小麦黄色素合成途径中Psy基因的克隆与分子特性及黄色素含量DH群体的构建
|
摘要
长期以来,我国人民对面粉及面制食品的色泽要求很高。普通小麦(Ttiticumaestivum L.2n=6x=AABBDD)籽粒黄色素含量(Yellow Pigment Content,YPC)是造成面粉色泽降低的重要因素之一,而且YPC对蒸煮面食品的色泽以及小麦的营养品质等均具有显著影响。虽然YPC受环境因素的影响,但基因型是其主要决定因素。黄色素是由多种物质组成的混合物,其主要成分是类胡萝卜素,类胡萝卜素的生物合成是在八氢番茄红素合成酶(Phytoene synthase,Psy)的作用下进行的,此酶为类胡萝卜素生物合成途径中的首要限速酶(Primary Rate-limiting Enzyme,PRH),其基因也是类胡萝卜素基因操作的首选基因,其他作物(尤其是玉米)中对Pay基因的研究已较为深入,但对普通小麦Psy基因的研究还未开展。本研究以已经发表的玉米Psy基因为基础,从YPC差异显著的中国小麦品种(系)中克隆普通小麦Psy基因,并进行分子特性和YPC分析;同时选择YPC差异显著的小麦品种组配F1与玉米远缘杂交,诱导可进行黄色素含量遗传分析的小麦双单倍体(Double Haploid,DH)群体,旨在明确Psy基因对小麦YPC的影响、开发小麦YPC的分子标记,同时为小麦YPC的遗传分析、高(低)YPC小麦新品种的选育提供遗传材料和新种质。主要结果如下:
1.小麦黄色素合成途径中Pay基因的克隆及分子特性利用玉米Pay基因和小麦EST序列设计引物克隆小麦Pay基因并测序,同时和玉米Pay基因进行序列比对分析。结果表明,5对引物Psy01、Psy03～Psy06在YPC不同的小麦样品中分别扩增出989bp、785 bp、1192 bp、690 bp和302 bp的5条带,同一引物在不同YPC品种中的PCR产物序列一致,经比对分析,5条序列均为Pay基因(片段),并由此组装成一条全长3437bp、包括6个外显子和5个内含子的小麦Psy基因。文中以引物Psy04为例对其扩增的PCR产物进行了详细的分子特性分析。
2.小麦Pay基因的等位变异及其对黄色素含量的影响根据已发表的麦族植物Pay基因序列的保守区设计引物Psy02,克隆小麦Psy基因。结果表明,该引物下PCR产物出现3种带型:196bp、233bp和489bp,其中196bp和233bp条带涵盖了小麦Psy基因第2外显子全部序列,相差的37bp为Psy基因第2内含子中的一段插入序列,可反映不同YPC,属小麦Pay基因的等位变异。验证试验表明,248份试材(包括229份中国小麦微核心种质和19份品质性状典型的小麦品种)中有153份材料(占样品数的65.7%)扩增出196bp条带,群体内YPC均值7.314mg/kg,属高YPC范畴;另有95份材料(占样品总数的38.3%)扩增出233bp条带,群体内YPC均值为5.207mg/kg,属低YPC范畴,方差分析表明二者YPC差异达1%极显著水平差异,说明上述37bp的插入序列是导致小麦品种间YPC产生差异的原因之一,因此该引物扩增的Psy基因对小麦YPC具有显著影响,引物Psy02是对小麦YPC进行分子鉴定的重要标记。序列分析表明,196bp和233bp与设计引物的探针序列100%同源,489bp序列的BLAST搜索显示,与序列EU096094(未定位)的同源率最高,达97%,与7A染色体上序列EF600063和7B染色体上序列DQ642439的同源率均为85%,表明该基因片段可能为7A和7B染色体以外的另一Psy基因。结合该序列对应小麦品种的高YPC,推测该基因和7A染色体的Psy基因对黄色素含量存在积加效应。
3.普通小麦DD染色体上Psy基因的发现及其等位变异以扩增小麦Psy基因的相应引物Psy01～Psy06,在节节麦(2X=DD=14)中进行PCR反应。结果表明,仅引物Psy02和Psy06能扩增出特异条带。引物Psy02的扩增产物长206bp,与普通小麦中扩增的196bp序列同源率为93.0%,对应的第2外显子区域内仅有1SNP;引物Psy06的PCR产物长305bp,与普通小麦中扩增的302bp序列同源率达95.77%,对应的第6外显子区域内无SNP,说明在小麦DD染色组中存在Psy基因,同时也排除了将引物Psy01、Psy03、Psy04、Psy05的扩增产物和引物Psy02的489bp扩增产物定位于D染色体的可能。
4.小麦黄色素含量DH群体的构建为获得一定规模的小麦单倍体以构建小麦黄色素含量的DH群体,用高/低YPC小麦品种组配F1,F1再和玉米远缘杂交诱导小麦单倍体,研究了环境温度对单倍体成胚率、幼胚取材时期、幼胚大小、4℃处理时间、暗处理时间对单倍体成苗率的影响。结果表明,获得最高成胚率的环境温度为21～23℃,授粉后12～16天取材的幼胚成苗率无明显差异;0.5～1.0mm大小的幼胚成苗率显著高于0～0.5mm和1.0～1.5mm大小的幼胚成苗率;1～3天短期4℃处理对胚萌发具有一定促进作用,但处理3天后,出愈率和成苗率都将降低;胚培养过程中12天左右的24h暗处理能有效提高成苗率,在此基础上建立了一套高效、可靠、重复性好的小麦×玉米单倍体诱导系统。
For a long time,color of wheat flour and flour foods have been required highly for people in China.Yellow pigment content(YPC) of wheat(Ttiticum aestivum L. 2n=6x=AABBDD) is one of important factors reducing flour color,and yellow pigment content is significant influence to cooked products and nutritional quality in wheat. Although environmental factors have significant effect on yellow pigment content,gene type is main decisive factors.Yellow pigment content is mixture composed of various substances,and its main components is carotenoids,biosynthesis of carotenoids need Phytoene synthase(Psy),the enzyme is the primary rate-limiting enzyme in biosynthetic pathway of carotenoids,and the gene is first selected gene to carotenoids genetic manipulation.Psy gene has been further studied in many crops(especially maize),but the research has not been developed in wheat.In this paper,wheat Psy gene with different significant YPC was cloned in 248 Chinese wheat cultivars based on maize Psy gene registered in GenBank,and the molecular characteristics of Psy gene and YPC of different wheat cultivars were analyzed too.At the same time,wheat F1 with high and low YPC parents were crossed by maize pollen in order to obtain wheat haploid embryos,then induced wheat double haploid for genetic analysis of yellow pigment content through embryo rescue and haploid chromosome doubling,the purposes are clearing the effect of Psy gene to wheat YPC,developing molecular markers of YPC in wheat,and creating genetic materials and new germplasm for wheat YPC genetic analysis and wheat YPC breeding.The results were as follows:
1.Cloning and molecular characterization of Psy gene in wheat yellow pigment biosynthesi Phytoene synthase(Psy) gene,a primary rate-limiting enzyme in the biosynthesis pathway of wheat yellow pigment(YP),was cloned and sequenced through Psy gene of maize and wheat ESTs in this study,at the same time,the Psy gene sequences between wheat and maize were compared.The results showed that,5 DNA fragments with 981bp,751bp,1192bp,690bp and 302bp was amplified by polymerase chain reaction in DNA samples extracted from the high YP and low YP content wheat cultivars,the sequences of high YPC wheat are same as those of low YPC with same primer.Sequences alignment showed that all the 5 DNA fragments were Psy gene(fragment) of wheat,and the full length gene with 6 exons and 5 introns,3437bp was assembled.Detailed molecular characterization of PCR products was analyzed with a case study of primer Psy04,and preliminary locating 5 Psy gene fragments on 7A chromosome in wheat.
2.Allelic variation of wheat Psy gene and effect on yellow pigment content Primer Psy02 was designed according to sequence conservative areas of Psy gene in Ttiticum,cloning Psy gene of wheat.The results showed that,3 kinds of PCR products were amplified with 196bp,233bp and 489bp,among them,196bp and 233bp bands cover all sequences of the second exon of wheat Psy gene,37bp difference sequence is a insertion sequence in the second exon of Psy gene,which display different YPC in wheat. Verification test showed,153 samples of 248 wheat cultivars(including 229 micro core germplasms and 19 cultivars with typical quality characters) amplified 196bp band, account for 61.7%,the YPC average value is 7.314mg/kg belonging to high YPC rang,and 95 samples amplified 233bp band,account for 38.3%,the YPC average value is 5.207mg/kg belonging to low YPC rang.Variance analysis showed that the YPC difference reached 1%significant level,which proved the 37bp insertion sequence is one of the reasons of different YPC in cultivars,therefore,primer Psy02 is a important molecular marker identifying YPC,and the gene effects on wheat YPC significantly.Sequence analysis showed that 196bp and 233bp were 100%homologous to primer probe sequence, and the two sequences were located on 7A chromosome.489bp sequence was high homologous to Psy-B1 of EU096094 with 97%identity,and 85%identity to the Psy-A1 of EF600063 on 7A chromosome and DQ642439 on 7B,thus the gene can not lacate on 7A or 7B chromosome.Combining with high YPC of 489bp,conjecturing the Psy genes on 7A and other chromosomes have additive effect to YPC.
