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多倍化遗传效应对拟南芥同源染色体减数分离及基因重组的影响
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摘要
在植物界,大约70%的被子植物都曾发生过多倍化现象。多倍化在植物界物种进化过程中有着非常重要的意义,是植物进化和变异的重要推动力量,也是保持物种多样性和稳定性的重要途径。因此,在多倍体植物中,深入研究和探索多倍化效应对植物生殖和发育的影响,对揭示多倍体植物的进化和变异有着重要的意义。正是基于此,本研究充分利用模式植物拟南芥生殖和发育优越性,采用人工同源或异源多倍化技术,成功合成出同源和异源四倍体拟南芥,并结合与植物有性生殖紧密相关的减数分裂过程,通过分子及细胞遗传学手段,深入研究了多倍化效应对基因突变、基因重组、配子传递等的影响,获得以下重要结论:
     1.对拟南芥二倍体中3种减数分裂突变体spo11-1、dmc1和asy1的细胞遗传学研究和相应表达蛋白质产物序列比对分析发现,spo11-1功能是在减数分裂间期启动DNA双链产生缺口,dmc1与其他修复蛋白构成修复复合体,对产生缺口的DNA单链进行同源依赖性修复,asy1对减数分裂前期I同源染色质(体)联会和中期I同源染色体配对有重要作用。虽然3个基因功能不同和表达时期不同,但在细胞遗传学上具有相似性,3个基因突变都能导致同源染色体联会和配对失败,最终减数分裂期间以单价体形式出现。但是,3种基因突变导致的细胞遗传变化程度不同,spo11-1和dmc1导致的突变效应较为强烈,减数分裂期同源染色体完全以单价体出现,而asy1突变体中,平均每个减数分裂细胞能产生1.26个二价体。此外,序列比对发现,3种基因在植物界较为保守,对保证植物正常的有性生殖有重要意义。
     2.通过人工多倍化,本研究成功合成了同源四倍体拟南芥A.thaliana。对多倍化后的突变体dmc1和asy1减数分裂过程研究发现,在同源四倍体中,dmc1基因导致的突变效应较为强烈,减数分裂过程未发现二价体或四价体的形成;而同源四倍体突变体asy1中,减数分裂过程中,二价体数量增多,并能检测到部分四价体现象。因此,本实验认为,多倍化可以缓冲基因突变带来的不良影响,为多倍体通过有性生殖延续后代提供条件。
     3.对同源四倍体突变体asy1有性生殖过程中非整倍体配子传递规律的研究发现,在细胞遗传学上,减数分裂后配子染色体数目决定了配子传递的方式,非整倍性亚倍体配子传递主要通过雄性配子,而超倍体配子传递可以通过雌雄配子,雌配子体主要传递超倍体。此外,双受精后,来自不同亲本的染色体数目和父母本间基因组含量相对平衡性对植物发育有重要影响
     4.在同源多倍体中,以一对紧密连锁基因GFP和RFP(G-R,物理距离约为5Mb)为模型,估计了同源多倍体减数分裂过程中多价体联会和配对对基因重组频率影响,并在实际研究中得到了证实。对于一对物理距离较近的连锁基因G-R,减数分裂期同源染色体间的多价体,如三价体和四价体的形成并不影响基因重组的频率。
     5.在二倍体拟南芥中,对连锁基因G-R的重组频率的研究发现,位于拟南芥3号染色体上的平均重组率为16.8%,父本重组率为20.2%,母本重组率为7.4%;位于5号染色体上的G-R连锁基因,平均重组率为14.9%,父本重组率为18.1%,母本重组率为14.0%,因此,本研究认为,同一对基因G-R重组率受染色体、染色体上位置的影响。此外,基因重组也受性别的影响,一般父本重组率较高。
     6.利用连锁基因G-R,通过同源和异源多倍化手段,分别对拟南芥同源四倍体A.thaliana和异源四倍体A.suecica基因重组频率进行了研究,研究结果表明,与二倍体相比较,同源和异源多倍化都可以导致基因重组频率的显著提高。同源四倍体A.thaliana中,连锁基因G-R的平均重组率为23.2%,父本重组率为28.0%,母本重组率为15.0%。异源四倍体A.suecica中,连锁基因G-R的平均重组率为28.2%,父本重组率为29.8%,母本重组率为13.0%。因此,多倍化后,性别特异性基因重组频率有所变化,都受到多倍化效应的影响。
     7.结合荧光原位杂交技术,针对不同来源拟南芥染色体组,通过利用特异性探针,研究结果表明,在新合成异源四倍体拟南芥A.suecica中,由于基因组冲击(genomic shock)的影响,尽管基因表达受到某种程度的影响,但是异源四倍体形成后,可能由于类似Ph1基因的存在,在减数分裂过程中,能进行正常的减数分裂,同源染色体联会和配对的机制不受干扰,保证了同源染色体间的等位基因交叉互换,为产生功能性配子提供细胞学物质基础。
     8.细胞遗传学研究表明,在异源多倍体中,减数分裂中同源染色体多价体的联会和配对并不能导致基因重组频率提高,因此,关于多倍化效应导致基因重组频率改变的内在原因有待于进一步研究。
In plant kingdom, approximately 70% of flowering plants have experienced polyploidization. Polyploidization has a meaningful role in plant evolutionary history, and is a driving force for evolution and variation in plants, and is a pathway for plants to maintain genetic diversification and genomic stability. Therefore, to explore the effects of polyploidy on plant sexual reproduction and development can undoubtedly contribute to our understandings of tremendous events of polyploidization in plants. In the present study, the model plant Arabidopsis was used for its fast growth and development to artificially synthesize polyploid pants including autotetraplod A.thaliana and allotetraploid A.suecica (by crossing between A.thaliana and A.arenosa). The meiotic progression of newly synthesized allotetraploid A.suecica was analysed. In combination with 3 meiosis-related genes spo11-1, dmc1 and asy1, the effects of polyploidy on meiosis and transmission of sexually reproductive gametes were also exploited. Besides, aided with a pair of cis-linked transgenic fluorescent marks G-R, the frequency of meiotic recombination was investigated as well. The results were presented as follows:
     1. In diploid A.thaliana, 3 mutant genes were investigated during meiosis. Spo11-1 was involved in initiation of double-strand breaks (DSBs), which further affected the meiotic recombination. Dmc1 played a key role in repairing DSBs by contributing to fixing the protruded 3’single DNA strands. Asy1 was a key component controlling synapsis and pairing of homologous chromatins (chromosomes). The results showed that, although the 3 genes examined here have distinct functions during meiosis respectively, the cytological observations of meiotic chromosomes were very similar. Mutation of the 3 genes can result in the failure of synapsis and pairing of homologous chromosomes, and chromosomes displayed as a unique formation of univalents in spo11-1 and dmc1, whereas in asy1 the rare formation of bivalents was infrequently observed at 1.26 per cell. Besides, the alignment of amino acid sequences of spo11-1, dmc1 and asy1 from other systems such as human, yeast, wheat and rice showed that 3 genes are highly conserved in sequence similarity which indicated an important role performed by these 3 genes in plant sexual life.
     2. After polyploidization, the autotetraploid Arabidopsis was obtained in the mutants dmc1 and asy1. The effects of polyploidization on meiotic progression in tetraploid dmc1 and asy1 were observed. The results demonstrated that the defects caused by mutation of dmc1 was more severe than that of asy1, and no improved meiosis was observed in tetraploid dmc1, but an increasing number of bivalents, even quadrivalents were frequently observed. These results indicated that polyploidy of plants can buffer the defectiveness brought about by specific mutations.
