植物基因组减数分裂重组热点作图研究进展
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摘要
减数分裂过程中同源染色体配对并进行遗传物质的交换与重组,这是形成遗传多样性的来源和进化的原动力,也是育种的遗传基础。重组起始于程序性DNA双链断裂(DSB),其修复过程可导致染色体交换(CO)。DSB和CO在基因组中并非随机分布,存在重组的热点。植物重组热点分布及其产生机制研究较少,但对育种有很大价值。在植物减数分裂重组途径的不同阶段可对重组热点进行作图,一类方法是通过重组后代分析CO位置,另一类方法是对重组发生时的DNA断点序列进行直接检测。本文结合酵母与动物中的工作对这些技术方法的原理、其对植物重组热点研究的贡献及其局限性进行综述。新的技术路线可提高重组热点的作图效率,对认识和利用植物重组现象具有重要意义。
Homologous chromosome pairing,synapsis and genetic recombination are achieved in meiosis. Meiotic recombination is the main source of genetic variation in sexually reproducing organisms because the recombination speeds up adaptation,and lay the foundation for traditional plant breeding. The recombination is initiated by the programmed DNA double-stranded breaks( DSBs) that are processed to form chromosome crossovers( COs).DSBs and COs predominantly occur at discrete genomic loci called recombination hotspots. The knowledge about the distribution of recombination hotspots in plant genome as well as the features associated with these areas is invaluable to breeding,yet still largely unknown. Here,methodologies to profile hotspots at different steps of the meiotic recombination pathway,their contributions to understanding of hotspots and their limitations were reviewed. Novel and high-efficiency approaches will not only facilitate the mapping of plant recombination hotspots,but also provide insights about the formation and the control of these hotspots.
Homologous chromosome pairing,synapsis and genetic recombination are achieved in meiosis. Meiotic recombination is the main source of genetic variation in sexually reproducing organisms because the recombination speeds up adaptation,and lay the foundation for traditional plant breeding. The recombination is initiated by the programmed DNA double-stranded breaks( DSBs) that are processed to form chromosome crossovers( COs).DSBs and COs predominantly occur at discrete genomic loci called recombination hotspots. The knowledge about the distribution of recombination hotspots in plant genome as well as the features associated with these areas is invaluable to breeding,yet still largely unknown. Here,methodologies to profile hotspots at different steps of the meiotic recombination pathway,their contributions to understanding of hotspots and their limitations were reviewed. Novel and high-efficiency approaches will not only facilitate the mapping of plant recombination hotspots,but also provide insights about the formation and the control of these hotspots.
引文
[1]LONGHESE M P,BONETTI D,GUERINE I,et al.DNA double-strand breaks in meiosis:Checking their formation,processing and repair[J].DNA Repair,2009,8(9):1127-1138.
[2]MA H.A molecular portrait of Arabidopsis meiosis[C]//SOMERVILLE C R,MEYEROWITZ E M.The Arabidopsis Book.Rockville,MD:American Society of Plant Biologists,2006:1-39.
[3]OSMAN K,HIGGINS J D,SANCHEZ-MORAN E,et al.Pathways to meiotic recombination in Arabidopsis thaliana[J].New Phytologist,2011,190(3):523-544.
[4]LUO Q,LI Y,SHEN Y,et al.Ten years of gene discovery for meiotic event control in rice[J].J Genet Genomics,2014,41(3):125-137.
[5]GRELON M,VEZON D,GENDROT G,et al.At SPO11-1 is necessary for efficient meiotic recombination in plants[J].EMBOJ,2001,20(3):589-600.
[6]HARTUNG F,WURZ-WILDERSINN R,FUCHS J,et al.The catalytically active tyrosine residues of both SPO11-1 and SPO11-2 are required for meiotic double-strand break induction in Arabidopsis[J].Plant Cell,2007,19(10):3090-3099.
[7]STACEY N J,KUROMORI T,AZUMI Y,et al.Arabidopsis SPO11-2 functions with SPO11-1 in meiotic recombination[J].Plant J,2006,48(2):206-216.
[8]VRIELYNCK N,CHAMBON A,VEZON D,et al.A DNA topoisomerase VI-like complex initiates meiotic recombination[J].Science,2016,351(6276):939-943.
