利用回交和标记辅助选择快速培育高油酸花生品种及其评价
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摘要
【目的】高油酸育种是花生品质改良的重要方向,利用回交育种结合标记选择可快速实现现有推广品种的高油酸化改良,探讨利用这一技术体系进行花生高油酸遗传改良的实践和效率。【方法】以目前推广的优质高产抗病品种中花16、中花21、泉花551、徐花13为轮回亲本(母本,基因型AABB),以高油酸材料冀花13为非轮回亲本(父本,基因型aabb)配制4个杂交组合,一年种植两季并进行人工杂交或自交,夏季在武汉种植,冬季在湛江南繁基地种植,通过1次杂交、4次回交和1次自交得到BC4F2后代。利用PCR产物测序方法,鉴定杂交和回交后代的基因型:根据回交后代基因型分离规律及ah FAD2A与ah FAD2B序列高度同源性的特点,用引物F0.7/R3在一个PCR反应内同时高效扩增F1和回交后代(BC1F1-BC4F1)的ah FAD2A和ah FAD2B片段,并利用R3作为测序引物进行反向测序,读取测序峰图判别基因型,在回交后代中筛选基因型Aa Bb的后代作为下代回交父本。自交后代(BC4F2-BC4F3)基因型鉴定采用KASP分型,获得高油酸(基因型aabb)后代。对获得的基因型为aabb的高油酸后代与其对应轮回亲本进行重要农艺性状、品质和重要抗病性的调查和SSR标记检测。【结果】在3年时间内,4个组合分别获得10、5、6、8株BC4F2高油酸纯合隐性基因型(aabb)单株,通过一代自交获得相应的BC4F3株系,对获得的高油酸株系与轮回亲本进行植物学、农艺性状、品质和青枯病抗性的考察,最终,4个组合均获得了与轮回亲本综合性状最接近的株系,分别为ZJ019、ZJ109、ZJ160和ZJ805,其油酸含量为82.54%、79.85%、79.22%、和78.94%,可作为轮回亲本对应的高油酸新品种。另外,本研究还对中花16回交组合中获得的高油酸株系的遗传背景进行了SSR分子检测,发现ZJ019株系的回复率达94.8%,在该组合中回复率最高,这一结果与植物学、农艺性状鉴定的结果一致。【结论】利用连续回交、南繁加代和分子标记辅助选择等技术可在3年内快速实现现有推广花生品种的高油酸化改良。
【Objective】 The objectives of this research were(i) to establish an effective backcrossing approach combined with marker-assisted selection(MAS) of peanut high oleate breeding,(ii) to transform the elite peanut cultivars with normal oleate(NO) into high oleate(HO) ones in three years.【Method】Four cross combinations were designed named as BO-01, BO-02, BO-03 and BO-04 by selecting the elite peanut cultivars ZH16, ZH21, QH551 and XH13 with AABB genotype as recurrent parents, while JH13 with aabb genotype was selected as the high oleate donor parent. Via 1 cross, 4 backcrosses and 1 self-crossing, BC4 F2 progenies were obtained. According to the separation rule of backcross and high homology sequence of ah FAD2 A and ah FAD2 B, F1 and backcross progenies(BC1 F1-BC4 F1) were genotyped by sequencing of PCR products, in which ah FAD2 A and ah FAD2 B were amplified in one PCR reaction with F0.7/R3 primer pairs. Aa Bb genotype progenies were selected and used as male parent in next generation backcross. Self-crossing progenies(BC4 F2-BC4 F3) were examined by KASP(kompetitive allele specific PCR) to obtain aabb genotype progenies. Main agronomic traits, quality characters and disease resistance of aabb genotype progenies were investigated. The genetic backgrounds of progenies with aabb genotype derived from ZH 16×JH13 were evaluated with SSR markers. 【Result】In summary, during 3 years, we obtained 10, 5, 6 and 8 high oleate progenies with aabb genotype in 4 cross combinations, respectively. By evaluating their agronomic traits, quality and bacterial wilt resistance, high oleate peanut cultivars were obtained, which were highly consistent with their corresponding recurrent female parent. ZJ019, ZJ109, ZJ160 and ZJ805 with oleic content as 82.54%, 79.85%, 79.22% and 78.94%, are regarded as high oleic cultivars of ZH16, ZH21, QH551 and XH13.【Conclusion】By using backcross, off-season multiplication for advancing generation and molecular marker selection, normal oleate peanut cultivars were successfully and quickly transited into high oleate ones within 3 years.