3.Identifying Psy gene with Aegilops tauschii(Triticum tauschii L.2x=DD=14) and allelic variation Psy gene were cloned by primer Psy01-Psy06 in Aegilops tauschii (Triticum tauschii L.),the results showed that only primer Psy02 and Psy06 amplified specific bands,the PCR product of primer Psy was 206bp,with 93.0%sequence homologous to wheat PCR product,there was only 1SNP in the second exon.And the PCR product of primer Psy06 in Aegilops tauschii was 305bp,with 95.77%sequence homologous to wheat PCR product,there was no SNP in the 6th exon,which showed that there were Psy genes in DD genome of wheat,and the possibilities of locating primer Psy01,Psy03,Psy04,Psy05 and 489bp gene fragment of Psy02 on the D chromosome were eliminated.
4.Establishment of DH groups used to study inheritance laws in wheat yellow pigment In order to obtain a large number of wheat haploids to establish the DH groups of wheat yellow pigment content,wheat and maize cross were used to induce wheat haploid embryos and then the haploid plants were produced via young embryo culture.The factors affecting wheat embryos and the regeneration of young plants from young embryos, such as environmental temperature,different period of peeling embryo,size and development stage of young embryos,the treatment time at 4℃and dark treatment time were studied.The results showed:the embryos frequency was the highest when environmental temperature was 21～23℃,there was not significant difference on the frequency of young plants production when haploid embryos was peeled in different time from 12d to 16d;the frequency of young plants production of the size of 0.5～1.0mm is much higher than that of the size of 0～0.5mm and 1.0～1.5mm;the short time treatment between 1～3 days at 4℃would accelerate the regeneration of young embryo,but after 3 days the frequency of callus induction and plants improve the frequency of plants production during the process of young embryo culture.Production would decrease;dark treatment of all 24 hours for 12 days could effectively.And on this condition,a high efficiency and repetitive regeneration system of wheat haploid from wheat×maize was established.
1.小麦黄色素合成途径中Pay基因的克隆及分子特性利用玉米Pay基因和小麦EST序列设计引物克隆小麦Pay基因并测序,同时和玉米Pay基因进行序列比对分析。结果表明,5对引物Psy01、Psy03～Psy06在YPC不同的小麦样品中分别扩增出989bp、785 bp、1192 bp、690 bp和302 bp的5条带,同一引物在不同YPC品种中的PCR产物序列一致,经比对分析,5条序列均为Pay基因(片段),并由此组装成一条全长3437bp、包括6个外显子和5个内含子的小麦Psy基因。文中以引物Psy04为例对其扩增的PCR产物进行了详细的分子特性分析。
2.小麦Pay基因的等位变异及其对黄色素含量的影响根据已发表的麦族植物Pay基因序列的保守区设计引物Psy02,克隆小麦Psy基因。结果表明,该引物下PCR产物出现3种带型:196bp、233bp和489bp,其中196bp和233bp条带涵盖了小麦Psy基因第2外显子全部序列,相差的37bp为Psy基因第2内含子中的一段插入序列,可反映不同YPC,属小麦Pay基因的等位变异。验证试验表明,248份试材(包括229份中国小麦微核心种质和19份品质性状典型的小麦品种)中有153份材料(占样品数的65.7%)扩增出196bp条带,群体内YPC均值7.314mg/kg,属高YPC范畴;另有95份材料(占样品总数的38.3%)扩增出233bp条带,群体内YPC均值为5.207mg/kg,属低YPC范畴,方差分析表明二者YPC差异达1%极显著水平差异,说明上述37bp的插入序列是导致小麦品种间YPC产生差异的原因之一,因此该引物扩增的Psy基因对小麦YPC具有显著影响,引物Psy02是对小麦YPC进行分子鉴定的重要标记。序列分析表明,196bp和233bp与设计引物的探针序列100%同源,489bp序列的BLAST搜索显示,与序列EU096094(未定位)的同源率最高,达97%,与7A染色体上序列EF600063和7B染色体上序列DQ642439的同源率均为85%,表明该基因片段可能为7A和7B染色体以外的另一Psy基因。结合该序列对应小麦品种的高YPC,推测该基因和7A染色体的Psy基因对黄色素含量存在积加效应。
3.普通小麦DD染色体上Psy基因的发现及其等位变异以扩增小麦Psy基因的相应引物Psy01～Psy06,在节节麦(2X=DD=14)中进行PCR反应。结果表明,仅引物Psy02和Psy06能扩增出特异条带。引物Psy02的扩增产物长206bp,与普通小麦中扩增的196bp序列同源率为93.0%,对应的第2外显子区域内仅有1SNP;引物Psy06的PCR产物长305bp,与普通小麦中扩增的302bp序列同源率达95.77%,对应的第6外显子区域内无SNP,说明在小麦DD染色组中存在Psy基因,同时也排除了将引物Psy01、Psy03、Psy04、Psy05的扩增产物和引物Psy02的489bp扩增产物定位于D染色体的可能。
4.小麦黄色素含量DH群体的构建为获得一定规模的小麦单倍体以构建小麦黄色素含量的DH群体,用高/低YPC小麦品种组配F1,F1再和玉米远缘杂交诱导小麦单倍体,研究了环境温度对单倍体成胚率、幼胚取材时期、幼胚大小、4℃处理时间、暗处理时间对单倍体成苗率的影响。结果表明,获得最高成胚率的环境温度为21～23℃,授粉后12～16天取材的幼胚成苗率无明显差异;0.5～1.0mm大小的幼胚成苗率显著高于0～0.5mm和1.0～1.5mm大小的幼胚成苗率;1～3天短期4℃处理对胚萌发具有一定促进作用,但处理3天后,出愈率和成苗率都将降低;胚培养过程中12天左右的24h暗处理能有效提高成苗率,在此基础上建立了一套高效、可靠、重复性好的小麦×玉米单倍体诱导系统。
For a long time,color of wheat flour and flour foods have been required highly for people in China.Yellow pigment content(YPC) of wheat(Ttiticum aestivum L. 2n=6x=AABBDD) is one of important factors reducing flour color,and yellow pigment content is significant influence to cooked products and nutritional quality in wheat. Although environmental factors have significant effect on yellow pigment content,gene type is main decisive factors.Yellow pigment content is mixture composed of various substances,and its main components is carotenoids,biosynthesis of carotenoids need Phytoene synthase(Psy),the enzyme is the primary rate-limiting enzyme in biosynthetic pathway of carotenoids,and the gene is first selected gene to carotenoids genetic manipulation.Psy gene has been further studied in many crops(especially maize),but the research has not been developed in wheat.In this paper,wheat Psy gene with different significant YPC was cloned in 248 Chinese wheat cultivars based on maize Psy gene registered in GenBank,and the molecular characteristics of Psy gene and YPC of different wheat cultivars were analyzed too.At the same time,wheat F1 with high and low YPC parents were crossed by maize pollen in order to obtain wheat haploid embryos,then induced wheat double haploid for genetic analysis of yellow pigment content through embryo rescue and haploid chromosome doubling,the purposes are clearing the effect of Psy gene to wheat YPC,developing molecular markers of YPC in wheat,and creating genetic materials and new germplasm for wheat YPC genetic analysis and wheat YPC breeding.The results were as follows:
1.Cloning and molecular characterization of Psy gene in wheat yellow pigment biosynthesi Phytoene synthase(Psy) gene,a primary rate-limiting enzyme in the biosynthesis pathway of wheat yellow pigment(YP),was cloned and sequenced through Psy gene of maize and wheat ESTs in this study,at the same time,the Psy gene sequences between wheat and maize were compared.The results showed that,5 DNA fragments with 981bp,751bp,1192bp,690bp and 302bp was amplified by polymerase chain reaction in DNA samples extracted from the high YP and low YP content wheat cultivars,the sequences of high YPC wheat are same as those of low YPC with same primer.Sequences alignment showed that all the 5 DNA fragments were Psy gene(fragment) of wheat,and the full length gene with 6 exons and 5 introns,3437bp was assembled.Detailed molecular characterization of PCR products was analyzed with a case study of primer Psy04,and preliminary locating 5 Psy gene fragments on 7A chromosome in wheat.