[9]KLIMYUK V I,JONES J D.At DMC1,the Arabidopsis homologue of the yeast DMC1 gene:characterization,transposoninduced allelic variation and meiosis-associated expression[J].Plant J,1997,11(1):1-14.
[10]DOUTRIAUX M P,COUTEAU F,BERGOUNIOUX C,et al.Isolation and characterisation of the RAD51 and DMC1 homologs from Arabidopsis thaliana[J].Mol Gen Genet,1998,257(3):283-291.
[11]COUTEAU F,BELZILE F,HORLOW C,et al.Random chromosome segregation without meiotic arrest in both male and female meiocytes of a dmc1 mutant of Arabidopsis[J].Plant Cell,1999,11(9):1623-1634.
[12]LI W,CHEN C,MARKMANN-MULISCH U,et al.The Arabidopsis AtRAD51 gene is dispensable for vegetative development but required for meiosis[J].Proc Natl Acad Sci U S A,2004,101(29):10596-10601.
[13]KURZBAUER M T,UANSCHOU C,CHEN D,et al.The recombinases DMC1 and RAD51 are functionally and spatially separated during meiosis in Arabidopsis[J].Plant Cell,2012,24(5):2058-2070.
[14]高玲,慕小倩,林煜,等.真核生物减数分裂重组热点的研究进展[J].遗传,2005(4):641-650.
[15]李亚非,程祝宽.植物减数分裂同源染色体重组的分子机理[J].中国科学:生命科学,2015(6):537-543.
[16]HIGGINS J D,PERRY R M,BARAKATE A,et al.Spatiotemporal asymmetry of the meiotic program underlies the predominantly distal distribution of meiotic crossovers in barley[J].Plant Cell,2012,24(10):4096-4109.
[17]EDLINGER B,SCHLOGELHOFER P.Have a break:determinants of meiotic DNA double strand break(DSB)formation and processing in plants[J].J Exp Bot,2011,62(5):1545-1563.
[18]SHILO S,MELAMED-BESSUDO C,DORONE Y,et al.DNA crossover motifs associated with epigenetic modifications delineate open chromatin regions in Arabidopsis[J].Plant Cell,2015,27(9):2427-2436.
[19]BORDE V,ROBINE N,LIN W,et al.Histone H3 lysine 4 trimethylation marks meiotic recombination initiation sites[J].EMBO J,2009,28(2):99-111.
[20]BAKER Z,SCHUMER M,HABA Y,et al.Repeated losses of PRDM9-directed recombination despite the conservation of PRDM9 across vertebrates[J].Elife,2017,6:e24133.
[21]郭晓强.PRDM9在哺乳动物基因重组热点中的作用[J].生物化学与生物物理进展,2010,37(9):929-931.
[22]MYERS S,BOWDEN R,TUMIAN A,et al.Drive against hotspot motifs in primates implicates the PRDM9 gene in meiotic recombination[J].Science,2010,327(5967):876-879.
[23]MYERS S,FREEMAN C,AUTON A,et al.A common sequence motif associated with recombination hot spots and genome instability in humans[J].Nat Genet,2008,40(9):1124-1129.
[24]HAYASHI K,YOSHIDA K,MATSUI Y.A histone H3 methyltransferase controls epigenetic events required for meiotic prophase[J].Nature,2005,438(7066):374-378.
[25]BAUDAT F,BUARD J,GREY C,et al.PRDM9 is a major determinant of meiotic recombination hotspots in humans and mice[J].Science,2010,327(5967):836-840.
[26]CHOI K,HENDERSON I R.Meiotic recombination hotspots-a comparative view[J].Plant J,2015,83(1):52-61.
[27]CHOI K,ZHAO X,KELLY K A,et al.Arabidopsis meiotic crossover hot spots overlap with H2A.Z nucleosomes at gene promoters[J].Nat Genet,2013,45(11):1327-1336.
[28]WIJNKER E,VELIKKAKAM J G,DING J,et al.The genomic landscape of meiotic crossovers and gene conversions in Arabidopsis thaliana[J].Elife,2013,2:e01426.
[29]LI X,LI L,YAN J.Dissecting meiotic recombination based on tetrad analysis by single-microspore sequencing in maize[J].Nat Commun,2015,6:6648.
[30]PERRELLA G,CONSIGLIO M F,AIESE-CIGLIANO R,et al.Histone hyperacetylation affects meiotic recombination and chromosome segregation in Arabidopsis[J].Plant J,2010,62(5):796-806.
