>ts took up around 15% <strong>of</strong> the total recomb<strong>in</strong>ant <strong>chromosome</strong>s (Tam et al. 2011), thepercentage <strong>of</strong> <strong>chromosome</strong>s with more than 1 recomb<strong>in</strong>ant site <strong>in</strong> lily is considerably low.There are three potential reasons for the low frequency <strong>of</strong> <strong>chromosome</strong>s with two or morerecomb<strong>in</strong>ant sites <strong>in</strong> lily hybrids: 1) the genomes <strong>of</strong> the lily parents are more divergentcompared those <strong>in</strong> cotton and as a result, complicated cross<strong>in</strong>g overs with multiplerecomb<strong>in</strong>ant sites on each <strong>chromosome</strong> are suppressed; 2) gene conversion, which usuallygives rise to two or more recomb<strong>in</strong>ant sites <strong>in</strong> genetic mapp<strong>in</strong>g and can be detected by mRNAsequenc<strong>in</strong>g, occurs frequently <strong>in</strong> cotton; or 3) s<strong>in</strong>ce the limited resolution <strong>of</strong> GISH, such geneconversions or small <strong>in</strong>trogressed <strong>chromosome</strong> segments cannot be detected by <strong>molecular</strong><strong>cytogenetic</strong> methods, which gives an underestimation <strong>of</strong> recomb<strong>in</strong>ant sites on <strong>chromosome</strong>s.Genomic shock, iso<strong>chromosome</strong> formation and B <strong>chromosome</strong>orig<strong>in</strong> dur<strong>in</strong>g sexual polyploidizationGenomes fac<strong>in</strong>g stress will suffer genomic shock which, on a chromosomal level, leads tostructure remodel<strong>in</strong>g (McCl<strong>in</strong>tock 1984). All k<strong>in</strong>ds <strong>of</strong> structure remodel<strong>in</strong>g (structurevariation) experience a process that <strong>in</strong>volves double strand breaks (DSBs, <strong>chromosome</strong>breakage <strong>in</strong> <strong>cytogenetic</strong>s) and error-reunions. DSBs can happen at centromere (centric fission),<strong>in</strong> <strong>in</strong>terstitial or term<strong>in</strong>al regions on a <strong>chromosome</strong>, and error-reunion <strong>of</strong> broken <strong>chromosome</strong>sgive rise to the production <strong>of</strong> structure variation. A simple example is that <strong>chromosome</strong>breakage followed by the fusion <strong>of</strong> broken arms from different <strong>chromosome</strong>s leads to thegeneration <strong>of</strong> so-called Robertsonian translocation <strong>in</strong> humans (Perry et al. 2004). In view <strong>of</strong>this, genomic shock is the driver <strong>of</strong> <strong>chromosome</strong> breakage, which causes erroneous repair <strong>in</strong>plants. It is not surpris<strong>in</strong>g that <strong>in</strong>terspecific hybridization leads to spontaneous <strong>chromosome</strong>breakage, which has been detected <strong>in</strong> Chapter 4. As a second step, error-reunion leads tovarious types <strong>of</strong> <strong>chromosome</strong> rearrangements, <strong>in</strong>clud<strong>in</strong>g chromosomal <strong>in</strong>versions, deletions,translocations, and duplication (Britt 1999).81
- Page 3 and 4:
A molecular cytogenetic analysis of
- Page 5:
Table of ContentsChapter 1General I
- Page 8 and 9:
Chapter 1LilyLilies belong to genus
- Page 10 and 11:
Chapter 1counterparts (Finnegan 200
- Page 12 and 13:
Chapter 1renewed interest in detect
- Page 14 and 15:
Chapter 1recombination, mechanisms
- Page 16 and 17:
Chapter 1is only an equational segr
- Page 18 and 19:
Chapter 1quite divergent with vario
- Page 21 and 22:
Chapter 2An assessment of chromosom
- Page 23 and 24:
Chromosome rearrangements in Lilium
- Page 25 and 26:
Chromosome rearrangements in Lilium
- Page 27 and 28:
Chromosome rearrangements in Lilium
- Page 29 and 30:
Chromosome rearrangements in Lilium
- Page 31 and 32:
Chromosome rearrangements in Lilium
- Page 33 and 34:
Chromosome rearrangements in Lilium
- Page 35 and 36: Chromosome rearrangements in Lilium
- Page 37 and 38: Chapter 3Elucidation of intergenomi
- Page 39 and 40: Intergenomic recombination and chro
- Page 41 and 42: Intergenomic recombination and chro
- Page 43 and 44: Intergenomic recombination and chro
- Page 45 and 46: Intergenomic recombination and chro
- Page 47 and 48: Intergenomic recombination and chro
- Page 49: Intergenomic recombination and chro
- Page 52 and 53: Chapter 4AbstractMeiotic abnormalit
- Page 54 and 55: Chapter 4There are some criteria to
- Page 56 and 57: Chapter 4FISH experiments were perf
- Page 58 and 59: Chapter 4was apparently shorter, wi
- Page 60 and 61: Chapter 4bridges is explained as U-
- Page 62 and 63: Chapter 4of the sexual polyploidize
- Page 64 and 65: Chapter 4fragment have the same len
- Page 66 and 67: Chapter 5AbstractSupernumerary (B)
- Page 68 and 69: Chapter 5their origin, the structur
- Page 70 and 71: Chapter 5Fig. 5.1. Discovery of B c
- Page 72 and 73: Chapter 5In order to investigate th
- Page 74 and 75: Chapter 5Centric breakage and fusio
- Page 76 and 77: Chapter 5It has not, however, been
- Page 78 and 79: Chapter 6The results presented in t
- Page 80 and 81: Chapter 6model for molecular cytoge
- Page 82 and 83: Chapter 6Fig. 6.2. The meiosis proc
- Page 84 and 85: Chapter 6over events during FDR mei
- Page 88 and 89: Chapter 6Another feature caused by
- Page 91 and 92: ReferencesAbe, H.A., Nakano, M.N.,
- Page 93 and 94: ReferencesChen, Q., and Armstrong,
- Page 95 and 96: ReferencesHartlerode, A.J., and Scu
- Page 97 and 98: ReferencesLarson, S.R., Kishii, M.,
- Page 99 and 100: ReferencesMcClintock, B. 1931. Cyto
- Page 101 and 102: ReferencesRai, R., Zheng, H., He, H
- Page 103 and 104: ReferencesStewart, R.N. 1947. The m
- Page 105: ReferencesZhang, L., Pickering, R.,
- Page 108 and 109: Summarychromosome rearrangements. T
- Page 111 and 112: SamenvattingLelie (Lilium) is in de
- Page 113 and 114: Samenvattingaantal 35 met daarnaast
- Page 115 and 116: 摘 要百 合 系 百 合 科 百
- Page 117 and 118: Acknowledgements淡 看 世 事 去
- Page 119: Curriculum VitaeSonglin Xie was bor
- Page 123: Education Statement of the Graduate