     3. The transmission of aneuploids was investigated in autotetraploid asy1, and the results showed that chromosome number in the produced gametes determines the pathway of aneuploid transmission. Transmission of hypoploids was mainly dependent on male gametes, whereas hyperploids have a dual pathway of transmission, through male or female. And chromosome number and relative ratio balance of genomic contents between male and female determine on the fate of development after fertilization.
     4. The statistical model on estimation of frequency of meiotic recombination based on a pair of cis-linked markers GFP and RFP was established on condition that the linked markers were closely located on a chromosme (5 Mb). The model indicated that the formation of multivalents in autotetraploid contributed little to the frequency change of meiotic recombination, which was proved further with our solid data.
     5. In diploid Arabidopsis, the recombination frequency of the cis-linked markers was estimated on chromosome 3 and chromosome 5 respectively. The results showed that on chromosome 3, the averaged frequency of homologous recombination was 16.8%, and the frequency in male was 20.2% and 7.4% in female. However, on chromosome 5, the averaged frequency of homologous recombination was 14.9%, and the frequency in male was 18.1% and 14.0% in female. These data indicated that recombination frequency was affected by a series of factors including chromosome of genes, locus of gene on chromosome. Besides, sex-related difference of recombination was generally observed with a higher frequency in male than female.
     6. After polyploidization including autopolyploidization and allopolyploidization, the recombination frequency of the same cis-linked markers was investigated in autotetraploid and allotetraploid Arabidopsis. The data indicated that in contrast with diploid Arabidopsis, autotetraploid has averaged 23.2% of recombination frequency, 28.0% in male and 15.0% in female. Comparatively, in allotetraploid, the averaged frequency of recombination was 28.2%, and 29.8% in male, and 13.0% in female. These results indicate that polyploidization contributes much to the increased frequency of meiotic recombination, and sex-related difference of recombination existed as well in polyploid plants.
     7. Aided with fluorescence in situ hybridization (FISH), the newly formed allotetraploid Arabidopsis was analyzed during meiosis. The observations showed that normal meiotic process was guaranteed with possible presence of wheat Ph1-like gene albeit the genomic shock of dirrerent chromosome resources. The synapsis and pairing of homologous chromosomes progress without interference of homoeologous chromosomes, which provided the opportunity for recombination and a basis for production of functional gametes after meiosis.
     8. Cytogenetic analysis demonstrated that formation of multivalents in polyploid plants can not account for the increased frequency of recombination, thus the further work was needed for unraveling the mechanism of increased recombination frequency brought about by polyploidy associated effects.
引文
鲍文奎,秦瑞珍,吴德瑜,等.高产四倍体水稻无性系[J].中国农业科学,1985,18(6):64-66.
    蔡得田,陈建国,陈冬玲,等.两个具多倍体减数分裂稳定性的多倍体水稻品系的选育[J].中国科学(C辑),2007,37(2):217-226.
    蔡得田,陈冬玲.水稻无融合生殖研究进展[J].华中农业大学学报,1992,11(l):15-20.
    常金华,罗耀武.同源四倍体高粱与约翰逊草杂交及其后代表现[J].草业学报,2002,11(1):56-58.
    崔平,潘荣.甜菜种质的性状改良与创新[J].中国糖料,2004(3):51-53.
    陈志勇,吴德瑜,宋文昌,等.同源四倍体水稻育种研究的近期进展[J].中国农业科学,1987,20(1):20-24
    陈劲枫,雷春,钱春桃,等.黄瓜多倍体育种中同源四倍体的合成和鉴定[J].植物生理学通讯,2004,40(2):149-152.
    黄群策,秦广雍,梁秋霞.同源四倍体水稻与狼尾草杂交结实的胚胎学证据[J].杂交水稻,2004,19(l):56-59.
    黄群策,向茂成,汤国雄.同源四倍体水稻与假稻杂交结籽的胚胎学机理研究[J].作物学报,2002,28(2):286-288.
    金佩芳,李世诚,蒋爱丽,等.早熟大粒葡萄新品种"申秀"育成[J]上海农业学报,1998,14(2):34
    李树贤,吴志娟,杨志刚,等.同源四倍体茄子品种新茄一号的选育[J].中国农业科学,2002,35(6):686-689
    李树贤.同源四倍体茄子多亲本轮回选择C5代遗传变异性的分析[J].西北农业学报,2007, 16(5):145-149.
    李树贤,吴志娟,赵萍.同源四倍体茄子自交亲和性遗传的初步分析[J].西北农业学报,2007,16 (6):170-173.
    李树贤,杨志刚,吴志娟,等.同源四倍体茄子育种的选择Ⅰ畸形僵果性状及植株结实力的选择[J].西北农业学报,2003,12(1)48- 52.
    李树贤.植物同源多倍体育种的几个问题[J].西北植物学报,2003,23(10): 1829-1841.
    黎垣庆,叶秀麟,陈泽濂,等.水稻无融合生殖材料PDER选育的初步研究[J].杂交水稻1995,4:6-8.
    马三梅,王永飞,赵男先,等.植物无融合生殖鉴定方法研究进展[J].西北植物学报,2002,22(4):985-993.
    谭素英,黄贞光,梁毅.同源四倍体西瓜自交花粉萌发生长与内源激素的关系[J].华北农学报,2000,15(4):73-79.
    王兰,刘向东,卢永根.同源四倍体水稻胚乳发育:极核融合和胚乳细胞化[J].中国水稻科学,2004, 18(4):281-289.
    王兰,刘向东,卢永根,冯九焕,徐雪宾,徐是雄.同源四倍体水稻胚乳发育:糊粉层细胞壁纤维素物质发育、胚乳淀粉积累及胼胝质套的形成[J].中国水稻科学,2004,18(6):28-32.
    王卓伟,余茂德,鲁成.桑树二倍体及人工诱导的同源四倍体遗传差异的AFLP分析[J].植物学通报,2002,19(2):194-200
    王志伟,李荣田,郭德栋.植物无融合生殖研究进展[J].中国生物工程杂志,2004,24(6):34-42.
    殷朝珍,王兆龙,葛才林.草地早熟禾无融合生殖及其育种利用研究进展[J].草原与草坪,2006(1):18-23.
    杨寅桂,庄勇,陈龙正,娄群峰,陈劲枫.蔬菜多倍体育种及其应用[J].江西农业大学学报,2006, 28(4):534-538.
    廖飞熊,李育农.变叶海棠无融合生殖胚胎发育的特征与结果[J].江西农业大学学报,1996,18 (3):287-191.
    王艳,许秋生,叶秀麟.大黍无融合生殖研究概况[J].热带亚热带植物学报,2003,11(1):83-86.
    杨继.植物多倍体基因组的形成与进化[J].植物分类学报,2001,39(4):357-371.
    Adams KL. (2007). Evolution of duplicate gene expression in polyploid and hybrid plants [J]. J Hered. 98:136–141.
    Adams KL, Cronn R, Percifield R, et al. (2003). Genes duplicated by Polyploid show unequal contributions to the transcriptome and organ-specific reciprocal silencing [J]. Proc. Natl. Acad. Sci. USA 100: 4649-4654.
    Adams KL, Percifield R, Wendel JF. (2004). Organ-specific silencing of duplicated genes in a newly synthesized cotton allotetraploid [J]. Genetics 168: 2217- 2226.
    Adams KL, Wendel JF. (2004). Exploring the genomic mysteries of polyploidy in cotton [J]. Biol. J. Linn. Soc. 82: 573-581.