[31]SIDHU,GAGANPREET K,FANG CELESTIA,et al.Recombination patterns in maize reveal limits to crossover homeostasis[J].Proc Natl Acad Sci U S A,2015,112(52):15982-15987.
[32]CHELYSHEVA L,VEZON D,CHAMBON A,et al.The Arabidopsis HEI10 is a new ZMM protein related to Zip3[J].PLoSGenet,2012,8(7):e1002799.
[33]SINGER T,FAN Y,CHANG H S,et al.A high-resolution map of Arabidopsis recombinant inbred lines by whole-genome exon array hybridization[J].PLoS Genet,2006,2(9):e144.
[34]KIANIAN P M A,WANG M,SIMONS K,et al.High-resolution crossover mapping reveals similarities and differences of male and female recombination in maize[J].Nat Commun,2018,9(1):2370.
[35]KIM S,PLAGNOL V,HU T T,et al.Recombination and linkage disequilibrium in Arabidopsis thaliana[J].Nat Genet,2007,39(9):1151-1155.
[36]ZHOU Q,MIAO H,LI S,et al.A sequencing-based linkage map of cucumber[J].Mol Plant,2015,8(6):961-963.
[37]QI J,CHEN Y,COPENHAVER G P,et al.Detection of genomic variations and DNA polymorphisms and impact on analysis of meiotic recombination and genetic mapping[J].Proc Natl Acad Sci U S A,2014,111(27):10007-10012.
[38]TERASAWA M,SHINOHARA A,HOTTA Y,et al.Localization of Rec A-like recombination proteins on chromosomes of the lily at various meiotic stages[J].Genes Dev,1995,9(8):925-934.
[39]FRANKLIN A E,MCELVER J,SUNJEVARIC I,et al.Three-dimensional microscopy of the Rad51 recombination protein during meiotic prophase[J].Plant Cell,1999,11(5):809-824.
[40]PAWLOWSKI W P,GOLUBOVSKAYA I N,CANDE W Z.Altered nuclear distribution of recombination protein RAD51 in maize mutants suggests the involvement of RAD51 in meiotic homology recognition[J].Plant Cell,2003,15(8):1807-1816.
[41]CHELYSHEVA L,GRANDONT L,VRIELYNCK N,et al.An easy protocol for studying chromatin and recombination protein dynamics during Arabidopsis thaliana meiosis:immunodetection of cohesins,histones and MLH1[J].Cytogenet Genome Res,2010,129(1-3):143-153.
[42]HIGGINS J D,SANCHEZ-MORAN E,ARMSTRONG S J,et al.The Arabidopsis synaptonemal complex protein ZYP1 is required for chromosome synapsis and normal fidelity of crossing over[J].Genes Dev,2005,19(20):2488-2500.
[43]SANCHEZ-MORAN E,SANTOS J L,JONES G H,et al.ASY1 mediates At DMC1-dependent interhomolog recombination during meiosis in Arabidopsis[J].Genes Dev,2007,21(17):2220-2233.
[44]HE Y,SIDHU G,PAWLOWSKI W P.Chromatin immunoprecipitation for studying chromosomal localization of meiotic proteins in maize[J].Methods Mol Biol,2013,990:191-201.
[45]BAUDAT F,NICOLAS A.Clustering of meiotic double-strand breaks on yeast chromosome III[J].Proc Natl Acad Sci U S A,1997,94(10):5213-5218.
[46]ZENVIRTH D,ARBEL T,SHERMAN A,et al.Multiple sites for double-strand breaks in whole meiotic chromosomes of Saccharomyces cerevisiae[J].EMBO J,1992,11(9):3441-3447.
[47]GERTON J L,DERISI J,SHROFF R,et al.Global mapping of meiotic recombination hotspots and coldspots in the yeast Saccharomyces cerevisiae[J].Proc Natl Acad Sci U S A,2000,97(21):11383-11390.
[48]BLITZBLAU H G,BELL G W,RODRIGUEZ J,et al.Mapping of meiotic single-stranded DNA reveals double-stranded-break hotspots near centromeres and telomeres[J].Curr Biol,2007,17(23):2003-2012.
[49]BUHLER C,BORDE V,LICHTEN M.Mapping meiotic single-strand DNA reveals a new landscape of DNA double-strand breaks in Saccharomyces cerevisiae[J].PLoS Biol,2007,5(12):e324.