【Objective】 The objectives of this research were(i) to establish an effective backcrossing approach combined with marker-assisted selection(MAS) of peanut high oleate breeding,(ii) to transform the elite peanut cultivars with normal oleate(NO) into high oleate(HO) ones in three years.【Method】Four cross combinations were designed named as BO-01, BO-02, BO-03 and BO-04 by selecting the elite peanut cultivars ZH16, ZH21, QH551 and XH13 with AABB genotype as recurrent parents, while JH13 with aabb genotype was selected as the high oleate donor parent. Via 1 cross, 4 backcrosses and 1 self-crossing, BC4 F2 progenies were obtained. According to the separation rule of backcross and high homology sequence of ah FAD2 A and ah FAD2 B, F1 and backcross progenies(BC1 F1-BC4 F1) were genotyped by sequencing of PCR products, in which ah FAD2 A and ah FAD2 B were amplified in one PCR reaction with F0.7/R3 primer pairs. Aa Bb genotype progenies were selected and used as male parent in next generation backcross. Self-crossing progenies(BC4 F2-BC4 F3) were examined by KASP(kompetitive allele specific PCR) to obtain aabb genotype progenies. Main agronomic traits, quality characters and disease resistance of aabb genotype progenies were investigated. The genetic backgrounds of progenies with aabb genotype derived from ZH 16×JH13 were evaluated with SSR markers. 【Result】In summary, during 3 years, we obtained 10, 5, 6 and 8 high oleate progenies with aabb genotype in 4 cross combinations, respectively. By evaluating their agronomic traits, quality and bacterial wilt resistance, high oleate peanut cultivars were obtained, which were highly consistent with their corresponding recurrent female parent. ZJ019, ZJ109, ZJ160 and ZJ805 with oleic content as 82.54%, 79.85%, 79.22% and 78.94%, are regarded as high oleic cultivars of ZH16, ZH21, QH551 and XH13.【Conclusion】By using backcross, off-season multiplication for advancing generation and molecular marker selection, normal oleate peanut cultivars were successfully and quickly transited into high oleate ones within 3 years.
引文
[1]http://www.fao.org/faostat/en/
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[3]JANILA P,PANDEY M K,SHASIDHAR Y,VARIATH M T,SRISWATHI M,KHERA P,MANOHAR S S,NAGESH P,VISHWAKARMA M K,MISHRA G P,RADHAKRISHNAN T,MANIVANNAN N,DOBARIYA K L,VASANTHI R P,VARSHNEY R K.Molecular breeding for introgression of fatty acid desaturase mutant alleles(ah FAD2A and ah FAD2B)enhances oil quality in high and low oil containing peanut genotypes.Plant Science an International Journal of Experimental Plant Biology,2016,242:203-213.
[4]BRADDOC J C,SIMS C A,O'KEEFE S F.Flavor and oxidative stability of roasted high oleic acid peanuts.Journal Food Science,1995,60:489-493.
[5]MUGENDI J B,SIMS C A,GORGET D W,O'KEEFE S F.Flavor stability of high-oleic peanuts stored at low humidity.Journal of the American Oil Chemists Society,1998,75:21-25.
[6]O’BRYNE D J,KNAUFT D A,SHIREMAN R B.Low fatmonounsaturated rich diets containing high-oleic peanuts improve serum lipoprotein profiles.Lipids 1997,32:687-695.
