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

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Chapter 6Another feature caused by <strong>in</strong>terspecific hybridization is the occurrence <strong>of</strong> univalentsdur<strong>in</strong>g meiosis <strong>in</strong> <strong>in</strong>terspecific hybrids. Univalents, which arose from association failure, havebeen found <strong>in</strong> many <strong>in</strong>terspecific hybrids, and are considered to be the ma<strong>in</strong> reason <strong>of</strong> thereduced fertility (Asano 1982; Lee et al. 2011; Lukaszewski 2010; Sears 1950). Meioticunivalents not only randomly move to one pole when segregat<strong>in</strong>g at anaphase I, but also havea tendency to misdivide at the centromere (Lukaszewski 2010). Centromere misdivision givesrise to centric translocation, production <strong>of</strong> telocentric and iso<strong>chromosome</strong>s, which have beenfound <strong>in</strong> maize and wheat (Kaszas et al. 2002; Lukaszewski 2010). Chapter 5 reported theproduction <strong>of</strong> iso<strong>chromosome</strong>s, which were derived from centric fission and fusion dur<strong>in</strong>gmeiosis <strong>of</strong> the maternal parent. Meanwhile, <strong>chromosome</strong> breakage has been found not only <strong>in</strong>univalents, but also <strong>in</strong> bivalents (Chapter 4). As a result, it is still not known whether thenewly-generated iso<strong>chromosome</strong>s <strong>in</strong> the backcross progenies are the result <strong>of</strong> centric breakageand fusion from either an univalent or a bivalent.B <strong>chromosome</strong>s, which extensively exist <strong>in</strong> many flower<strong>in</strong>g plants, are probably derivedfrom aberrant <strong>chromosome</strong>s. It is already well accepted that B <strong>chromosome</strong>s orig<strong>in</strong>ate frommeiotic errors <strong>in</strong> which <strong>in</strong>terspecific hybridization provides an ideal platform, and this type <strong>of</strong><strong>chromosome</strong>s are deduced to be escaped from standard <strong>chromosome</strong>s (Jones and Houben2003). However, what should be noticed is that the orig<strong>in</strong> <strong>of</strong> B <strong>chromosome</strong>s is not a one-stepprocess, which has been shown by Dhar et al. (Dhar et al. 2002) <strong>in</strong> Plantago. Comb<strong>in</strong>ed withthe fact that most <strong>of</strong> the species are <strong>in</strong>volved <strong>in</strong> at least one round polyploidization, it can beconcluded that B <strong>chromosome</strong>s arose <strong>in</strong> the process <strong>of</strong> speciation <strong>of</strong> polyploids <strong>in</strong><strong>in</strong>terspecific hybrids, which has been shown by the production <strong>of</strong> small aberrant<strong>chromosome</strong>s dur<strong>in</strong>g sexual polyploidization <strong>of</strong> lily hybrids.Conclusions and future perspectivesAs presented <strong>in</strong> this thesis, it has been shown that not only <strong>in</strong>tergenomic recomb<strong>in</strong>ation whichis derived from crossovers, but also <strong>chromosome</strong> rearrangements causes genetic variation <strong>in</strong>backcross progenies <strong>of</strong> lily. Moreover, <strong>chromosome</strong> breakage and fusion lead to theproduction <strong>of</strong> <strong>chromosome</strong> bridges at anaphase I stage dur<strong>in</strong>g meiosis and the generation <strong>of</strong>small aberrant <strong>chromosome</strong>s <strong>in</strong> the backcross progenies. However, to apply these results <strong>in</strong>practical breed<strong>in</strong>g, the follow<strong>in</strong>g research should also be done <strong>in</strong> the future:Although crossover events have been studied <strong>in</strong> this thesis, it is necessary to study it on thelevel <strong>of</strong> <strong>in</strong>dividual <strong>chromosome</strong>s. S<strong>in</strong>ce recomb<strong>in</strong>ation sites on different <strong>chromosome</strong>s arehighly uneven (Khan et al. 2009a), it is a precondition to make an accurate identification <strong>of</strong><strong>in</strong>dividual <strong>chromosome</strong>s. Traditional methods to identify <strong>chromosome</strong>s are based on<strong>chromosome</strong> length, arm length, arm length ratio and so on, which makes it difficult torecognize <strong>chromosome</strong>s with short arms s<strong>in</strong>ce most lily <strong>chromosome</strong>s are morphologicalsimilar (Noda 1978; Stewart 1947). Later on, a few efforts were made to dist<strong>in</strong>guish<strong>chromosome</strong>s with different band<strong>in</strong>g techniques (Smyth et al. 1989; Von Kalm and Smyth82

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