[50]PAN J,SASAKI M,KNIEWEL R,et al.A hierarchical combination of factors shapes the genome-wide topography of yeast meiotic recombination initiation[J].Cell,2011,144(5):719-731.
[51]SMAGULOVA F,GREGORETTI I V,BRICK K,et al.Genome-wide analysis reveals novel molecular features of mouse recombination hotspots[J].Nature,2011,472(7343):375-378.
[52]PRATTO F,BRICK K,KHIL P,et al.Recombination initiation maps of individual human genomes[J].Science,2014,346(6211):1256442.
[53]HE Y,WANG M,DUKOWIC-SCHULZE S,et al.Genomic features shaping the landscape of meiotic double-strand-break hotspots in maize[J].Proc Natl Acad Sci U S A,2017,114(46):12231-12236.
[54]SERRENTINO M E,CHAPLAIS E,SOMMERMEYER V,et al.Differential association of the conserved SUMO ligase Zip3with meiotic double-strand break sites reveals regional variations in the outcome of meiotic recombination[J].PLoS Genet,2013,9(4):e1003416.
[55]MELAMED-BESSUDO C,SHILO S,LEVY A A.Meiotic recombination and genome evolution in plants[J].Curr Opin Plant Biol,2016,30:82-87.
[2]MA H.A molecular portrait of Arabidopsis meiosis[C]//SOMERVILLE C R,MEYEROWITZ E M.The Arabidopsis Book.Rockville,MD:American Society of Plant Biologists,2006:1-39.
[3]OSMAN K,HIGGINS J D,SANCHEZ-MORAN E,et al.Pathways to meiotic recombination in Arabidopsis thaliana[J].New Phytologist,2011,190(3):523-544.
[4]LUO Q,LI Y,SHEN Y,et al.Ten years of gene discovery for meiotic event control in rice[J].J Genet Genomics,2014,41(3):125-137.
[5]GRELON M,VEZON D,GENDROT G,et al.At SPO11-1 is necessary for efficient meiotic recombination in plants[J].EMBOJ,2001,20(3):589-600.
[6]HARTUNG F,WURZ-WILDERSINN R,FUCHS J,et al.The catalytically active tyrosine residues of both SPO11-1 and SPO11-2 are required for meiotic double-strand break induction in Arabidopsis[J].Plant Cell,2007,19(10):3090-3099.
[7]STACEY N J,KUROMORI T,AZUMI Y,et al.Arabidopsis SPO11-2 functions with SPO11-1 in meiotic recombination[J].Plant J,2006,48(2):206-216.
[8]VRIELYNCK N,CHAMBON A,VEZON D,et al.A DNA topoisomerase VI-like complex initiates meiotic recombination[J].Science,2016,351(6276):939-943.
[9]KLIMYUK V I,JONES J D.At DMC1,the Arabidopsis homologue of the yeast DMC1 gene:characterization,transposoninduced allelic variation and meiosis-associated expression[J].Plant J,1997,11(1):1-14.
[10]DOUTRIAUX M P,COUTEAU F,BERGOUNIOUX C,et al.Isolation and characterisation of the RAD51 and DMC1 homologs from Arabidopsis thaliana[J].Mol Gen Genet,1998,257(3):283-291.
[11]COUTEAU F,BELZILE F,HORLOW C,et al.Random chromosome segregation without meiotic arrest in both male and female meiocytes of a dmc1 mutant of Arabidopsis[J].Plant Cell,1999,11(9):1623-1634.
[12]LI W,CHEN C,MARKMANN-MULISCH U,et al.The Arabidopsis AtRAD51 gene is dispensable for vegetative development but required for meiosis[J].Proc Natl Acad Sci U S A,2004,101(29):10596-10601.
[13]KURZBAUER M T,UANSCHOU C,CHEN D,et al.The recombinases DMC1 and RAD51 are functionally and spatially separated during meiosis in Arabidopsis[J].Plant Cell,2012,24(5):2058-2070.
[14]高玲,慕小倩,林煜,等.真核生物减数分裂重组热点的研究进展[J].遗传,2005(4):641-650.
[15]李亚非,程祝宽.植物减数分裂同源染色体重组的分子机理[J].中国科学:生命科学,2015(6):537-543.