[7]赵术珍,侯蕾,李长生,赵传志,任丽,李爱芹,邓丽,夏晗,王兴军.分子标记辅助回交选育高油酸花生新种质.中国油料作物学报,2017,39(1):30-36.ZHAO S Z,HOU L,LI C S,ZHAO C Z,REN L,LI A Q,DENG L,XIA H,WANG X J.Development of high oleic acid peanut from molecular marker assisted selection.Chinese Journal of Oil Crop Sciences,2017,39(1):30-36.(in Chinese)
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[9]NORDEN A J,GORBET D W,KNAUFT D A,YOUNG C T.Variability in oil quality among peanut genotypes in the Florida breeding program.Peanut Science,1987,14:7-11.
[10]OKULEY J,LIGHTNER J,FELDMANN K,YADAV N,LARK E,BROWSE J.Arabidopsis FAD2 gene encodes the enzyme that is essential for polyunsaturated lipid synthesis.The Plant Cell,1994,6:147-158.
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[12]WANG C T,HU D Q,DING F Y,YU H T,TANG Y Y,WANG X Z,ZHANG J C,CHEN D X.A new set of allele-specific PCR primers for identification of true hybrids in normal oleate×high oleate crosses in groundnut.Journal of SAT Agricultural Research,2011,12(9):1-4.
[13]CHEN Z,WANG M L,BARKLEY N A,PITTMAN R N.A simple allele-specific PCR assay for detecting FAD2 alleles in both A and B genomes of the cultivated peanut for high-oleate trait selection.Plant Molecular Biology Reporter,2010,28:542-548.
[14]BARKLEY N A,CHAMBERLIN K D C,WANG M L,PITTMAN R N.Development of a real-time PCR genotyping assay to identify high oleic acid peanuts(Arachis hypogaea L.).Molecular Breeding,2010,25:541-548.
[15]CHU Y,RAMOS L,HOLBROOK C C,OZIAS-AKINS P.Frequency of a Loss-of-Function Mutation in Oleoyl-PC Desaturase(ah FAD2A)in the Mini-Core of the U.S.Peanut Germplasm Collection.Crop Science.2007,47(6):2372-2378.
[16]曹永强,孙石.回交在转基因作物育种中的应用.作物杂志,2014(1):9-14.CAO Y Q,SUN S.Backcross breeding genetically modified crops in the application.Crops,2014(1):9-14.(in Chinese)
[17]王传堂,王秀贞,唐月异,吴琪,孙全喜,张建成,崔凤高,李利民,苗昊翠.中国高油酸花生种质创制、品种选育进展与建议.花生学报,2015(2):49-53.WANG C T,WANG X Z,TANG Y Y,WU Q,SUN Q X,ZHANG J C,CUI F G,LI L M,MIAO H C.High-oleic peanut germplasm enhancement and cultivar releases in China:Main achievements and suggestions.Journal of Peanut Science,2015(2):49-53.(in Chinese)
[18]成良强.花生遗传图谱构建及产量相关性状的QTL分析[D].北京:中国农业科学院,2014.CHENG L Q.Construction of genetic linkage map and QTL analysis for yield related traits in peanut(Arachis hypogaea L.)[D].Beijing:Chinese Academy of Agricultural Sciences,2014.(in Chinese)
[19]王传堂,王秀贞,唐月异,张建成,陈殿绪,崔凤高,禹山林,于树涛.花生健康组织和病组织简便快速DNA提取方法:中国,CN101805730 A.200910255786.0.WANG C T,WANG X Z,TANG Y Y,ZHANG J C,CHEN D X,CUI F G,YU S L,YU S T.A simple and rapid DNA extraction method from healthy and disease tissues in peanut:China,CN101805730 A,200910255786.0.(in Chinese)
[20]廖伯寿,雷永,姜慧芳,晏立英,王志慧,万丽云,淮东欣.一种花生高油酸回交育种后代基因型的快速精准鉴定方法:中国,CN105200142 A.201510675886.4.LIAO B S,LEI Y,JIANG H F,YAN L Y,WANG Z H,WAN L Y,HUAI D X.A rapid and accurate method for identifying the genotype of backcross progenies from high oleate peanut breeding:China,CN105200142 A.201510675886.4.(in Chinese)
[21]ZHAO S Z,LI A Q,LI C S,XIA H,ZHAO C Z,ZHANG Y,LEI H,WANG X J.Development and application of KASP marker for high throughput detection of Ah FAD2,mutation in peanut.Electronic Journal of Biotechnology,2017,25(C):9-12.