    Adams KL, Wendel JF (2005) Polyploidy and genome evolution in plants [J]. Curr Opin Plant Biol 8:135-141
    Ahloowalia BS (1971) Frequency, origin, and survival of aneuploids in tetraploid ryegrass [J]. Genetica 42:129-138
    Ali HB, Lysak MA, Schubert I. (2004) Genomic in situ hybridization in plants with small genomes is feasible and elucidates the chromosomal parentage in interspecific Arabidopsis hybrids [J]. Genome 47(5):954-960.
    Arabidopsis Genome Initiative (2000) Analysis of the genome sequence of the flowering plant Arabidopsis thaliana [J]. Nature 408:796-815.
    Armstrong SJ, Jones GH (2003) Meiotic cytology and chromosome behaviour in wild-type Arabidopsis thaliana [J]. J Exp Bot 54:1-10.
    Armstrong SJ, Caryl AP, Jones GH, Franklin FC (2002) Asy1, a protein required for meiotic chromosome synapsis, localizes to axis-associated chromatin in Arabidopsis and Brassica [J]. J Cell Sci 115: 3645–3655.
    Armstrong SJ, Sanchez-Moran E, Franklin FC.(2009) Cytological analysis of Arabidopsis thaliana meiotic chromosomes [J]. Methods Mol Biol. 558:131-145.
    Bailey JA, Gu Z, Clark RA, et al. (2002). Recent segmental duplications in the human genome [J]. Science 297:1003–1007.
    Ballington JR, Galletta GJ. (1976). Potential fertility levels in four diploid Vaccinium species [J]. J Am Soc Hortic Sci 101: 507-509.
    Barton AB, Pekosz MR, Kurvathi RS, et al. (2008). Meiotic recombination at the ends of chromosomes in Saccharomyces cerevisiae [J]. Genetics 179:1221-1235.
    Barth, S., Melchinger, A.E., Devezi-Savula, B. et al. (2000) A high-throughput system for genome-wide measurement of genetic recombination in Arabidopsis thaliana based on transgenic markers. Funct. Integr. Genomics, 1: 200–206.
    Baumel A, Ainouche M, Kalender R, et al. (2002). Retrotransposons and genomic stability in populations of the young allopolyploid species Spartina anglica C. E. Hubbard (Poaceae) [J]. Mol. Biol. Evol. 19: 1218-1227.
    Beaulieu J, Jean M, Belzile F.(2009) The allotetraploid Arabidopsis thaliana–Arabidopsis lyrata subsp. petraea as an alternative model system for the study of polyploidy in plants [J]. Mol Genet Genomics 281:421-435.
    Belling J. (1925). The origin of chromosomal mutations in Uvularia [J]. J Genet 15: 245-266.
    Bernstein J. H. Bernstein, F.A. Hopf and R.E. Michod. (1988) Is meiotic recombination an adaptation for repairing DNA, producing genetic variation or both?. In: R.E. Michod and B.R. Levin, Editors, The Evolution of Sex, Sinauer, Sunderland, MA (1988), pp. 139–160.
    Birchler JA, Yao H, Chudalayandi S (2007) Biological consequences of dosage dependent gene regulatory systems [J]. Biochim Biophys Acta 1769:422–428.
    Bishop DK. (1994) RecA homologs Dmc1 and Rad51 interact to form multiple nuclear complexes prior to meiotic chromosome synapsis [J]. Cell 79:1081-1092.
    Blanc G, Barakat, A, Guyot R, et al. (2000). Extensive duplication and reshuffling in the Arabidopsis genome [J]. Plant Cell 12: 1093-1101.
    Blanc G, Hokamp K, Wolfe KH. (2003). A recent polyploidy superimposed on older large-scale duplications in the Arabidopsis genome [J]. Genome Research 13: 137–144.
    Blanc G, Wolfe KH. (2004a). Widespread paleopolyploidy in model plant species inferred from age distributions of duplicate genes [J]. Plant Cell 16: 1667-1678.
    Blanc G., Wolfe KH. (2004b). Function divergence of duplicated genes formed by polyploidy during Arabidopsis evolution [J]. Plant Cell 16: 1679-1691.
    Bousalem M, Arnau G, Hochu I, et al. (2006). Microsatellite segregation analysis and cytogenetic evidence for tetrasomic inheritance in the American yam Dioscorea trifida and a new basic chromosome number in the Dioscoreae [J]. Theor Appl Genet. 113:439-451.
    Bowers J, Chapman BA, Rong J, etal. ( 2003). Unravelling angiosperm genome evolution by phylogenetic analysis of chromosomal duplication events [J]. Nature 422:433-438.
    Boyd WJR, Sisodia NS, Larter EN (1970) A comparative study of the cytological and reproductive behaviour of wheat and triticale subjected to two temperature regimes [J]. Euphytica 19:490-497.
    Bozza CG, Pawlowski WP. (2008). The cytogenetics of homologous chromosome pairing in meiosis in plants [J]. Cytogenet Genome Res. 120:313-319.
    Brochmann C, Nilsson T, Gabrielsen TM. (1996). A classic example of postglacial allopolyploid speciation reexamined using RAPD markers and nucleotide sequences: Saxifraga osloensis (Saxifragaceae) [J]. Symb Bot Upsala 31: 75-89.
    Brandham PE. (1977). The meiotic behaviour of inversions in polyploid aloineae [J]. Chromosoma 62:69-84.
    Brandham PE, Johnson MAT. (1977).The meiotic behaviour of inversions in polyploid aloineae [J].Chromosoma 62:85-91
    Cai D, Chen J, Chen D, et al. (2007) The breeding of two polyploid rice lines with the characteristic of polyploid meiosis stability [J]. Sci China C Life Sci 50:356-366.
    Cai X, Xu S S. (2007). Meiosis-driven genome variation in plants [J]. Curr Genomics 8:151-161.
    Caryl AP, Armstrong SJ, Jones GH, Franklin FC (2000) A homologue of the yeast HOP1 gene is inactivated in the Arabidopsis meiotic mutant asy1 [J]. Chromosoma 109:62-71.
    Causier B, Castillo R, Zhou JL, et al. (2005). Evolution in action: Following function in duplicated floral homeotic genes [J]. Curr. Biol. 15: 1508- 1512.
    Chen Z J. (2007). Genetic and epigenetic mechanisms for gene expression and phenotypic variation in plant polyploids [J]. Annu Rev Plant Biol 58:377–406.
    Chen Q, Armstrong KC. (1994). Genomic in situ hybridization in Avena sativa [J]. Genome 37: 607-612.
    Chen ZJ, Comai L, Pikaard CS. (1998). Gene dosage and stochastic effects determine the severity and direction of uniparental ribosomal RNA gene silencing (nucleolar dominance) in Arabidopsis allopolyploids [J]. Proc Natl Acad Sci USA 95: 14891-14896.
    Chen ZJ, Pikaard CS. (1997). Epigenetic silencing of RNA polymeraseⅠtranscription: A role for DNA methylation and histon modification in nucleolar domince [J].Genes Dev. 11: 2124-2136.
    Chen ZJ, Pikaard CS. (1997). Transcriptional analysis of nucleolar dominance in polyploid plants: biased expression silencing of progenitor rRNA genes is developmentally regulated in Brassica [J]. Proc Natl Acad Sci USA 94: 3442-3447.
    Chen Q, Armstrong K. (1994).Genomic in situ hybridization in Avena sativa [J].Genome 37:607-612.
    Clough, S.J. and Bent, A.F. (1998) Floral dip: a simplified method for Agrobacterium-mediated transformation of Arabidopsis thaliana. Plant J. 16, 735–743.
    Coleman L. (1950). Nuclear conditions in normal stem tissue of Vicia faba [J]. Can J Res 28: 382-391.