2.Allelic variation of wheat Psy gene and effect on yellow pigment content Primer Psy02 was designed according to sequence conservative areas of Psy gene in Ttiticum,cloning Psy gene of wheat.The results showed that,3 kinds of PCR products were amplified with 196bp,233bp and 489bp,among them,196bp and 233bp bands cover all sequences of the second exon of wheat Psy gene,37bp difference sequence is a insertion sequence in the second exon of Psy gene,which display different YPC in wheat. Verification test showed,153 samples of 248 wheat cultivars(including 229 micro core germplasms and 19 cultivars with typical quality characters) amplified 196bp band, account for 61.7%,the YPC average value is 7.314mg/kg belonging to high YPC rang,and 95 samples amplified 233bp band,account for 38.3%,the YPC average value is 5.207mg/kg belonging to low YPC rang.Variance analysis showed that the YPC difference reached 1%significant level,which proved the 37bp insertion sequence is one of the reasons of different YPC in cultivars,therefore,primer Psy02 is a important molecular marker identifying YPC,and the gene effects on wheat YPC significantly.Sequence analysis showed that 196bp and 233bp were 100%homologous to primer probe sequence, and the two sequences were located on 7A chromosome.489bp sequence was high homologous to Psy-B1 of EU096094 with 97%identity,and 85%identity to the Psy-A1 of EF600063 on 7A chromosome and DQ642439 on 7B,thus the gene can not lacate on 7A or 7B chromosome.Combining with high YPC of 489bp,conjecturing the Psy genes on 7A and other chromosomes have additive effect to YPC.
3.Identifying Psy gene with Aegilops tauschii(Triticum tauschii L.2x=DD=14) and allelic variation Psy gene were cloned by primer Psy01-Psy06 in Aegilops tauschii (Triticum tauschii L.),the results showed that only primer Psy02 and Psy06 amplified specific bands,the PCR product of primer Psy was 206bp,with 93.0%sequence homologous to wheat PCR product,there was only 1SNP in the second exon.And the PCR product of primer Psy06 in Aegilops tauschii was 305bp,with 95.77%sequence homologous to wheat PCR product,there was no SNP in the 6th exon,which showed that there were Psy genes in DD genome of wheat,and the possibilities of locating primer Psy01,Psy03,Psy04,Psy05 and 489bp gene fragment of Psy02 on the D chromosome were eliminated.
4.Establishment of DH groups used to study inheritance laws in wheat yellow pigment In order to obtain a large number of wheat haploids to establish the DH groups of wheat yellow pigment content,wheat and maize cross were used to induce wheat haploid embryos and then the haploid plants were produced via young embryo culture.The factors affecting wheat embryos and the regeneration of young plants from young embryos, such as environmental temperature,different period of peeling embryo,size and development stage of young embryos,the treatment time at 4℃and dark treatment time were studied.The results showed:the embryos frequency was the highest when environmental temperature was 21～23℃,there was not significant difference on the frequency of young plants production when haploid embryos was peeled in different time from 12d to 16d;the frequency of young plants production of the size of 0.5～1.0mm is much higher than that of the size of 0～0.5mm and 1.0～1.5mm;the short time treatment between 1～3 days at 4℃would accelerate the regeneration of young embryo,but after 3 days the frequency of callus induction and plants improve the frequency of plants production during the process of young embryo culture.Production would decrease;dark treatment of all 24 hours for 12 days could effectively.And on this condition,a high efficiency and repetitive regeneration system of wheat haploid from wheat×maize was established.
引文
[1]刘建军,何中虎,赵振东,等小麦品质性状与干面条品质参数关系的研究[J],作物学报,2002,28(6):738-742
[2]He,Z.H.,Liu,A.H.,Pena,R.J.,and Rajaram,S.Suitability of Chinese wheat cultivars for production of northern style Chinese steamed bread[J].Euphytiea.2003,131:155-163
[3]何中虎,庄巧生.编.中国小麦品种改良及系谱分析[A].北京:中国农业出版社.2003.519-542
[4]常成.小麦及近缘种属籽粒硬度、多酚氧化酶性状的分子机理研究[D].北京,2007,中国农业大学博士学位论文
[5]张立平,葛秀秀,何中虎,等.普通小麦多酚氧化酶活性的QTL分析[J].作物学报,2005,31(1):7-10
[6]杨朝柱,马传喜,司红起,等.普通小麦品种多酚氧化酶活性的变异[J].中国农业科学,2004,37(11):1713-1717
[7]孙道杰,何中虎,张立平,等.小麦多酚氧化酶(PPO)活性的SSR标记筛选与验证[J].中国农业科学,2005,38(7):1295-1299
[8]Anderson,J.V.and C.F.Morris.An improved whole-seed assay for screen-ing wheat germplasm for polyphenol oxidase activity[J].Crop Sci.2001,41:1697-1705
[9]Okot-Kotber,M.A.,Liavoga,A.,Yong,K.J.,and Bagorogoza,K.Activity and inhibition of polyphenol oxidase in extracts of bran and other milling fractions from a variety of wheat eultivars[J].Creal Chem.2001,78:514-520
[10]Demeke T,and Morris C F.Moleeulax characterization of wheat poly-phenol oxidase (PPO)[J].Thero.Appl.Genet.,2002,104:813-818
[11]Zhang LP,Yan J,Xia XC et al.,2006.QTL mapping for kernel yellow pigment content in common wheat[J].Aeta Agronomica Siniea,32(11):51-45
[12]W.Zhang,J.Dubcovsky,2008.Association between allelie variation at the Phytoene synthase 1 gene and yellow pigment content in the wheat grain[J].Theor Appl Genet, 116(5):635-45.
[13]Naik PS,Khurana SM,Kalloo G,2003.Genetic manipulation of carotenoid pathway in higher plants[J].Current Science,85(1):1423-1430
[14]Sharp PJ,2001.Validation of molecular markers for wheat breeding[J].Australian Journal of Agricultural Research,52:1357-1366
[15]Busch M.,Seuter A.,and Haln R.Functional analysis of the early steps of carotenoid biosynthesis in tobacco[J].Plant Physiol.,2002,128:439-453
[16]Ducreux L.J.,Morris W.L.,Hedley P.E.,Shepherd T.,Davies H.V.,Millam S.,and Taylor M.A.Metabolic engineering of high carotenoid potato tubers containing enhanced levels of beta-carotene and lutein[J].Exp.Bot.,2005,56(409):81-89
[17]吕欣,毛忠贵.玉米黄色素研究进展[J].中国食品添加剂,2003,3:57-60
[18]Liehtenthaler H.K.,Schwender J.,Disch A.,and Rohmer M.Biosynthesis of isoprenoids in higher plants chloroplasts proceeds via a mevalonate independent pathway[J],FEBS Lett.,1997,40(3):271-274
[19]Hirschberg J.Carotenoid biosynthesis in flowering plants,Curr[J].Opin.Plant Biol.,2001,4(3):210-218
[20]Isaaeson T.,Ronen G.,Zamir D.,and Hirschberg J.Cloning of tangerine from tomato reveals a carotenoid isomerase essential for the production of β-carotene and xanthophylls in plants[J].Plant Cell,2002,14(2):333-342
[21]Park H.,Kreunen S.S.,Cuttriss A.J.,DellaPerma D.,and Pogson B.J.Identification of the carotene isomerase provides insight into carotenoid biosynthesis,prolamellar body formation,and photomorphogenesis[J].Plant Cell,2002,14:321-332
[22]Cunningham F.X.,and Gantt E.One ring or two? Determination of ring number in earotenoids by lyeopene epsilon-eyclases[J].Proe.Natl.Aead.Sei.,2001,98(5):2905-2910
[23]Pogson B.,McDonald K.A.,Truong M.,Britton G.,and DellaPerma D.,1996,Arabidopsis carotenoid mutants demonstrate that lutein is not essential for photosynthesis in higher plants[J].Plant Cell,8(9):1627-1639
[24]Albrecht M,Misawa N,Sandmann G.Metabolic engineering of the terpenoid biosynthetic pathway of Escherichia coli for production of the carotenoids beta-carotene and zeaxanthin[J].Biotechnol Lett,1999,21:791-795
[25]Alicia N.Massa·Brian Beecher·Craig F.Morris Polyphenol oxidase(PPO) in wheat and wild relatives:molecular evidence for a multigene family[J].Theor Appl Genet.2007,114:1239-1247
[26]Johnson E A,Schroeder W A.Microbial carotenoids[J].Adv Biochem Eng Biotechnol,1995,53:119-178
[27]Hirschberg J.Carotenoid biosynthesis in flowering plants[j].Currt Opin Plant Biol,2001,4:210-218
[28]Dvorak J,McGuire PE,Cassidy B.Apparent sources of the Agenomes of wheats inferred from polymorphism in abundance and restriction fragment length of repeated nucleotide sequences[J].Genome.1998,30,680-689.