[22]姜慧芳,段乃雄.花生种质资源描述规范和数据标准.北京:中国农业出版社,2006.JIANG H F,DUAN N X.Descriptors and Dada Standard for Peanut.Beijing:China Agriculture Press,2006.(in Chinese)
[23]郭建斌.花生含油量及脂肪酸组成的QTL分析[D].武汉:华中农业大学,2016.GUO J B.QTL Analysis for oil content and fatty acid traits in Peanut(Arachis hypogaea L.)[D].Wuhan:Huazhong Agricultural University,2016.(in Chinese)
[24]段红梅.利用大豆SSR标记辅助遗传背景选择的效果分析[D].北京:中国农业科学院,2002.DUAN H M.Efficiency selection of genetic background using SSR markers in soybean[D].Beijing:Chinese Academy of Agricultural Sciences,2002.(in Chinese)
[25]SHIRASAWA K,BERTIOLI D J,VARSHNEY R K,MORETZSOHN M C,LEAL-BERTIOLI S C M,THUDI M,PANDEY M K,RAMI,J F,FONCéKA,D,GOWDA M V C,QIN H D,GUO B Z,HONG Y B,LIANG X Q,HIRAKAWA H,TABATA S,ISOBE S.Integrated consensus map of cultivated peanut and wild relatives reveals structures of the A and B Genomes of Arachis and divergence of the legume genomes.DNA Research an International Journal for Rapid Publication of Reports on Genes&Genomes,2013,20(2):173.
[26]ZHOU X J,YANG D,ZHAO J J,HUANG L,REN X P,CHEN Y N,HUANG S M,LIAO B S,LEI Y,YAN L Y,JIANG H F.Genomic survey sequencing for development and validation of single-locus SSR markers in peanut(Arachis hypogaea L.).BMC Genomics,2016,17(1):420.
[27]李丽,何美敬,崔顺立,侯名语,陈焕英,杨鑫雷,王鹏超,刘立峰,穆国俊.高油酸、中果型花生新材料的创制与鉴定.中国农业科学,2014,47(19):3898-3906.LI L,HE M J,CUI S L,HOU M Y,CHEN H Y,YANG X L,WANG P C,LIU L F,MU G J.The development and identification of new peanut germplasm materials with high oleic acid and medium pod.Scientia Agricultura Sinica,2014.(in Chinese)
[28]XU S S,CHU C G,HARRIS M O,WILLIAMS C E..Comparative analysis of genetic background in eight near-isogenic wheat lines with different H genes conferring resistance to Hessian fly.Genome,2011,54(1):81.
[29]JUNGMAN B S.The effect of fatty acid profiles on peanut seed germination at low soil temperatures[D].Lubbock:Texas Tech University,2000.
[30]XUE H Q,ISLEIB T G,PAYNE G A,WILSON R F,NOVITZKY W P,BRIAN G O.Comparison of aflatoxin production in normal-and high-oleic backcross-derived peanut lines.Plant Disease,2003,87(11):1360-1365.
[31]HOLBROOK C C,WILSON D M,MATHERON M E,HUNTER J E,KNAUFT D A,GORBET D W.Aspergillus colonization and aflatoxin contamination in peanut genotypes with reduced linoleic acid composition.Plant Disease,2000,84(2):148-150.
[32]BARKLEY N A,ISLEIB T G,WANG M L,PITTMAN R N.Genotypic effect of ah FAD2 on fatty acid profiles in six segregating peanut(Arachis hypogaea L.)populations.BMC Genetics,2013,14(1):62.