    Comai L (2005) The advantages and disadvantages of being polyploid [J]. Nat Rev Genet 6:836-846
    Comai L, Madlung A, Josefsson C, Tyagi A.(2003) Do the different parental 'heteromes' cause genomic shock in newly formed allopolyploids [J]. Philos Trans R Soc Lond B Biol Sci. 358(1434):1149-1155.
    Comai L. (2000). Genetic and epigenetic interactions in allopolyploid plants [J]. Plant Mol Biol 43: 387-399.
    Comai L, Tyagi AP, Winter K, et al. (2000). Phenotypic instability and rapid gene silencing in newly formed Arabidopsis allotetraploids [J]. Plant Cell 12: 1551-1568.
    Comai L, Tyagi AP, Lysak MA. (2003). FISH analysis of meiosis in Arabidopsis allopolyploids [J]. Chromosome Res. 11:217–226.
    Cooke J, Nowak MA, Boerlijst M, et al. (1997). Evolutionary origins and maintenance of redundant gene expression during metazoan development [J]. Trends Genet 13: 360-364.
    Couteau F, Belzile F, Horlow C, et al.(1999) Random chromosome segregation without meiotic arrest in both male and female meiocytes of a dmc1 mutant of Arabidopsis [J]. Plant Cell 11:1623-1634
    Davis L, Smith GR. (2001). Meiotic recombination and chromosome segregation in Schizosac charomyces pombe [J]. Proc. Natl. Acad. Sci. USA 98:8395–402.
    Dawe RK. (1998). Meiotic chromosome organization and segregation in plants [J]. Annu Rev Plant Physiol49:371-395.
    De Haan A, Maceira N O, Lumaret R, et al. (1992). Production of 2n gametes in diploid subspecies of Dactylis glomerata L . Occurrence and frequency of 2n eggs [J]. Ann Bot 69: 345-350.
    Dilkes BP, Spielman M, Weizbauer R, et al. (2008) The maternally expressed WRKY transcription factor TTG2 controls lethality in interploidy crosses of Arabidopsis [J]. PLoS Biol 6:2707-2720.
    Ding DQ, Yamamoto A, Haraguchi T, et al. (2004). Dynamics of homologous chromosome pairing during meiotic prophase in fission yeast [J]. Dev Cell. 6:329-341.
    Drouaud J, Mercier R, Chelysheva L, et al.(2007)Sex-specific crossover distributions and variations in interference level along Arabidopsis thaliana chromosome 4 [J].PLoS Genet. 3(6):e106.
    Doebley J, Stec A, Hubbard L. (1997). The evolution of apical dominance in maize [J]. Nature 386: 485-488.
    Dorsey E. (1936) Induced polyploidy in wheat and rye. Chromosome doubling in Triticum , Secale and Triticum Secale hybrids produced by temperature changes[J]. J Hered 27: 155-160.
    Dover GA, Riley R. (1972). Prevention of Pairing of Homoeologous Meiotic Chromosomes of Wheat by an Activity of Supernumerary Chromosomes of Aegilops [J]. Nature 240, 159-161
    Doyle JJ, Flagel LE, Paterson AH,et al.(2008) Evolutionary genetics of genome merger and doubling in plants [J]. Annu Rev Genet. 42:443-461.
    Duesberg P (2007) Chromosomal chaos and cancer. Sci Am 296:53–59.
    Ermolaeva MD, Wu M, Eisen JA, et al. (2003). The age of the Arabidopsis thaliana genome duplication [J]. Plant Mol Biol 51:859-866.
    Feldman M, Liu B, Segal G, et al. (1997). Rapid elimination of low copy DNA sequences in polyploid wheat: a possible mechanismfor differentiation of homoeologous chromosomes [J]. Genetics 147: 1318-1387.
    Feldman M, Levy AA (2005) Allopolyploidy--a shaping force in the evolution of wheat genomes [J]. Cytogenet Genome Res 109:250-258.
    Frary A, Nesbitt TC, Grandillo S, et al. (2000). fw2.2: A quantitative trait locus key to the evolution of tomato fruit size [J]. Science 289: 85-88.
    Fras A, Maluszynska J (2004) The correlation between the chromosome variation in callus and genotype of explants of Arabidopsis thaliana [J]. Genetica 121:145-154.
    Gaut BS, Doebley JF (1997). DNA sequence evidence for the segmental allopolyploid origin of maize [J]. Proc. Natl. Acad. Sci. USA 94: 6808-6814.
    Gaeta RT, Pires JC, Iniguez-Luy F, et al. (2007). Genomic changes in resynthesized Brassica napus and their effect on gene expression and phenotype [J]. Plant Cell 19:3403-17.
    Galili G, Feldman M. (1984). Intergenomic suppression of endosperm protein genes in common wheat [J]. Can J Genet Cytol 26: 651-656.
    Galitski T, Saldanha AJ, Styles CA, et al. (1999). Ploidy regulation of gene expression [J]. Science 285:251–254.
    Gaut BS, Doebley JF. (1997). DNA sequence evidence for the segmental allotetraploid origin of maize [J].Proc Natl Acad Sci USA 94: 6808-6814.
    Gibson TJ, Spring J. (1998). Genetic redundancy in vertebrates: polyploidy and persistence of genes encoding multidomain proteins [J]. Trends Genet 14: 46-49.
    Gill BS. (1991). Nucleocytoplasmic interaction (NCT) hypothesis of genome evolution and speciation in polyploid plants [R]. Yokohama, Proceedings of the Kihara Memorial International Symposium on Cytoplasmic Engineering in Wheat.
    Gottlieb LD, Ford VS. (1997). A recently silenced duplicate PgiC locus in Clarkia [J]. Mol Biol Evol 14: 125-132.
    Goodman HM, Ecker JR, Dean C (1995) The genome of Arabidopsis thaliana [J]. Proc Natl Acad Sci USA 24: 10831-10835.
    Gorman P, Roylance R. (2006) Fluorescence in situ hybridization and comparative genomic hybridization [J]. Methods Mol Med. 120:269-295.
    Grandbastien MA, Spielmann A, Caboche M. (1989). Tntl, a mobile retroviral like transposable element of tobacco isolated by plant cell genetics [J]. Nature 337: 376-380.
    Grant V. (1952). Cytogenetics of the hybrid Gilia millefoliata×achilleaefolia.Ⅰ. Variations in meiosis and polyploidy rate as affected by nutritional and genetic conditions [J]. Chromosoma 5: 372-390.
    Grey C, Baudat F, de Massy B (2009) Genome-wide control of the distribution of meiotic recombination. PLoS Biol 7: 327–339.
    Griffiths S, Sharp R, Foote TN, et al. (2006). Molecular characterization of Ph1 as a major chromosome pairing locus in polyploid wheat [J]. Nature 439: 749-752.
    Guo M, Davis D, Birchler JA. (1996). Dosage effects on gene expression in a maize ploidy series [J]. Genetics 142:1349-1355.
    Guo M, Birchler JA (1994) Trans-acting dosage effects on the expression of model gene systems in maize aneuploids [J]. Science 266: 1999–2002.
    Gottlieb LD (2003) Plant polyploidy: gene expression and genetic redundancy [J]. Heredity 91:91-92.
    Hackett CA, Bradshaw JE, McNicol JW.(2001)Interval mapping of quantitative trait loci in autotetraploid species [J].Genetics 159(4):1819-32.
    Hagerup O. (1974) The spontaneous formation of haploid, polyploid and aneuploid embryos in some orchids [J]. Kongel Danshe Videnskab Selskab Biol Meddelelser 20:1-22.