[16]HIGGINS J D,PERRY R M,BARAKATE A,et al.Spatiotemporal asymmetry of the meiotic program underlies the predominantly distal distribution of meiotic crossovers in barley[J].Plant Cell,2012,24(10):4096-4109.
[17]EDLINGER B,SCHLOGELHOFER P.Have a break:determinants of meiotic DNA double strand break(DSB)formation and processing in plants[J].J Exp Bot,2011,62(5):1545-1563.
[18]SHILO S,MELAMED-BESSUDO C,DORONE Y,et al.DNA crossover motifs associated with epigenetic modifications delineate open chromatin regions in Arabidopsis[J].Plant Cell,2015,27(9):2427-2436.
[19]BORDE V,ROBINE N,LIN W,et al.Histone H3 lysine 4 trimethylation marks meiotic recombination initiation sites[J].EMBO J,2009,28(2):99-111.
[20]BAKER Z,SCHUMER M,HABA Y,et al.Repeated losses of PRDM9-directed recombination despite the conservation of PRDM9 across vertebrates[J].Elife,2017,6:e24133.
[21]郭晓强.PRDM9在哺乳动物基因重组热点中的作用[J].生物化学与生物物理进展,2010,37(9):929-931.
[22]MYERS S,BOWDEN R,TUMIAN A,et al.Drive against hotspot motifs in primates implicates the PRDM9 gene in meiotic recombination[J].Science,2010,327(5967):876-879.
[23]MYERS S,FREEMAN C,AUTON A,et al.A common sequence motif associated with recombination hot spots and genome instability in humans[J].Nat Genet,2008,40(9):1124-1129.
[24]HAYASHI K,YOSHIDA K,MATSUI Y.A histone H3 methyltransferase controls epigenetic events required for meiotic prophase[J].Nature,2005,438(7066):374-378.
[25]BAUDAT F,BUARD J,GREY C,et al.PRDM9 is a major determinant of meiotic recombination hotspots in humans and mice[J].Science,2010,327(5967):836-840.
[26]CHOI K,HENDERSON I R.Meiotic recombination hotspots-a comparative view[J].Plant J,2015,83(1):52-61.
[27]CHOI K,ZHAO X,KELLY K A,et al.Arabidopsis meiotic crossover hot spots overlap with H2A.Z nucleosomes at gene promoters[J].Nat Genet,2013,45(11):1327-1336.
[28]WIJNKER E,VELIKKAKAM J G,DING J,et al.The genomic landscape of meiotic crossovers and gene conversions in Arabidopsis thaliana[J].Elife,2013,2:e01426.
[29]LI X,LI L,YAN J.Dissecting meiotic recombination based on tetrad analysis by single-microspore sequencing in maize[J].Nat Commun,2015,6:6648.
[30]PERRELLA G,CONSIGLIO M F,AIESE-CIGLIANO R,et al.Histone hyperacetylation affects meiotic recombination and chromosome segregation in Arabidopsis[J].Plant J,2010,62(5):796-806.
[31]SIDHU,GAGANPREET K,FANG CELESTIA,et al.Recombination patterns in maize reveal limits to crossover homeostasis[J].Proc Natl Acad Sci U S A,2015,112(52):15982-15987.
[32]CHELYSHEVA L,VEZON D,CHAMBON A,et al.The Arabidopsis HEI10 is a new ZMM protein related to Zip3[J].PLoSGenet,2012,8(7):e1002799.
[33]SINGER T,FAN Y,CHANG H S,et al.A high-resolution map of Arabidopsis recombinant inbred lines by whole-genome exon array hybridization[J].PLoS Genet,2006,2(9):e144.
[34]KIANIAN P M A,WANG M,SIMONS K,et al.High-resolution crossover mapping reveals similarities and differences of male and female recombination in maize[J].Nat Commun,2018,9(1):2370.
[35]KIM S,PLAGNOL V,HU T T,et al.Recombination and linkage disequilibrium in Arabidopsis thaliana[J].Nat Genet,2007,39(9):1151-1155.
[36]ZHOU Q,MIAO H,LI S,et al.A sequencing-based linkage map of cucumber[J].Mol Plant,2015,8(6):961-963.