[29]Busch M.,Seuter A.,and Hain R.Functional analysis of the early steps of carotenoid biosynthesis in Tobacco[J].Plant Physiol.,2002,128:439-453
[30]Boze H,Moulin G,Galzy P.Production of food and fodder yeasts[J].Crit Rev Biotechnol,1992,12:65-86
[31]A.W.Campbell,W.B.Griffin.The Effects of Temperature and Light Intensity on Embryo Numbers in Wheat Double Haploid Production through Wheat×Maize crosses[J].Annals of Botany 1998,82:29-33
[32]Atienza SG,Ram'lrez MC,Hern'andez P,Martin A.Chromosomal location of genes for carotenoid content in Hordeum chilens[J]e.Plant Breeding,2004,123,303-304
[33]Artley G E,Scolnik P A.Plant carotenoids:Pigments for photo protection,visual attraction and human health[J].Plant Cell,1995,7:1027-1038
[34]姜文侯,单志萍,孟好.微生物产生的类胡萝卜素及其工业化[J].食品与发酵工业.1999,3:46-48
[35]J oppa LR,Williams ND.Langdon durum disomic substitution lines and aneuploid analysis in tetraploid wheat[J].Genome,1998,30,222-228.
[36]Misawa N,Satomi Y,Kondo K et al.Structure and functional analysis of a marine bacterial carotenoid biosynthesis gene cluster and astaxanthin biosynthetic pathway proposed at the gene level[J].Bactefio1,1995,177:6575-65842
[37]Park H.,Kreunen S.S.,Cuttriss A.J.,DellaPenna D.,and Pogson B.J.Identification of the carotene isomerase provides insight into carotenoid biosynthesis,prolamellar body formation,and photomorphogenesis[J].Plant Cell,2002,14:321-332
[38]Krinaky N I.Carotenoids in medicine.In:carotenoids.Chemistry and Biochemistry[J].Plenum,New York,1989,279-291
[39]Riera-Lizarazu O,Mujeeb-Kasi A and William Mdhm.Maize mediated wheat polyhaploid production using various crossing methods[J].Cereal Res.Commun,1992,46:335-346
[40]Truesdale M.,Bramley M.,and Schuch W.Cloning and characterization of gene involved in phytoene synthesis from tomato[J].Plant Mol.Biol.,1987,19(3):401-404
[41]Yang J,Zhang Y,He ZH,Yan J,Wang DS,Liu JJ,Wang MF.Association between wheat quality traits and performance of pan bread and dry white Chinese noodle[J].Acta Agron Sin,2004,30:739-744
[42]Suenaga K,Nakajima K.Variation in doubled haploid plants of wheat obtained through wheat(Triticum aestivum)×maize(Zea mays L.) crosses[J].Plant Breeding,1993,111:120-124
[43]P A Scolnik and G E Bartley,1994.Nucleotide sequence of an Arabidopsis cDNA for phytoene synthase[J].Plant Physiol.104(4):1471-1472
[44]Misawa N,Shimada H.Metabolic engineering for the production of carotenoids in non2carotenogenic bacteria and yeasts.J Biotechnol,1998,59:169-181
[45]范晓岚,糜漫天,杨慧.类胡萝卜素与衰老[J].辽宁医学杂志,2003,17(6):41-42
[46]Sies H,Stahl W.Vitamin E and C,B-carotene,and other carotenoids as antioxidants[J].Am J Clin Nutr,1995,62(1):1315-1321
[47]Burtun G W.Antioxidant action of carotenoids[J].Nutr,1989,119:109-111
[48]Kohlmeier,Hastings S B.Epidem iologic ev idence of a role of carotenoids in cardiovascular disease prevention[J].AM J C lin Nutr,1995,62:1370-1376
[49]Buba,W atza B,Abraham.Moderate intervention with carotenoid-rich vegetable product reduces lipid peroxidation in men[J].Nutr,2000,130:2200-2206
[50]Ronen G,Carmel,Goren L,Zamir D,et al.,2000.An alternative pathway to β-carotene formation in plant chromoplasts discovered by map-based cloning of beta and old-gold color mutations in tomato[J].Proceedings of the National Academy of Sciences of the United States of America,97:11102-11107.
[51]潘天红,苗玲,金慧芳,等.脑梗死患者血清B-胡萝卜素及低密度脂蛋白的含量分析[J].中风与神经疾病杂志,2000,17(2):75-76
[52]Stahl,W,Heinrich U,Jungm ann H,et al.Carotenoids and carotenoids plus vitam in E protect against ultraviolet light-induced erythem in humans[J].Am J Clin Nutr,2000,71:795-798
[53]张敬,张军,刘艺敏,等.B-胡萝卜素对染镍肺泡巨噬细胞MDA及SOD的影响[J].中国公共卫生,2002,18(8):898-899
[54]Kondo K,Salto T,Kajiwara S et al.A transformation systemfor the yeast Candida utilis:use of a modified endogenous ribosomal protein gone as a drug2resistant marker and ribosomal DNA as an integration target for vector DNA[J].Bactedol,1995,177:7171-7177
[55]Hirschberg J.Carotenoid biosynthesis in flowering plants[J].Plant Biol.,2001,4(3);210-218
[56]Gr(u|¨)newald K,Hagen C:β2carotene is the intermediate exported from the chloroplast during accumulation of secondary carotenoids in Haematococcus pluvialis[J].Applied Phycol,2001,13:89-93
[57]Ebiamadon Andi Brisibe,Alena Gajdosova,Annette Olesen and Sven Bode Andersen.Cytodifferentiation and transformation of embryogenic callus lines derived from anther culture of wheat[J].Journal of Experimental Botany,2000,51(343):187-196
[58]Cunningham FX,Gantt E.Genes and enzymes of carotenoid biosynthesis in plants[J].Annual Review of Plant Physiology and Plant Molecular Biology,1998,49,557-583
[59]Al-Babili S.,Hartung W.,Kleinig H.,and Beyer P.CPTA modulates levels of carotenogenic proteins and their mRNAs and affects carotenoid and ABA content as well as chromo last structure in Narcissus pseudo narcissus flowers[J],Plant Biol.,1999,1:607-612
[59]Chamovitz D.,Sandmann G.,and Hirschberg J.Molecular and biochemical characterization of herbicide-resistant mutants of cyanobactefia reveals that phytoene desaturation is a rote-limiting step in carotenoid biosynthesis[J].Biol.Chem.,1993,268(23):17348-17353
[60]Shewmaker C K,Sheehy J A,Daley M,et al,.Seed specific overexpression of phytoebe synthase:increase in carotenoids and other metabolic effects[J].Plant,1999,20(4):401-412
[61]Cunningham EX.,Pogson B.,Sun Z.,McDonald K.A.,DellaPenna D.,and Gantt E.Functional analysis of the β and slycopene cyclase enzymes of Arabidopsis reveals amcchanism for control of cyclic carotenoid formation[J].Plant Cell,1996,8(9):1613-1626
[62]Fraser P D,Mcr S,Shipton C A,et al..Biochemical evaluation of transgcnic tomato plants expressing an addition phytoene synthase in a fruit specific manner[J].Proceedings of the National Academy of Sciences of the United States of America,2002,99:1092-I097
[63]Goodwin T W,Britton G.Distribution and analysis of carotenoids.In:Goodwin TW (Ed.)[J].Plant Pigments.Academic Press,London,1988,61-132
[64]Jyonouchi H,Hill J,Tornita Yet al.Studies of immunomodulating actions of carotenoids.Ⅰ.Effects of β2carotene and astaxanthin on murine lymphocyte functions and cell surface marker expression in in vivo culturesystcm[J].Nutr Cancer,1991,16:93-105
[65]Murdock D,Ensley B D,Serdar C et al.Construction of metabolic opcrons catalyzing the de novo biosynthesis of indigo in Eschcrichia coli[J]. Biotechnology,1993,11:381-386
[66]朱长甫,陈星,王英典.植物类胡萝卜素生物合成有其相关基因的基因工程中的应用[J].植物生物与分子生物学报,2004,30(6):609-618
[67]Giovannucci E,Aschedo A,Rimm E B et al.Intake of carotenoids and retinol in relation to risk of prostate cancer[J].J Natl Cancer Inst,1995,87:1767-1776
[68]Bird C R,Ray J A,Fletcher J D,et al,.Using antisense RNA to study gene fuction:inhibition of carotenoid biosynthesis in transgenic tomatoes[J].Bio Technol,1991,9(7):635-639
[69]Bramley P.,Teulieres C.,Blain I.,Bird C.,and Schuch W.Biochemical characterization of transgenic tomato in which carotenoid biosynthesis has been inhibited through the expression of antisense RNA to pTOMS[J].Plant,1992,2:343-349
[70]陈选阳,袁照年,张招娟,等.甘薯块根总RNA提取与psy基因保守序列的克隆[J].福建农林大学学报,2005,34(1):259-263
[71]Palaisa KA,Morgante M,Williams M,et al.,2003.Contrasting effects of selection on sequence diversity and linkage disequilibrium at two phytoene synthase loc[J].Plant Cell,15:1795-1806
[72]Hirschberg J.Carotenoid biosynthesis in flowering plants[J].Current Opinion in Plant Biology,2001,4,210-218
[73]Ichii T,Takchara S,Konno H et al.Development of a new commercial-scale airlift fermentor for rapid growth of yeast[J].Ferment Bioeng,1993,75:375-379
[74]Fraser PD,Bramlcy PM.The biosynthesis and nutritional uses of carotenoids[J].Progress in Lipid Research.2004,43,228-265
[75]Fraser P.D.,Kiano J.W.,Truesdale M.R.,Schuch W.,and Branley P.M.Phytoene synthase-2 enzyme activity in tomato does not contribute to carotenoid biosynthesis in ripening fruit[J].Plant Mol.Biol.,1999,40(4):687-698
[76]He,Z.H.,Yang J.,Zhang Y.,Quail K.J.and Pena R.J.Pan bread and dry white Chinese noodle quality in Chinese winter wheats[J].Euphytica.2004.139:257-267
[77]Fraser ED.,Truesdale M.R.,Bird C.R.,Schuch W.,and BramleyP.M.Carotenoid biosynthesis during tomato fruit development[J].Plant Physiol.,1994,105(1):405-413
[78]Buekner B,San Miguel P,Janick-Buekner D,Bennetzen JL.The ylgene of maize codes for phytoene synthase[J].Genetics,1996,143,479-488.