    Haldane JSB. (1919) The combination of linkage values and the calculation of distances between the loci of linked factors [J]. J Genet 8:299-309.
    Han F, Gao Z, Yu W, et al. (2007). Minichromosome analysis of chromosome pairing, disjunction, and sister chromatid cohesion in maize [J]. Plant Cell.19:3853-3863.
    Hanson RE, Zhao XP, Islam-Faridi MN, et al. (1998). Evolution of interspersed repetitive elements in Gossypium (Malvaceae) [J]. Am J Bot 85: 1364-13687.
    Harberd NP, Flavell RB, Thompson RD. (1987). Identification of a transposon like insertion in a Glu-1 allele of wheat [J]. Mol Gen Genet 209: 326-332.
    Harland SC. (1936).The genetical conception of the species [J] biol reviews 11:83-112.
    Haufler CH. (1987).Electrophoresis is modifying our concepts of evolution in homosporous pteridophytes [J].Am J Bot 74:953-966.
    Havekes FW, Jong JH, Heyting C (1997) Comparative analysis of female and male meiosis in three meiotic mutants of tomato [J]. Genome 40:879-886.
    Hegarty M J, Hiscock S J. (2008). Genomic clues to the evolutionary success of polyploid plants [J]. Curr Biol.18:R435-44.
    Helentjaris T, Weber D, Wright S. (1998). Identification of the genomic locations of duplicate nucleotide sequences inmaize by analysis of restriction fragment length polymorphisms [J]. Genetics 118: 353-363.
    He P, Friebe BR, Gill BS, et al. (2003). Allopolyploidy alters gene expression in the highly stable hexaploid wheat [J]. Plant Mol. Biol. 52: 401-414.
    Henikoff S, Matzke MA. (1997). Exploring and explaining epigenetic effects [J]. Trends Genet.13: 293-295.
    Henry IM, Dilkes BP, Comai L (2007) Genetic basis for dosage sensitivity in Arabidopsis thaliana [J]. PLoS Genet 3:e70.
    Henry IM, Dilkes BP, Young K, Watson B, Wu H, Comai L (2005) Aneuploidy and genetic variation in the Arabidopsis thaliana triploid response [J]. Genetics 170:1979–1988.
    Higgins JD, Armstrong SJ, Franklin FC, et al.(2004). The Arabidopsis MutS homolog AtMSH4 functions at an early step in recombination: evidence for two classes of recombination in Arabidopsis [J]. Genes Dev.18:2557-2570.
    Holliday R, Ho T. (1998). Evidence for gene silencing by endogenous DNA methylation [J]. Proc Natl Acad Sci USA 95: 8727-8732.
    Hong D Y (1990). Cytotaxonomy of Plants. Beijing. Science Press.
    Hughes AL, (1994). The evolution of functionally novel proteins after gene duplication [J]. Proc R Soc Lond B 256: 119-124.
    Hughes MK, Hughes AL. (1993). Evolution of duplicate genes in a tetraploid animal Xenopus laevis [J]. Mol Biol Evol 10:1360-1369.
    Huettel B, Kreil DP, Matzke M, Matzke AJ (2008) Effects of aneuploidy on genome structure, expression, and interphase organization in Arabidopsis thaliana [J]. PLoS Genet 4(10):e1000226.
    Hollingsworth NM, Goetsch L, Byers B. (1990). The HOP1 gene encodes a meiosis-specific component of yeast chromosomes [J]. Cell 61:73-84.
    Houchins K, O’Dell M, Flavell RB, et al. (1997). Cytosine methylation and nucleolar dominance in cereal hybrids [J]. Mol. Gen. Genet. 255: 294-301.
    Hovav R, Udall JA, Chaudhary B, et al. (2008). Partitioned expression of duplicated genes during development and evolution of a single cell in a polyploid plant [J]. Proc Natl Acad Sci USA. 105:6191-6195.
    Jellen EN, Gill BS, Cox TS. (1994). Genomic in situ hybridization differentiates between A/D- and C-genome chromatin and detects intergenomic translocations in polyploid oat species (genus Avena)[J]. Genome 37: 613-618.
    Jenuwein T, Allis CD. (2001). Translating the histon code [J]. Science 293: 1074-1080.
    Jiang J, Gill BS (1994). Different species specific chromosome translocations in Triticum timopheevii and T. turgidum support the diphyletic origin of polyploid wheats [J]. Chromosome Res 2: 59-64.
    Jiang J M, Gill B S. (2006) Current status and the future of fluorescence in situ hybridization (FISH) in plant genome research [J]. Genome 49: 1057–1068
    Jones GH, Armstrong SJ, Caryl AP, et al. (2003). Meiotic chromosome synapsis and recombination in Arabidopsis thaliana; an integration of cytological and molecular approaches [J]. Chromosome Res. 11:205-215.
    Jones GH. (1978).Giemsa C-banding of rye meiotic chromosomes and the nature of terminal chiasmata [J].Chromosoma 66:45-57.
    Karpechenko GD. (1927). The production of polyploid gametes in hybrids [J]. Hereditas 9: 349-368.
    Kashkush K, Feldman M, Levy AA. (2002). Gene loss, silencing and activation in newly synthesized wheat allopolyploid [J]. Genetics 160: 1651-1659.
    Kashkush K, Feldman M, Levy AA. (2003). Transcriptional activation of retrotransposons alters the expression of adjacent genes in wheat [J]. Nat. Genet. 33: 102-106.
    Kellis M, Birren BW, Lander ES. (2004). Proof and evolutionary analysis of ancient genome duplication in the yeast Saccharomyces cereviseae [J]. Nature 428:617-624.
    Kenton A, Parokonny AS, Gleba YY,et al. (1993). Characterization of the Nicotiana tabacum L. genome by molecular cytogenetics [J]. Mol Gen Genet 240: 159-169.
    Klinga K (1986) Aneuploidy in induced autotetraploid populations of Festuca pratensis, Lolium multiflorum and Lolium perenne [J]. Hereditas 104:121-130.
    Lagercrantz U, Lydiate DJ. (1996). Comparative genome mapping in Brassica [J]. Genetics 144: 1903-1910
    Larhammer D, Risinger C. (1994). Why so few pseudogenes in tetraploid species [J] Trends Genet 10: 418-419.
    Lee HS, Chen ZJ. (2001). Protein-coding genes are epigenetically regulated in Arabidopsis polyploids [J]. Proc. Natl. Acad. Sci. USA 98: 6753-6758.
    Leitch AR, Leitch I J. (2008). Genomic plasticity and the diversity of polyploid plants [J]. Science. 320:481-483.
    Leitch I J, Bennett MD. (1997). Polyploidy in angiosperms [J]. Trends Plant Sci 2: 470-476.
    Lenormand T, Dutheil J (2005) Recombination difference between sexes: a role for haploid selection [J]. PLoS Biol 3:396–403.
    Lenormand T.(2003)The evolution of sex dimorphism in recombination [J].Genetics 163(2):811-22.
    Lerceteau-K?hler E, Guérin G, Laigret F, et al. (2003). Characterization of mixed disomic and polysomic inheritance in the octoploid strawberry (Fragaria x ananassa) using AFLP mapping [J]. Theor Appl Genet. 107:619-628.
    Levsky JM, Singer RH (2003). Fluorescence in situ hybridization: past, present and future [J]. J Cell Sci. 116(Pt14):2833-2838.
    Levy AA, Feldman M. (2004). Genetic and epigenetic reprogramming of the wheat genome upon allopolyploidization [J]. Biol J Linn Soc 82:607-613.