[37]QI J,CHEN Y,COPENHAVER G P,et al.Detection of genomic variations and DNA polymorphisms and impact on analysis of meiotic recombination and genetic mapping[J].Proc Natl Acad Sci U S A,2014,111(27):10007-10012.
[38]TERASAWA M,SHINOHARA A,HOTTA Y,et al.Localization of Rec A-like recombination proteins on chromosomes of the lily at various meiotic stages[J].Genes Dev,1995,9(8):925-934.
[39]FRANKLIN A E,MCELVER J,SUNJEVARIC I,et al.Three-dimensional microscopy of the Rad51 recombination protein during meiotic prophase[J].Plant Cell,1999,11(5):809-824.
[40]PAWLOWSKI W P,GOLUBOVSKAYA I N,CANDE W Z.Altered nuclear distribution of recombination protein RAD51 in maize mutants suggests the involvement of RAD51 in meiotic homology recognition[J].Plant Cell,2003,15(8):1807-1816.
[41]CHELYSHEVA L,GRANDONT L,VRIELYNCK N,et al.An easy protocol for studying chromatin and recombination protein dynamics during Arabidopsis thaliana meiosis:immunodetection of cohesins,histones and MLH1[J].Cytogenet Genome Res,2010,129(1-3):143-153.
[42]HIGGINS J D,SANCHEZ-MORAN E,ARMSTRONG S J,et al.The Arabidopsis synaptonemal complex protein ZYP1 is required for chromosome synapsis and normal fidelity of crossing over[J].Genes Dev,2005,19(20):2488-2500.
[43]SANCHEZ-MORAN E,SANTOS J L,JONES G H,et al.ASY1 mediates At DMC1-dependent interhomolog recombination during meiosis in Arabidopsis[J].Genes Dev,2007,21(17):2220-2233.
[44]HE Y,SIDHU G,PAWLOWSKI W P.Chromatin immunoprecipitation for studying chromosomal localization of meiotic proteins in maize[J].Methods Mol Biol,2013,990:191-201.
[45]BAUDAT F,NICOLAS A.Clustering of meiotic double-strand breaks on yeast chromosome III[J].Proc Natl Acad Sci U S A,1997,94(10):5213-5218.
[46]ZENVIRTH D,ARBEL T,SHERMAN A,et al.Multiple sites for double-strand breaks in whole meiotic chromosomes of Saccharomyces cerevisiae[J].EMBO J,1992,11(9):3441-3447.
[47]GERTON J L,DERISI J,SHROFF R,et al.Global mapping of meiotic recombination hotspots and coldspots in the yeast Saccharomyces cerevisiae[J].Proc Natl Acad Sci U S A,2000,97(21):11383-11390.
[48]BLITZBLAU H G,BELL G W,RODRIGUEZ J,et al.Mapping of meiotic single-stranded DNA reveals double-stranded-break hotspots near centromeres and telomeres[J].Curr Biol,2007,17(23):2003-2012.
[49]BUHLER C,BORDE V,LICHTEN M.Mapping meiotic single-strand DNA reveals a new landscape of DNA double-strand breaks in Saccharomyces cerevisiae[J].PLoS Biol,2007,5(12):e324.
[50]PAN J,SASAKI M,KNIEWEL R,et al.A hierarchical combination of factors shapes the genome-wide topography of yeast meiotic recombination initiation[J].Cell,2011,144(5):719-731.
[51]SMAGULOVA F,GREGORETTI I V,BRICK K,et al.Genome-wide analysis reveals novel molecular features of mouse recombination hotspots[J].Nature,2011,472(7343):375-378.
[52]PRATTO F,BRICK K,KHIL P,et al.Recombination initiation maps of individual human genomes[J].Science,2014,346(6211):1256442.
[53]HE Y,WANG M,DUKOWIC-SCHULZE S,et al.Genomic features shaping the landscape of meiotic double-strand-break hotspots in maize[J].Proc Natl Acad Sci U S A,2017,114(46):12231-12236.
[54]SERRENTINO M E,CHAPLAIS E,SOMMERMEYER V,et al.Differential association of the conserved SUMO ligase Zip3with meiotic double-strand break sites reveals regional variations in the outcome of meiotic recombination[J].PLoS Genet,2013,9(4):e1003416.
[55]MELAMED-BESSUDO C,SHILO S,LEVY A A.Meiotic recombination and genome evolution in plants[J].Curr Opin Plant Biol,2016,30:82-87.