[79]王玉萍,刘庆昌.翟红植物类胡萝卜素生物合成相关基因的表达调控及其在植物基因工程中的应用[J].分子植物育种,2006,4(1):103-110
[80]Belitz,H.D.and Grosch,W.Food chemistry[M].Springer Verlag.Berlin.1987
[81]Wang C W,Oh M K,Liao J C.Engineered isoprenoid pathway enhances astaxanthin production in Eschedchia coli[J].Biotechnol Bioeng,1999,62:235-241
[82]Mares D J,Campbell AW.Mapping components of flour colour in Australian wheat[J].Australian Journal of Agricultural Research.2001,52,1297-1309.
[83]张立平,阎俊,夏先春,何中虎,Mark W,Suthedand.普通小麦籽粒黄色素含量的QTL分析[J].作物学报,2006,32(11):41-45
[84]查如璧,国淑惠,赵中民,王德森,陆建中.我国硬粒小麦品质性状及质量分级标准研究[J].中国农业科学,1995,28(02):8-14
[85]W.Zhang and Dubcovsky,Association between allelie variation at the Phytoene synthase 1 gene and yellow pigment content in the wheat grain[J].Theor Appl Genet 2008,116:635-645
[86]Kuehel H,Langridge P,Mosionek L,et al..The genetic control of milling yield,dough rheology and baking quality of wheat[J].Theor Appl Genet,2006,112:1487-1495
[87]Miskelly DM.Flour components aflecting paste and noodle colour[J].J Sci Food Agrie,1984,35:463-471
[88]Baik B K,Czuchajowska Z,Pomeranz Y.Discoloration of dough for oriental noodle[J].Cereal Chem,1995,72:198-205
[89]Parker G D,Chalmers KJ,Rathjen A J,Langridge P.Mapping loci associated with flour colour in wheat(Tdticurn aestivum L.)[J].Theor Appl Genet,1998,97:238-245
[90]杨金,张艳,何中虎,阎俊,王德森,刘建军,王美芳.小麦品质性状与面包和面条品质关系分析[J].作物学报.2004.30(8):739-744
[91]Oliver,J.R.The colour of flour streams as related to ash and pigment contents[J].J.of Cereal Sci.1993.17:169-182
[92]Symons.S.J.Computer analysis of fluorescence for the measurement of flour refinement as determined by flour ash content,flour grade colour,and tristimulus colour measurements[J].Cereal Chem.1991.68:454-460
[93]葛秀秀,何中虎,杨金,张歧军.我国冬小麦品种多酚氧化酶活性的遗传变异及其与品质性状的相关分析[J].作物学报.2003.29:125-131
[94]Oliver,J.R.'Proceedings of the 38th Australian Cereal Chemistry Conference'[C],Royal Australian Chemical Institute,Parkville,Victoria,Australia,1988.216-218
[95]杨朝柱,张磊,司红起,马传喜.小麦面粉白度研究进展麦类作物学报[J].2002,22(3):74-77
[96]查如壁,国淑惠,赵中民,王德森,陆建中.我国硬粒小麦品质性状及质量分级标准研究[J].中国农业科学,1995,28(02):8-14
[97]Ward,S.Millikan,M.and Wootton,M.Examination of carotenoid pigments in wheat[A].Proceedings of the 46thAustralian Cereal Chemistry Conference',Royal Australian Chemical Institute,Parkville,Victoria,Australia,1996.145-149
[98]Ward,S.Millikan,M.and Wootton,M.Quantification of carotenoid compounds in Australian wheats[A].'Proceedings of the 47th Australian Cereal Chemistry Conference',Royal Australian Chemical Institute,Parkville,Vietoria,Australia,1997.294-298
[99]Mares,D.J.Wang,Y.and Cassidy,C.A.Separation,identification and tissue location of compounds responsible for the yellow colour of alkaline noodles[A].'Proceedings of the 47th Australian Cereal Chemistry Conference',Royal Australian Chemical Institute, North Melbourne,Victoria,Australia,1997.114-117
[100]Gallagher CE,Matthews PD,Li F,Wurtzel ET.Gene duplication inthe carotenoid biosynthetic pathway preceded evolution of the grasses[J].Plant Physiology,2004,135:1776-1783.
[101]Bailey J E.Toward a science of metabolic engineering[J].Science 1991,252:1668-1675
[102]Hirschberg J,Cohen M,Harker Met al.Molecular genetics of the carotenoid biosynthesis pathway in plants and algae[J].Pure Appl Chem,1997,69:2151-2158
[103]Inagaki,M and M.Tahir.Comparison of haploid production frequencies in wheat varieties crossed with Hordeum bulbosum L.and maize[J].Japan J Breed,1990,40:209-216
[104]Kajiwara S,Fraser P D,Kondo K et al.Expression of an exogenous isopentenyl diphosphate iso-merase gene enhances isoprenoid biosyn2 thesis in Escherichia coli[J].Biochem,1997,324:421-426
[105]Misawa N,Nakagawa M,Kobayashi K et al.Elucidation of the Erwinia uredevora carotenoid biosynthetic pathway by functional analysis of gene products expressed in Escherichia coli[J].Bacteriol,1990,172:6704-6712
[106]Elouafi I,Nachit MM,Martin LM.Identification of a microsatellite on chromosome 7B showing a strong linkage with yellow pigment in durum wheat(Triticum turgidum L.var.durum)[J].Hereditas,2001,135,255-261
[107]Yamano S,Ishii T,Nakagawa M et al.Metabolic engineering for production of b2carotene and lycopene in Saccharomyces cerevisiae[J].Biosci Biotechnol Biochem,1994,58:1112-1114
[108]Pozniak CJ,Knox RE,Clarke FR,et al.,2007.Identification of QTL and association of a Phytoene synthase gene with endosperm colour in durum wheat[J].Theor Appl Genet 114:525-537
[109]Shimada H,Kondo K,Fraser P et al.Increased carotenoid production by the food yeast Candida utilis through metabolic engineering of the isoprenoid pathway[J].Appl Environ Microbiol,1998,64:2676-2680
[110]Alvarez J B,Martin L M,Martin A.Chromosomal localization of genes for carotenoid pigments using addition lines of Hordeum chilense in wheat[J].Plant Breeding,1998.117,287-289
[111]Alvarez JB,Martin LM,Martin A.Genetic variation for carotenoid pigment content in the amphiploid Hordeum chilense×Triticum turgidum conv.durum[J].Plant Breeding,1995,118,187-189
[112]He,Z.H.,Yang J.,Zhang Y.,Quail K.J.and Pena R.J.Pan bread and dry white Chinese noodle quality in Chinese winter wheats[J].Euphytica.2004.139:257-267
[113]X.Y.He,Y.L.Zhang,Z.H.He.Characterization of phytoene synthase 1 gene(Psyl)located on common wheat chromosome 7A and development of a functional marker[J].Theor Appl Genet.2008,116:213-221
[114]杨紫薇,方震,李高潮,罗红炼,赵鹏涛.硬粒小麦面筋和黄色素含量的早代选择[J].麦类作物,1999,19(6):7-9
[115]陈新民,陈孝.小麦×玉米产生单倍体及双单倍体研究进展[J].麦类作物,1998,18(3)1-4
[116]Snape,J.W.,V.Chapman,J.Moss.The cross abilities of wheat varieties with Hordeumbulbousm[J].Heredity,1979,42:291-298
[117]Zenkteler M,Nitzsche W.Wide hybridization experiments in cereals[J].Theor Appl Genet,1984,68:311-315.
[118]Laurie D.A.Wheat×maize hybridization[J].Can.J.Genet.Cytol,1986,28:313-316.