[2]禹山林.中国花生品种及其系谱.上海:上海科技出版社,2008.YU S L.Chinese Peanut Varieties and Its Pedigree.Shanghai:Shanghai Science and Technology Press,2008.(in Chinese)
[3]JANILA P,PANDEY M K,SHASIDHAR Y,VARIATH M T,SRISWATHI M,KHERA P,MANOHAR S S,NAGESH P,VISHWAKARMA M K,MISHRA G P,RADHAKRISHNAN T,MANIVANNAN N,DOBARIYA K L,VASANTHI R P,VARSHNEY R K.Molecular breeding for introgression of fatty acid desaturase mutant alleles(ah FAD2A and ah FAD2B)enhances oil quality in high and low oil containing peanut genotypes.Plant Science an International Journal of Experimental Plant Biology,2016,242:203-213.
[4]BRADDOC J C,SIMS C A,O'KEEFE S F.Flavor and oxidative stability of roasted high oleic acid peanuts.Journal Food Science,1995,60:489-493.
[5]MUGENDI J B,SIMS C A,GORGET D W,O'KEEFE S F.Flavor stability of high-oleic peanuts stored at low humidity.Journal of the American Oil Chemists Society,1998,75:21-25.
[6]O’BRYNE D J,KNAUFT D A,SHIREMAN R B.Low fatmonounsaturated rich diets containing high-oleic peanuts improve serum lipoprotein profiles.Lipids 1997,32:687-695.
[7]赵术珍,侯蕾,李长生,赵传志,任丽,李爱芹,邓丽,夏晗,王兴军.分子标记辅助回交选育高油酸花生新种质.中国油料作物学报,2017,39(1):30-36.ZHAO S Z,HOU L,LI C S,ZHAO C Z,REN L,LI A Q,DENG L,XIA H,WANG X J.Development of high oleic acid peanut from molecular marker assisted selection.Chinese Journal of Oil Crop Sciences,2017,39(1):30-36.(in Chinese)
[8]NAWADE B,BOSAMIA T C,THANKAPPAN R,RATHNAKUMAR A,KUMAR A,DOBARIA J R,KUNDU R,MISHRA G P.Insights into the indian peanut genotypes for ah FAD2 gene polymorphism regulating its oleic and linoleic acid fluxes.Frontiers in Plant Science,2016,7(1271):173-184.
[9]NORDEN A J,GORBET D W,KNAUFT D A,YOUNG C T.Variability in oil quality among peanut genotypes in the Florida breeding program.Peanut Science,1987,14:7-11.
[10]OKULEY J,LIGHTNER J,FELDMANN K,YADAV N,LARK E,BROWSE J.Arabidopsis FAD2 gene encodes the enzyme that is essential for polyunsaturated lipid synthesis.The Plant Cell,1994,6:147-158.
[11]JUNG S,SWIFT D,SENGOKU E,PATEL M,TEULéF,POWELL G,MOORE K,ABBOTT A.The high oleate trait in the cultivated peanut[Arachis hypogaea,L.]:I.Isolation and characterization of two genes encoding microsomal oleoyl-PC desaturases.Molecular&General Genetics Mgg,2000,263(5):796-805.
[12]WANG C T,HU D Q,DING F Y,YU H T,TANG Y Y,WANG X Z,ZHANG J C,CHEN D X.A new set of allele-specific PCR primers for identification of true hybrids in normal oleate×high oleate crosses in groundnut.Journal of SAT Agricultural Research,2011,12(9):1-4.
[13]CHEN Z,WANG M L,BARKLEY N A,PITTMAN R N.A simple allele-specific PCR assay for detecting FAD2 alleles in both A and B genomes of the cultivated peanut for high-oleate trait selection.Plant Molecular Biology Reporter,2010,28:542-548.
[14]BARKLEY N A,CHAMBERLIN K D C,WANG M L,PITTMAN R N.Development of a real-time PCR genotyping assay to identify high oleic acid peanuts(Arachis hypogaea L.).Molecular Breeding,2010,25:541-548.
[15]CHU Y,RAMOS L,HOLBROOK C C,OZIAS-AKINS P.Frequency of a Loss-of-Function Mutation in Oleoyl-PC Desaturase(ah FAD2A)in the Mini-Core of the U.S.Peanut Germplasm Collection.Crop Science.2007,47(6):2372-2378.