    Li WH. (1985). Accelerated evolution following gene duplication and its implications for the neutralist selectionist controversy. In: OhtaT, AokiK eds. Population Genetics and Molecular Evolution. Berlin: Spinger-Verlag. 333-353.
    Lister C, Jackson D, Martin C. (1993). Transposon2indued inversion in Antirrhinum modifies nivea gene expression to give a novel flower color pattern under the control of Cycloidea radialis [J]. Plant Cell 5: 1541-1553.
    Liu, B, Brubaker CL, Mergeai G, et al. (2001). Ployploid formation in cotton is not accompanied by rapid genomic changes [J]. Genome 44: 321-330.
    Liu B, Vega JM, Feldman M (1998a) Rapid genomic changes in newly synthesized amphiploids of Triticum and Aegilops .Ⅱ. Changes in low copy coding DNA sequence [J]. Genome 41: 535-542.
    Liu B, Vega JM, Segal G, et al. (1998b). Rapid genomic changes in newly synthesized amphiploids of Triticum and Aegilops.Ⅰ. Changes in low copy non-coding DAN sequences [J]. Genome 41: 272-277.
    Liu B, Wendel JF (2003). Epigenetic phenomena and the evolution of plant allopolyploids [J]. Mol Phylo genet Evol 29:365-379.
    Lorch PD.(2005) Sex differences in recombination and mapping adaptations [J].Genetica 123(1-2):39-47.
    Luig NH (1960) Differential transmission of gametes in wheat [J]. Nature 185:636– 637.
    Lundin LG. (1993). Evolution of the vertebrate genome as reflected in paralogous chromosome regions in man and the house mouse [J]. Genomics 16: 1–19.
    Lukens L N, Pires J C, Leon E, et al. (2006). Patterns of sequence loss and cytosine methylation within a population of newly resynthesized Brassica napus allopolyploids [J]. Plant Physiol 140:336-348.
    Lu X, Liu X, An L, et al. (2008). The Arabidopsis MutS homolog AtMSH5 is required for normal meiosis [J]. Cell Res. 18:589-599.
    Luo ZW,Zhang RM,Kearsy MJ.(2004)Theoretical basis for genetic linkage analysis in autotetraploid species [J]. Proc Natl Acad Sci USA 101(18):7040-7045.
    Maceira NO, De Haan AA, Lumaret R. (1992). Production of 2n gametes in diploid subspecies of Dactylis glomerata L. 1. Occurrence and frequency of 2n pollen [J]. Ann Bot 69: 335-343.
    Madlung A, Masuelli RW, Watson B, et al. (2002). Remolding of DNA methylation and phenotypic and transcriptional changes in synthetic Arabidopsis allotetraploids [J]. Plant Physiol. 129: 733- 746.
    Madlung A, Tyagi AP, Watson B, Jiang H, et al.(2005) Genomic changes in synthetic Arabidopsis polyploids [J]. Plant J 41(2):221-230.
    Makarevitch I, Phillips RL, Springer NM (2008) Profiling expression changes caused by a segmental aneuploid in maize [J]. BMC genomics 9:7
    Maestra BN, de Jong JH, Shepherd K, et al. (2002). Chromosome arrangement and behaviour of two rye homologous telosomes at the onset of meiosis in disomic wheat-5RL addition lines with and without the Ph1 locus [J]. Chromosome Res 10:655–667.
    Mancera E, Bourgon R, Brozzi A, et al. (2008). High-resolution mapping of meiotic crossovers andnon-crossovers in yeast [J]. Nature 454:479-485.
    Martinez-Perez E, Shaw P, Moore G. (2001). The Ph1 locus is needed to ensure specific somatic and meiotic centromere association [J]. Nature 411:204–207.
    Martienssen RA, Colot V. (2001). DNA methlation and epigenetic inheritance in plants and filamentous fungi [J]. Science 293: 1070-1074.
    Masterson J. (1994). Stomatal size in fossil plants: evidence for polyploidy in majority of angiosperms [J]. Science 264: 421-424.
    Matzke MA, Matzke AJM. (1998a). Gene silencing in plants: relevance for genome evolution and the acquisition of genomic methylation patterns. In: Epigenetics Novartis Foundation Symposium 214. Chichester: Wiley &Sons.168-186.
    Matzke MA, Matzke AM. (1998b). Epigenetic silencing of plant transgenes as a consequence of diverse cellular defense responses [J]. Cell Mol Life Sci 54: 94-103.
    Matzke MA, Mette MF, Kanno T, Matzke AJ (2003) Does the intrinsic instability of aneuploid genomes have a causal role in cancer [J]. Trends Genet 19:253–256.
    Meyer P, Saedler H. (1996). Homology-dependent gene silencing in plants [J]. Annu. Rev. Plant Physiol. Plant Mol. Biol. 47: 23-28.
    Michaels SD, Amasino RM. (1999). FLOWERING LOCUS C encodes a novel MADS-domain protein that acts as a repressor of flowering [J]. Plant Cell 11: 949-956.
    Mochida K, Yamazaki Y, Ogihara Y. (2003). Discrimination of homoeologous gene expression in hexaploid wheat by SNP analysis of contigs grouped from a large number of expressed sequence tags [J]. Mol. Genet. Genomics 270: 371-377.
    May BP, Dellaporta SL. (1998). Transposon sequences drive tissues pecific expression of the maize regulatory gene Rs [J]. Plant J 13: 241-247.
    McClintock B. (1984). The significance of responses of the genome to challenge [J]. Science 226: 792-801.
    McHale NA. (1983). Environmental induction of high frequency 2n pollen formation in diploid Solanum [J]. Can J Genet Cytol 25: 609-615.
    Mell JC, Wienholz BL, Salem A, et al. (2008). Sites of recombination are local determinants of meiotic homolog pairing in Saccharomyces cerevisiae. Genetics.179:773-784.
    Melamed-Bessudo C, Yehuda E, Stuitje AR, Levy AA.(2005)A new seed-based assay for meiotic recombination in Arabidopsis thaliana [J].Plant J 43(3):458-466.
    Mercier R, Jolivet S, Vezon D, et al. (2005). Two meiotic crossover classes cohabit in Arabidopsis: one is dependent on MER3, whereas the other one is not [J]. Curr Biol 15:692–701.
    Mikhailova EI, Phillips D, Sosnikhina SP, et al. (2006). Molecular assembly of meiotic proteins Asy1 and Zyp1 and pairing promiscuity in rye (Secale cereale L.) and its synaptic mutant sy10 [J]. Genetics 174:1247–1258.
    Ming R, Liu S C,. Moore PH, et al. (2001) QTL Analysis in a Complex Autopolyploid: Genetic Control of Sugar Content in Sugarcane. Genome Res. 11: 2075–2084.
    Mittelsten Scheid O, Afsar K, Paszkowski J. (2003) Formation of stable epialleles and their paramutation-like interaction in tetraploid Arabidopsis thaliana [J]. Nat. Genet. 34:450–454.
    Mittelsten Scheid O, Jakovleva L, Afsar K, et al. (1996) A change of ploidy can modify epigenetic silencing [J]. Proc Natl Acad Sci USA 93:7114–7119.
    Mok D WS, Peloquin SJ. (1975). Three mechanisms of 2n pollen formation in diploid potatoes [J]. Can J Genet Cytol 17: 217-225.
    Moens PB, Chen DJ, Shen Z,et al. (1997) Rad51 immunocytology in rat and mouse spermatocytes and oocytes [J]. Chromosoma 106:207-215.
    Moore G. (2002). Meiosis in allopolyploids– the importance of‘Teflon’chromosomes [J]. Trends in Genetics 18:456-463.