[119]Laurie D.A,Bennett MD.The effect of the crossability loci Kr1 and Kr2 on fertilization frequency in hexaploid wheat×maize crosses[J].Theor.Appl.Genet,1988,76:393-397.
[120]孙敬三,刘辉,路铁刚,等.利用小麦×玉米获得小麦单倍体[J].植物学报,1992,34(11):817-821.
[121]陈新民,徐惠君,周俊芳,等.提高小麦×玉米胚培养植株产生频率的研究[J].中 国农业科学,1996,29(4):29-32.
[122]Campbell A W,Griffi W B,Burdtt D J,Conner A J..The importance of light intensity for pollen tube growth and embryo survival in wheat×maize crosses[J].Annals of Botany,2001,87:517-522
[123]蔡华,马传喜,黄正来,等.小麦×玉米诱导小麦单倍体高效系统的建立[J].核农学报,2006,20(2):90-94
[124]闫林,王成社,王辉.不同处理对小麦×玉米产生小麦单倍体胚频率的影响[J].西北植物学报,2007,27(8):1593-1599
[125]来长凯,王成社,闫林.冬小麦×玉米远缘杂交产生小麦单倍体胚的研究[J].麦类作物学报,2007,27(2):193-196
[126]顾坚,刘琨,李绍祥,等.小麦×玉米杂交诱导小麦单倍体的割穗离体培养研究[J].麦类作物学报,2008,28(1):1-5
[127]Inagaki M.,Tahir M.Production of haploid wheat through intergeneric crosses[J].Hereditas.1992,116:117-120
[128]Inagaki,M.,M.Tahir.Comparison of haploid production frequence in wheat varieties crossed with Hordeum bulbosum L.and Malze[J].Japan J.Breed,1990,40:209-216
[129]Kisan N.S.et al.Production of haploid wheat plants from wheat×maize crosses[J].Plant Breed,1993,110:96-102.
[130]陈新民.提高小麦×玉米产生单倍体频率的研究.第六届全国作物育种讨论会论文集[D],北京,1997.
[131]蔡华,马传喜,周淼平,等.利用小麦与玉米远缘杂交诱导小麦双单倍体的研究.植物细胞工程与分子育种技术研究[D].北京:中国农业科学技术出版社,2003:68-71
[132]蔡华,马传喜,陆维忠.小麦与玉米远缘杂交诱导小麦单倍体的研究[J].麦类作物学报,2004:24(2):11-14
[133]王广金.小麦与玉米杂交产生单倍体频率的研究[J].麦类作物,1998,18(6):12-14
[134]陈新民,张文祥,崔淑兰,等.小麦×玉米产生小麦单倍体的染色体加倍研究[J].中国农业科学,2002,35(4):447-450
[135]Inagaki M.et al.Use of pollen storage and detached tiller culture in wheat polyploid production though wide crosses[J].Cereal Research Comm.1997,25:7-13.
[136]胡冬梅.利用小麦×玉米属间杂交产生小麦单倍体植株研究初报[J].青海农林科技,1998,3:6-9
[137]孙敬三,路铁刚,辛化伟.利用染色体消除法获得太谷核不育小麦纯合体[J].植物学报,1999,41(1):254-257
[138]蔡华,马传喜,陆维忠.小麦×玉米产生的小麦单倍体植株的染色体加倍研究[J].麦类作物学报,2005:25(3):20-22
[139]李振声,容珊.小麦远缘杂交.[M].科学出版社,北京,1985.
[140]Sharp PJ,2001.Validation of molecular markers for wheat breeding[J].Australian Journal of Agricultural Research,52:1357-1366
[141]Riera-Lizarazu O.Maize mediated wheat polyhaploid production using various crossing methods[J].Cereal Res.Commun.1991,18:339-346.
[142]N.Amrani.小麦×玉米产生单倍体的遗传变异性[J].国外农学-麦类作物,1994,2:2-5.
[143]陈新民,李学渊,陈孝,等.不同杂交技术对小麦×玉米产生单倍体的影响[J].作物学报,1998,24(6):743-746
[144]李大玮,欧阳平,邱纪文,等.染色体消失法在小麦育种中应用的研究[J].农业生物技术学报.2000,8(1):17-21
[145]Mujeeb-Kazia.Polyhaploidy in the Triticeae madited by crosses Triticum species with Zea mays and Tdpsacum datyloides[J].Int.Wheat Genet,1993,8:1119-1123.
[146]陈新民,张文祥等小麦×玉米产生小麦单倍体的染色体加倍研究[J].中国农业科学,2002,35(4):447-450.
[147]孙敬三,路铁刚,辛化伟.硬粒小麦单倍体高效再生体系的建立.中国农业科 学.1996,29(3):61-65
[148]刘辉,陈纯贤.小麦×玉米杂交后代的蛋白质及酯酶同工酶分析[J].植物学报,1999,41(1):55-59.
[149]G..Brazauskas,I.Pasakinskien,A.Jahoor.AFLP analysis indicates no introgression of maize DNA in wheat×maize crosses[J].Plant Breeding 2004,123:117-121.
[150]R(e|¨)mer S,Fraser P D,2005.Recent advances in carotenoid biosynthesis,regulation and manipulation[J].Planta.211:305-308
[151]R(o|¨)mer S,Hugueney P,Bouvier F,et al.,1993.Expression of the genes encoding the early carotenoid biosynthetic enzymes in Capsicum armuum[J].Biochem Biophys Res Commun,196:1414-1421
[152]Schmimidt,Darmert C.Engineering novel carotenoids in microorganisms[J].Curr Opin Biotechnol,2000,11:255-261
[153]Zoi Karvounil,Isaac Johnl,Jane E,et al.,1995.Turnerl and Don Grierson Isolation and eharaetedsation of a melon cDNA clone encoding phytoene synthase[J].Plant Molecular Biology,27:1153-1162
[154]Burkhardt PK,Beyer P,Warm J,et al.,1997.Transgenie rice(Oryza sativa)endosperm expressing daffodil(Narcissus pseudonarcissus) phytoene synthase accumulates phytoene,a key intermediate of provitamin A biosynthesis[J].Plant Physiol,11(5):1071-1078.
[155]Faqiang Li,Ratnakar Vallabhaneni,and Eleanore T.PSY3,a New Member of the Phytoene Synthase Gene Family Conserved in the Poaceae and Regulator of Abiotic Stress-Induced Root Carotenogenesis[J].Plant Physiol,2008,146(3):1333-1345
[156]Li Y Y,Ma C Z,Fu T D,et al..Construction of a molecular functional map ofrapeseed(Brassiea napus L.) using differentially expressed genesbetween hybrid and its parents[J].Euphytiea,2006,152:25-39.
[157]Ronen G,Carmel,Goren L,Zamir D,et al.,2000.An alternative pathway to β-carotene formation in plant ehromoplasts discovered by map-based cloning of beta and old-gold color mutations in tomato[J].Proceedings of the National Academy of Sciences of the United States of America,97:11102-11107.
[158]Wong J C,Lambert R J and Wurtzel E T,2004.QTL and candidate genes phytoene synthase and carotene desaturase associated with the accumulation of carotenoids in maize[J].Theor Appl Genet,108:349-359
[159]Gale MD,Devos KM Comparative genetics in the grasses[J].Proc Natl Acad Sci USA,1998,95:1971-1974
[160]孙道杰,何中虎,王辉.小麦面粉黄色素相关基因研究[J].西北植物学报,2006,26(4):0655-0660
[161]AACC(1995) Approved Methods of the American Association of Cereal Chemists,9th end.St.Paul,MN,USA
[162]Guo M,Rupe M A,Zinselmeier C,et al.Allelic variation of gene expression in maize hybrids[J].Plant Cell,2004,16(7):1707-1716.
[163]Knott DR.Translocations involving Triticum chromosomes and Agropyron chromosomes carrying rust resistance[J].Canadian Journal of Genetics and Cytology,1968,10:695-696.
[164]Riera-Lizarazu O,Rines H W,Phillips R L.Maize chromosome retention,sexual transimission,and introgression in plants generated by oat×maize crosses[J].In:Abstract of the second international conference on the plant genome.San Diego:California,1994.178
[165]Cenci A,Somma S,Chantret N,Dubcovsky J,Blanco A.PCR identification of durum wheat BAC clones containing genes coding for carotenoid biosynthesis enzymes and their chromosome localization[J].Genome,2004,47,911-917
[166]Ruther A,Misawa N,BEger P et al.Production of zeaxanthin in Eschedchia coli transformed with different carotenogenic plasmids[J].Appl Microbiol Biotechnol,1997,48:162-167
[167]Atienza SG,Avila CM,Ramirez MC,Martin A.Application of near infrared reflectance spectroscopy to the determination of carotenoid content in tritordeum for breeding purposes[J].Australian Journal of Agricultural Research,2005,56,85-89
[168]Atienza SG,Ballesteros J,Martin A,Hornero-M'endez D.Genetic variability of carotenoid concentration and degree of esterification among tritordeum(×Tritordeum Ascherson et Graebner) and durum wheat accessions.Journal of Agricultural and Food Chemistry[J],2007,55,4244-4251
[169]李懋学,陈瑞阳.关于植物核型分析的标准化问题[J].武汉植物学研究,1985,3(4):297-302.