[16]曹永强,孙石.回交在转基因作物育种中的应用.作物杂志,2014(1):9-14.CAO Y Q,SUN S.Backcross breeding genetically modified crops in the application.Crops,2014(1):9-14.(in Chinese)
[17]王传堂,王秀贞,唐月异,吴琪,孙全喜,张建成,崔凤高,李利民,苗昊翠.中国高油酸花生种质创制、品种选育进展与建议.花生学报,2015(2):49-53.WANG C T,WANG X Z,TANG Y Y,WU Q,SUN Q X,ZHANG J C,CUI F G,LI L M,MIAO H C.High-oleic peanut germplasm enhancement and cultivar releases in China:Main achievements and suggestions.Journal of Peanut Science,2015(2):49-53.(in Chinese)
[18]成良强.花生遗传图谱构建及产量相关性状的QTL分析[D].北京:中国农业科学院,2014.CHENG L Q.Construction of genetic linkage map and QTL analysis for yield related traits in peanut(Arachis hypogaea L.)[D].Beijing:Chinese Academy of Agricultural Sciences,2014.(in Chinese)
[19]王传堂,王秀贞,唐月异,张建成,陈殿绪,崔凤高,禹山林,于树涛.花生健康组织和病组织简便快速DNA提取方法:中国,CN101805730 A.200910255786.0.WANG C T,WANG X Z,TANG Y Y,ZHANG J C,CHEN D X,CUI F G,YU S L,YU S T.A simple and rapid DNA extraction method from healthy and disease tissues in peanut:China,CN101805730 A,200910255786.0.(in Chinese)
[20]廖伯寿,雷永,姜慧芳,晏立英,王志慧,万丽云,淮东欣.一种花生高油酸回交育种后代基因型的快速精准鉴定方法:中国,CN105200142 A.201510675886.4.LIAO B S,LEI Y,JIANG H F,YAN L Y,WANG Z H,WAN L Y,HUAI D X.A rapid and accurate method for identifying the genotype of backcross progenies from high oleate peanut breeding:China,CN105200142 A.201510675886.4.(in Chinese)
[21]ZHAO S Z,LI A Q,LI C S,XIA H,ZHAO C Z,ZHANG Y,LEI H,WANG X J.Development and application of KASP marker for high throughput detection of Ah FAD2,mutation in peanut.Electronic Journal of Biotechnology,2017,25(C):9-12.
[22]姜慧芳,段乃雄.花生种质资源描述规范和数据标准.北京:中国农业出版社,2006.JIANG H F,DUAN N X.Descriptors and Dada Standard for Peanut.Beijing:China Agriculture Press,2006.(in Chinese)
[23]郭建斌.花生含油量及脂肪酸组成的QTL分析[D].武汉:华中农业大学,2016.GUO J B.QTL Analysis for oil content and fatty acid traits in Peanut(Arachis hypogaea L.)[D].Wuhan:Huazhong Agricultural University,2016.(in Chinese)
[24]段红梅.利用大豆SSR标记辅助遗传背景选择的效果分析[D].北京:中国农业科学院,2002.DUAN H M.Efficiency selection of genetic background using SSR markers in soybean[D].Beijing:Chinese Academy of Agricultural Sciences,2002.(in Chinese)
[25]SHIRASAWA K,BERTIOLI D J,VARSHNEY R K,MORETZSOHN M C,LEAL-BERTIOLI S C M,THUDI M,PANDEY M K,RAMI,J F,FONCéKA,D,GOWDA M V C,QIN H D,GUO B Z,HONG Y B,LIANG X Q,HIRAKAWA H,TABATA S,ISOBE S.Integrated consensus map of cultivated peanut and wild relatives reveals structures of the A and B Genomes of Arachis and divergence of the legume genomes.DNA Research an International Journal for Rapid Publication of Reports on Genes&Genomes,2013,20(2):173.
[26]ZHOU X J,YANG D,ZHAO J J,HUANG L,REN X P,CHEN Y N,HUANG S M,LIAO B S,LEI Y,YAN L Y,JIANG H F.Genomic survey sequencing for development and validation of single-locus SSR markers in peanut(Arachis hypogaea L.).BMC Genomics,2016,17(1):420.
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