    Moore G, Devos KM, Wang Z. (1995). Grasses, line up and form a circle [J]. Curr Biol 5: 737-739.
    Murashige T, Skoog F (1962) A revised medium for rapid growth and bioassays with tobacco tissue cultures [J]. Physiol Plant 15: 473-497.
    Naranjo T, Corredor E.(2004).Clustering of centromeres precedes bivalent chromosome pairing of polyploid wheats.Trends Plant Sci 9(5):214-217.
    Neale MJ, Keeney S. (2006). Clarifying the mechanics of DNA strand exchange in meiotic recombination [J]. Nature 442:153–158.
    Newton WCF, Pellew C. (1929). Primula kewensis and its derivatives [J]. J Genet 20:405-467.
    Nonomura KI, Nakano M, Eiguchi M, et al. (2006). PAIR2 is essential for homologous chromosome synapsis in rice meiosis I [J]. J Cell Sci 119:217–225.
    Ohta T. (1991). Multigene families and the evolution of complexity [J]. J Mol Evol 33: 34-41.
    Ohta T. (1994). Further examples of evolution by gene duplication revealed through DNA sequence comparisons [J]. Genetics 138: 1331-1337.
    Osborn TC. (2004). The contribution of polyploid to variation in Brassica species [J]. Physiol. Plant.121: 531-536.
    Osborn TC, Pires JC, Birchler JA, et al. (2003). Understanding mechanisms of novel gene expression in polyploids [J]. Trends Genet. 19: 141-147.
    Ozkan H, Levy AA, Feldman M. (2001). Allopolyploidy-induced rapid genome evolution in the wheat (Aegilops-Triticum) group [J]. Plant Cell 13: 1735-1747.
    Ozkan H, Tuna M, Arumuganathan K. (2003). Nonadditive changes in genome size during allopolyploidization in the wheat (aegilops-triticum) group [J].J Hered 94:260-264.
    Papp I, Iglesias VA, Moscone EA, et al. (1996) Structural instability of a transgene locus in tobacco is associated with aneuploidy [J]. Plant J 10:469–478
    Parrott WA, Smith RR. (1984). Production of 2n pollen in red clover [J]. Crop Sci 24:469-472.
    Parry JM, Al-Obaidly A, Al-Walhaib M, et al. (2002) Spontaneous and induced aneuploidy, considerations which may influence chromosome malsegregation [J]. Mutat Res 504:119-129
    Pawlowski WP, Cande WZ. (2005). Coordinating the events of the meiotic prophase [J]. Trends Cell Biol. 15:674-681.
    Pickett FB, Meeks-Wagner DR. (1995). Seeing double: appreciating genetic redundancy [J]. Plant Cell, 7:1347-1356.
    Pignatta D, Dilkes B, Wroblewski T, et al. (2008). Transgene-induced gene silencing is not affected by a change in ploidy level [J]. PLoS ONE. 3:e3061.
    Pikaard CS. (2002). Genomic change and gene silencing in polyploids [J]. Trends Genet. 17, 657-677.
    Pires JC, Zhao J, Schranz ME, et al. (2004). Flowering time divergence and genomic rearrangements in resynthesized Brassica polyploids (Brassicaceae) [J]. Biol.J. Linn. Soc. 82, 675-688.
    Pontes O, Neves N, Silva M, et al.(2004). Chromosomal locus rearrangements are a rapid response to formation of the allotetraploid Arabidopsis suecica genome [J]. Proc Natl Acad Sci USA. 101:18240–18245.
    Ramsey J, Schemske DW. (1998). Pathways, mechanisms, and rates of polyploid formation in flowering plants [J]. Annu Rev Ecol Syst 29: 467-501.
    Randolph LF. (1932). Some effects of high temperature on polyploidy and other variations in maize [J]. Proc Natl Acad Sci USA 18: 222-229.
    Redei GP. (1969) A review of genetics and biology of Arabidopsis thaliana (L.) Heynh. Bibl. Genet 21:1-151.
    Reeder RH. (1985). Mechanisms of nucleolar dominance in animals and plants [J]. J Cell Biol 101: 2013-2016.
    Reinisch AJ, Dong J, Brubaker CL, et al. (1994). A detailed RFLP map of cotton, Gossypium hirsutum×G. barbadense: chromosome organization and evolution in a disomic polyploid genome [J]. Genetics 138: 829-847.
    Roberts MA, Reader SM, Dalgliesh C et al.(1999) Induction and characterization of Ph1 wheat mutants [J]. Genetics 153(4):1909-1918.
    Roelofs D, van Velzen J, Kuperus P, et al. (1997). Molecular evidence for an extinct parent of the tetraploid species Microseris acuminata and M. campestris (Asteraceae, Lactuceae) [J]. Mol Ecol 6: 641-649.
    Ross KJ, Fransz P, Jones GH (1996) A light microscopic atlas of meiosis in Arabidopsis thaliana [J]. Chromosome Res 4:507-516
    Ross KJ, Fransz P, Armstrong SJ, Vizir I, et al. (1997) Cytological characterization of four meiotic mutants of Arabidopsis isolated from T-DNA transformed lines [J]. Chromosome Res 5:551–559.
    Salina EA, Numerova OM, Ozkan H, etal. (2004). Alterations in subtelomeric tandem repeats during early stages of allopolyploidy in wheat [J]. Genome 47:860-867.
    Sanchez-Moran E, Osman K, Higgins JD, etal. (2008). ASY1 coordinates early events in the plant meiotic recombination pathway [J]. Cytogenet Genome Res. 120:302-312.
    Sanchez Moran E, Armstrong SJ, Santos JL, Franklin FC, Jones GH (2001) Chiasma formation in Arabidopsis thaliana accession Wassileskija and in two meiotic mutants [J]. Chromosome Res 9:121-128.
    Sanchez-Moran E, Santos JL, Jones GH, Franklin FC (2007) ASY1 mediates AtDMC1- dependent interhomolog recombination during meiosis in Arabidopsis [J]. Genes Dev 21: 2220-2233.
    Sang T, Crawford DJ, Stuessy TF. (1995). Documentation of reticulate evolution in peonies (Paeonia) usinginternal transcribed spacer sequences of nuclear ribosomal DNA: implications for bio-geography and concerted evolution [J]. Proc Natl Acad Sci USA 92: 6813-6817.
    Santos JL, Alfaro D, Sanchez-Moran E, et al. (2003) Partial diploidization of meiosis in autotetraploid Arabidopsis thaliana [J].Genetics 2165: 1533–1540.
    Scannell DR, Byrne KP, Gordon JL, et al. (2006). multiple rounds of speciation associated with reciprocal gene loss in polyploid yeasts [J]. Nature 440:341–345.
    Schranz ME, Osborn TC. (2009) Novel flowering time variation in the resynthesized polyploid Brassica napus [J].J Hered. 91(3):242-246
    Scheid OM, Jakovleva L, Afsar K, et al. (1996). A change in ploidy can modify epigenetic silencing [J]. Proc Natl Acad Sci USA 93: 7114-7119.
    Scott RJ, Spielman M, Bailey J, Dickinson HG (1998) Parent-of-origin effects on seed development in Arabidopsis thaliana [J]. Development 125:3329-3341.
    Shcherban AB, Badaeva ED, Amosova AV, et al.(2008). Genetic and epigenetic changes of rDNA in a synthetic allotetraploid, Aegilops sharonensis x Ae. umbellulata [J]. Genome 51:261-71.
    Shitsukawa N, Tahira C, Kassai K, et al. (2007). Genetic and epigenetic alteration among three homoeologous genes of a class E MADS box gene in hexaploid wheat [J]. Plant Cell 19:1723-1737.