[170]陈瑞阳,宋文芹,李秀兰,等.中国主要经济植物基因组染色体图谱(第3册)[J].北京:科学出版社,2003:1-152.
[171]安海龙,卫志明,黄健秋.小麦幼胚培养高效成株系统的建立[J].植物生理学报,2000,26(6):532-538.
[172]宋国琦,王成社,何培茹.小麦幼胚培养技术及其应用的研究进展[J].西安联合大学学报.2003,6(2):22-27
[173]陈耀锋,曹团武,丁云姣,等.普通小麦基因转化中良好受体系统的建立[J].西北农林科技大学学报(自然科学版).2004,32(9):1-3
[174]王子宁,张艳敏,郭北海,等.利用单倍体育种技术快速培育糯性小麦新品系[J].华北农学报,2001,16(1):1-6
[175]Comeau A,Nadeau P,Ploured A.Media for the in ovule culture of proembryos of wheat and wheat-derived interspecific hybrids or haploids[J].Plant Sci,1992,81:117-125
[176]陈纯贤,孙敬三,朱立煌.由小麦×玉米获得的普通小麦加倍单倍体后代的RFLP变异[J].植物学报,1999,41(1):55-59
[177]李振声.用于小麦染色体工程的蓝粒小麦单体系列材料的创制[J].遗传,2001,23(1):42-42
[2]He,Z.H.,Liu,A.H.,Pena,R.J.,and Rajaram,S.Suitability of Chinese wheat cultivars for production of northern style Chinese steamed bread[J].Euphytiea.2003,131:155-163
[3]何中虎,庄巧生.编.中国小麦品种改良及系谱分析[A].北京:中国农业出版社.2003.519-542
[4]常成.小麦及近缘种属籽粒硬度、多酚氧化酶性状的分子机理研究[D].北京,2007,中国农业大学博士学位论文
[5]张立平,葛秀秀,何中虎,等.普通小麦多酚氧化酶活性的QTL分析[J].作物学报,2005,31(1):7-10
[6]杨朝柱,马传喜,司红起,等.普通小麦品种多酚氧化酶活性的变异[J].中国农业科学,2004,37(11):1713-1717
[7]孙道杰,何中虎,张立平,等.小麦多酚氧化酶(PPO)活性的SSR标记筛选与验证[J].中国农业科学,2005,38(7):1295-1299
[8]Anderson,J.V.and C.F.Morris.An improved whole-seed assay for screen-ing wheat germplasm for polyphenol oxidase activity[J].Crop Sci.2001,41:1697-1705
[9]Okot-Kotber,M.A.,Liavoga,A.,Yong,K.J.,and Bagorogoza,K.Activity and inhibition of polyphenol oxidase in extracts of bran and other milling fractions from a variety of wheat eultivars[J].Creal Chem.2001,78:514-520
[10]Demeke T,and Morris C F.Moleeulax characterization of wheat poly-phenol oxidase (PPO)[J].Thero.Appl.Genet.,2002,104:813-818
[11]Zhang LP,Yan J,Xia XC et al.,2006.QTL mapping for kernel yellow pigment content in common wheat[J].Aeta Agronomica Siniea,32(11):51-45
[12]W.Zhang,J.Dubcovsky,2008.Association between allelie variation at the Phytoene synthase 1 gene and yellow pigment content in the wheat grain[J].Theor Appl Genet, 116(5):635-45.
[13]Naik PS,Khurana SM,Kalloo G,2003.Genetic manipulation of carotenoid pathway in higher plants[J].Current Science,85(1):1423-1430
[14]Sharp PJ,2001.Validation of molecular markers for wheat breeding[J].Australian Journal of Agricultural Research,52:1357-1366
[15]Busch M.,Seuter A.,and Haln R.Functional analysis of the early steps of carotenoid biosynthesis in tobacco[J].Plant Physiol.,2002,128:439-453
[16]Ducreux L.J.,Morris W.L.,Hedley P.E.,Shepherd T.,Davies H.V.,Millam S.,and Taylor M.A.Metabolic engineering of high carotenoid potato tubers containing enhanced levels of beta-carotene and lutein[J].Exp.Bot.,2005,56(409):81-89
[17]吕欣,毛忠贵.玉米黄色素研究进展[J].中国食品添加剂,2003,3:57-60
[18]Liehtenthaler H.K.,Schwender J.,Disch A.,and Rohmer M.Biosynthesis of isoprenoids in higher plants chloroplasts proceeds via a mevalonate independent pathway[J],FEBS Lett.,1997,40(3):271-274
[19]Hirschberg J.Carotenoid biosynthesis in flowering plants,Curr[J].Opin.Plant Biol.,2001,4(3):210-218
[20]Isaaeson T.,Ronen G.,Zamir D.,and Hirschberg J.Cloning of tangerine from tomato reveals a carotenoid isomerase essential for the production of β-carotene and xanthophylls in plants[J].Plant Cell,2002,14(2):333-342
[21]Park H.,Kreunen S.S.,Cuttriss A.J.,DellaPerma D.,and Pogson B.J.Identification of the carotene isomerase provides insight into carotenoid biosynthesis,prolamellar body formation,and photomorphogenesis[J].Plant Cell,2002,14:321-332
[22]Cunningham F.X.,and Gantt E.One ring or two? Determination of ring number in earotenoids by lyeopene epsilon-eyclases[J].Proe.Natl.Aead.Sei.,2001,98(5):2905-2910
[23]Pogson B.,McDonald K.A.,Truong M.,Britton G.,and DellaPerma D.,1996,Arabidopsis carotenoid mutants demonstrate that lutein is not essential for photosynthesis in higher plants[J].Plant Cell,8(9):1627-1639
[24]Albrecht M,Misawa N,Sandmann G.Metabolic engineering of the terpenoid biosynthetic pathway of Escherichia coli for production of the carotenoids beta-carotene and zeaxanthin[J].Biotechnol Lett,1999,21:791-795
[25]Alicia N.Massa·Brian Beecher·Craig F.Morris Polyphenol oxidase(PPO) in wheat and wild relatives:molecular evidence for a multigene family[J].Theor Appl Genet.2007,114:1239-1247
[26]Johnson E A,Schroeder W A.Microbial carotenoids[J].Adv Biochem Eng Biotechnol,1995,53:119-178
[27]Hirschberg J.Carotenoid biosynthesis in flowering plants[j].Currt Opin Plant Biol,2001,4:210-218
[28]Dvorak J,McGuire PE,Cassidy B.Apparent sources of the Agenomes of wheats inferred from polymorphism in abundance and restriction fragment length of repeated nucleotide sequences[J].Genome.1998,30,680-689.
[29]Busch M.,Seuter A.,and Hain R.Functional analysis of the early steps of carotenoid biosynthesis in Tobacco[J].Plant Physiol.,2002,128:439-453
[30]Boze H,Moulin G,Galzy P.Production of food and fodder yeasts[J].Crit Rev Biotechnol,1992,12:65-86
[31]A.W.Campbell,W.B.Griffin.The Effects of Temperature and Light Intensity on Embryo Numbers in Wheat Double Haploid Production through Wheat×Maize crosses[J].Annals of Botany 1998,82:29-33
[32]Atienza SG,Ram'lrez MC,Hern'andez P,Martin A.Chromosomal location of genes for carotenoid content in Hordeum chilens[J]e.Plant Breeding,2004,123,303-304
[33]Artley G E,Scolnik P A.Plant carotenoids:Pigments for photo protection,visual attraction and human health[J].Plant Cell,1995,7:1027-1038
[34]姜文侯,单志萍,孟好.微生物产生的类胡萝卜素及其工业化[J].食品与发酵工业.1999,3:46-48
[35]J oppa LR,Williams ND.Langdon durum disomic substitution lines and aneuploid analysis in tetraploid wheat[J].Genome,1998,30,222-228.
