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A molecular cytogenetic analysis of chromosome behavior in Lilium ...

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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

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