    Shoemaker RC, Polzin K, Labate J, et al. (1996). Gemome duplication in soybean (Glycine subgenus soja) [J]. Genetics 144, 329-338.
    Seoighe C. (2003). Turning the clock back on ancient genome duplication [J]. Curr Opin Genet Dev 13:636-643.
    Soltis DE, Soltis PS. (1989). Polyploidy, breeding systems and genetic differentiation in homosporous pteridophytes. In: Soltis D E, Soltis P S eds. Isozymes in Plant Biology. Oregan: Dioscorides Press.
    Soltis DE, Soltis PS. (1993). Molecular data and the dynamic nature of polyploidy [J]. Crit Rev Plant Sci 12: 243-273.
    Soltis DE, Soltis PS. (1999). Polyploidy : recurrent formation and genome evolution [J]. Trends Ecol Evol 9: 348-352.
    Soltis DE, Bell CD, Kim S, et al. (2008). Origin and early evolution of angiosperms [J]. Ann N Y Acad Sci.1133:3-25.
    Song K, Lu P, Tang K, et al. (1995). Rapid genome change in synthetic polyploids of Brassica and its implications for polyploid evolution [J]. Proc Natl Acad Sci USA 92: 7719-7723.
    Stacey NJ, Kuromori T, Azumi Y, et al. (2006). Arabidopsis SPO11-2 functions with SPO11-1 in meiotic recombination [J]. Plant J.48:206-216.
    Stebbins GL. (1971). Chromosomal Evolution in Higher Plants [J]. London: Edward Arnold. Stephens SG. (1951). Possible significance of duplication in evolution [J]. Adv Genet 4: 247-265.
    Stewart CN Jr, Via LE. (1993)A rapid CTAB DNA isolation technique useful for RAPD fingerprinting and other PCR applications [J]. Biotechniques 14(5):748-50.
    Tate JA, Ni Z, Scheen AC, et al. (2006). Evolution and expression of homeologous loci in Tragopogon miscellus (Asteraceae), a recent and reciprocally formed allopolyploid [J]. Genetics 173:1599–1611.
    Tavoletti S, Mariani A, Veronesi F. (1991). Phenotypic recurrent selection for 2n pollen and 2n egg production in diploid alfalfa [J]. Euphytica 57: 97-102.
    Taylor JS, Peer VD, Meyer A (2001). Genome duplication, divergent resolution and speciation [J]. Trends Genet 17:299-301.
    Tian CG., Xiong YQ, Liu TY, et al. (2005). Evidence for an ancient whole-genome duplication event in rice and other cereals [J]. Chin. J. Genet. 32: 519-527.
    Thompson JD, Lumaret R. (1992). The evolutionary dynamics of polyploid plants: origins, establishment and persistence [J]. Trends Ecol Evol 7: 303-306.
    Uchino A, Tanaka K (1995) Occurrence of aneuploid progenies from an asynaptic amphidiploid of Scilla scilloides (lindley) druce II mechanism of production of the various aneuploid progenies. J Plant Res 108: 185-194.
    Vandepoele K, Simillion C, vande Peer Y. (2003). Evidence that rice and other cereals are ancient aneuploids [J]. Plant Cell 15: 2192–2202.
    Vigfusson E. (1970). On polyspermy in the sunflower [J]. Hereditas 64: 1-52.
    Vinkenoog R, Bushell C, Spielman M, et al. (2003) Genomic imprinting and endosperm development in flowering plants [J]. Mol Biotechnol 25:149-184
    Vizir IY,Korol AB.(1990) sex difference in recombination frequency in Arabidopsis [J]. Heredity 65:379-383.
    Vuylsteke M, van Eeuwijk F, Van Hummelen P, et al. (2005). Genetic analysis of variation in gene expression in Arabidopsis thaliana [J]. Genetics 171:1267–1275.
    Wagner A. (1998). The fate of duplicated genes: loss or new function [J]. Bioessays 20: 785-788.
    Wang JL, Tian L, Lee HS, et al. (2006)Genomewide nonadditive gene regulation in Arabidopsis allotetraploids [J]. Genetics 172: 507-517.
    Weiss H, Maluszynska J. (2000). Chromosomal rearrangement in autotetraploid plants of Arabidopsis thaliana [J]. Hereditas 133:255-261.
    Wells JL, Pryce DW, McFarlane RJ. (2006). Homologous chromosome pairing in Schizosaccharomyces pombe [J]. Yeast. 23:977-989.
    Wendel JF. (2000). Genome evolution in polyploids [J]. Plant Mol Biol 42: 225-249.
    Wendel JF, Schnabel A, Seelanan T. (1995). Bidirectional interlocus concerted evolution following allopolyploid speciation in cotton (Gossypium) [J]. Proc Natl Acad Sci USA: 280-284.
    Werner JE, Peloquin SJ. (1991). Occurrence and mechanisms of 2n egg formation in 2x potato [J]. Genome 34: 975-982.
    White SE, Habera LF, Wessler SR. (1994). Retrotransposons in the flanking regions of normal plant genes: a role for copia like elements in the evolution of gene structure and expression [J]. Proc Natl Acad Sci USA 91: 11792-11796.
    Wijeratne AJ, Ma H. (2007) Genetic analyses of meiotic recombination in Arabidopsis [J]. J Integr Plant Biol 49(8):1199-1207.
    Williams BR, Prabhu VR, Hunter KE, et al. (2008) Aneuploidy affects proliferation and spontaneousimmortalization in mammalian cells [J]. Science 322:703-709.
    Wolffe AP, Matzke, MA (1999). Epigenetics: Regulation through repression [J]. Science 286: 481-486.
    Wolfe KH. (2001) Yesterday's polyploids and the mystery of diploidization [J]. Nature Reviews Genetics 2:333-341.
    Wong S, Butler G, Wolfe KH. (2002). Gene order evolution and paleopolyploidy in hemiascomycete yeasts [J]. Proc Natl Acad Sci. 99: 9272–9277.
    Wu R, Ma CX, Casella G.(2004)A bivalent polyploid model for mapping quantitative trait loci in outcrossing tetraploids [J].Genetics 166(1):581-95.
    Wu SS, Wu R, Ma CX, et al. (2001). A multivalent pairing model of linkage analysis in autotetraploids [J]. Genetics 159:1339-1350.
    Zhang CH, Li XF, Shen SX, et al. (2008) Determination of n+1 gamete transmission rate of trisomics and location of gene controlling 2n gamete formation in Chinese cabbage (Brassica rapa) [J]. J Integr Plant Biol 51:29-34.
    Zhang D, Sang T. (1999). Physical mapping of ribosomal RNA genes in peonies (Paeonia; Paeoniaceae) by fluorescent in situ hybridization: implications for phylogeny and concerted evolution [J]. Am J Bot 86: 735-740.
    Zhang L, Liu D, Yan Z, et al.(2004). Rapid changes of microsatellite flanking sequence in the allopolyploidization of newly synthesized hexaploid wheat [J]. Sci China C Life Sci. 47:553-561.
    Zhang L, Liu D, Lan X, et al. (2008). A synthetic wheat with 56 chromosomes derived from Triticum turgidum and Aegilops tauschii [J]. J Appl Genet 49:41-44.
    Zhao XP, Si Y, Hanson RE, et al. (1998). Dispersed repetitive DNA has spread to new genomes since polyploid formation in cotton [J]. Genome Res 8: 479-492.
    Zwierzykowski Z, Tayyar R, Brunell, et al. (1998). Genome recombination in intergeneric hybrids between tetraploid Festuca pratensis and Lolium multiflorum [J]. J Hered 89: 324-328.

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