[36]Misawa N,Satomi Y,Kondo K et al.Structure and functional analysis of a marine bacterial carotenoid biosynthesis gene cluster and astaxanthin biosynthetic pathway proposed at the gene level[J].Bactefio1,1995,177:6575-65842
[37]Park H.,Kreunen S.S.,Cuttriss A.J.,DellaPenna D.,and Pogson B.J.Identification of the carotene isomerase provides insight into carotenoid biosynthesis,prolamellar body formation,and photomorphogenesis[J].Plant Cell,2002,14:321-332
[38]Krinaky N I.Carotenoids in medicine.In:carotenoids.Chemistry and Biochemistry[J].Plenum,New York,1989,279-291
[39]Riera-Lizarazu O,Mujeeb-Kasi A and William Mdhm.Maize mediated wheat polyhaploid production using various crossing methods[J].Cereal Res.Commun,1992,46:335-346
[40]Truesdale M.,Bramley M.,and Schuch W.Cloning and characterization of gene involved in phytoene synthesis from tomato[J].Plant Mol.Biol.,1987,19(3):401-404
[41]Yang J,Zhang Y,He ZH,Yan J,Wang DS,Liu JJ,Wang MF.Association between wheat quality traits and performance of pan bread and dry white Chinese noodle[J].Acta Agron Sin,2004,30:739-744
[42]Suenaga K,Nakajima K.Variation in doubled haploid plants of wheat obtained through wheat(Triticum aestivum)×maize(Zea mays L.) crosses[J].Plant Breeding,1993,111:120-124
[43]P A Scolnik and G E Bartley,1994.Nucleotide sequence of an Arabidopsis cDNA for phytoene synthase[J].Plant Physiol.104(4):1471-1472
[44]Misawa N,Shimada H.Metabolic engineering for the production of carotenoids in non2carotenogenic bacteria and yeasts.J Biotechnol,1998,59:169-181
[45]范晓岚,糜漫天,杨慧.类胡萝卜素与衰老[J].辽宁医学杂志,2003,17(6):41-42
[46]Sies H,Stahl W.Vitamin E and C,B-carotene,and other carotenoids as antioxidants[J].Am J Clin Nutr,1995,62(1):1315-1321
[47]Burtun G W.Antioxidant action of carotenoids[J].Nutr,1989,119:109-111
[48]Kohlmeier,Hastings S B.Epidem iologic ev idence of a role of carotenoids in cardiovascular disease prevention[J].AM J C lin Nutr,1995,62:1370-1376
[49]Buba,W atza B,Abraham.Moderate intervention with carotenoid-rich vegetable product reduces lipid peroxidation in men[J].Nutr,2000,130:2200-2206
[50]Ronen G,Carmel,Goren L,Zamir D,et al.,2000.An alternative pathway to β-carotene formation in plant chromoplasts discovered by map-based cloning of beta and old-gold color mutations in tomato[J].Proceedings of the National Academy of Sciences of the United States of America,97:11102-11107.
[51]潘天红,苗玲,金慧芳,等.脑梗死患者血清B-胡萝卜素及低密度脂蛋白的含量分析[J].中风与神经疾病杂志,2000,17(2):75-76
[52]Stahl,W,Heinrich U,Jungm ann H,et al.Carotenoids and carotenoids plus vitam in E protect against ultraviolet light-induced erythem in humans[J].Am J Clin Nutr,2000,71:795-798
[53]张敬,张军,刘艺敏,等.B-胡萝卜素对染镍肺泡巨噬细胞MDA及SOD的影响[J].中国公共卫生,2002,18(8):898-899
[54]Kondo K,Salto T,Kajiwara S et al.A transformation systemfor the yeast Candida utilis:use of a modified endogenous ribosomal protein gone as a drug2resistant marker and ribosomal DNA as an integration target for vector DNA[J].Bactedol,1995,177:7171-7177
[55]Hirschberg J.Carotenoid biosynthesis in flowering plants[J].Plant Biol.,2001,4(3);210-218
[56]Gr(u|¨)newald K,Hagen C:β2carotene is the intermediate exported from the chloroplast during accumulation of secondary carotenoids in Haematococcus pluvialis[J].Applied Phycol,2001,13:89-93
[57]Ebiamadon Andi Brisibe,Alena Gajdosova,Annette Olesen and Sven Bode Andersen.Cytodifferentiation and transformation of embryogenic callus lines derived from anther culture of wheat[J].Journal of Experimental Botany,2000,51(343):187-196
[58]Cunningham FX,Gantt E.Genes and enzymes of carotenoid biosynthesis in plants[J].Annual Review of Plant Physiology and Plant Molecular Biology,1998,49,557-583
[59]Al-Babili S.,Hartung W.,Kleinig H.,and Beyer P.CPTA modulates levels of carotenogenic proteins and their mRNAs and affects carotenoid and ABA content as well as chromo last structure in Narcissus pseudo narcissus flowers[J],Plant Biol.,1999,1:607-612
[59]Chamovitz D.,Sandmann G.,and Hirschberg J.Molecular and biochemical characterization of herbicide-resistant mutants of cyanobactefia reveals that phytoene desaturation is a rote-limiting step in carotenoid biosynthesis[J].Biol.Chem.,1993,268(23):17348-17353
[60]Shewmaker C K,Sheehy J A,Daley M,et al,.Seed specific overexpression of phytoebe synthase:increase in carotenoids and other metabolic effects[J].Plant,1999,20(4):401-412
[61]Cunningham EX.,Pogson B.,Sun Z.,McDonald K.A.,DellaPenna D.,and Gantt E.Functional analysis of the β and slycopene cyclase enzymes of Arabidopsis reveals amcchanism for control of cyclic carotenoid formation[J].Plant Cell,1996,8(9):1613-1626
[62]Fraser P D,Mcr S,Shipton C A,et al..Biochemical evaluation of transgcnic tomato plants expressing an addition phytoene synthase in a fruit specific manner[J].Proceedings of the National Academy of Sciences of the United States of America,2002,99:1092-I097
[63]Goodwin T W,Britton G.Distribution and analysis of carotenoids.In:Goodwin TW (Ed.)[J].Plant Pigments.Academic Press,London,1988,61-132
[64]Jyonouchi H,Hill J,Tornita Yet al.Studies of immunomodulating actions of carotenoids.Ⅰ.Effects of β2carotene and astaxanthin on murine lymphocyte functions and cell surface marker expression in in vivo culturesystcm[J].Nutr Cancer,1991,16:93-105
[65]Murdock D,Ensley B D,Serdar C et al.Construction of metabolic opcrons catalyzing the de novo biosynthesis of indigo in Eschcrichia coli[J]. Biotechnology,1993,11:381-386
[66]朱长甫,陈星,王英典.植物类胡萝卜素生物合成有其相关基因的基因工程中的应用[J].植物生物与分子生物学报,2004,30(6):609-618
[67]Giovannucci E,Aschedo A,Rimm E B et al.Intake of carotenoids and retinol in relation to risk of prostate cancer[J].J Natl Cancer Inst,1995,87:1767-1776
[68]Bird C R,Ray J A,Fletcher J D,et al,.Using antisense RNA to study gene fuction:inhibition of carotenoid biosynthesis in transgenic tomatoes[J].Bio Technol,1991,9(7):635-639
[69]Bramley P.,Teulieres C.,Blain I.,Bird C.,and Schuch W.Biochemical characterization of transgenic tomato in which carotenoid biosynthesis has been inhibited through the expression of antisense RNA to pTOMS[J].Plant,1992,2:343-349
[70]陈选阳,袁照年,张招娟,等.甘薯块根总RNA提取与psy基因保守序列的克隆[J].福建农林大学学报,2005,34(1):259-263
[71]Palaisa KA,Morgante M,Williams M,et al.,2003.Contrasting effects of selection on sequence diversity and linkage disequilibrium at two phytoene synthase loc[J].Plant Cell,15:1795-1806
[72]Hirschberg J.Carotenoid biosynthesis in flowering plants[J].Current Opinion in Plant Biology,2001,4,210-218
[73]Ichii T,Takchara S,Konno H et al.Development of a new commercial-scale airlift fermentor for rapid growth of yeast[J].Ferment Bioeng,1993,75:375-379
[74]Fraser PD,Bramlcy PM.The biosynthesis and nutritional uses of carotenoids[J].Progress in Lipid Research.2004,43,228-265
[75]Fraser P.D.,Kiano J.W.,Truesdale M.R.,Schuch W.,and Branley P.M.Phytoene synthase-2 enzyme activity in tomato does not contribute to carotenoid biosynthesis in ripening fruit[J].Plant Mol.Biol.,1999,40(4):687-698
[76]He,Z.H.,Yang J.,Zhang Y.,Quail K.J.and Pena R.J.Pan bread and dry white Chinese noodle quality in Chinese winter wheats[J].Euphytica.2004.139:257-267
[77]Fraser ED.,Truesdale M.R.,Bird C.R.,Schuch W.,and BramleyP.M.Carotenoid biosynthesis during tomato fruit development[J].Plant Physiol.,1994,105(1):405-413
[78]Buekner B,San Miguel P,Janick-Buekner D,Bennetzen JL.The ylgene of maize codes for phytoene synthase[J].Genetics,1996,143,479-488.
[79]王玉萍,刘庆昌.翟红植物类胡萝卜素生物合成相关基因的表达调控及其在植物基因工程中的应用[J].分子植物育种,2006,4(1):103-110
[80]Belitz,H.D.and Grosch,W.Food chemistry[M].Springer Verlag.Berlin.1987
[81]Wang C W,Oh M K,Liao J C.Engineered isoprenoid pathway enhances astaxanthin production in Eschedchia coli[J].Biotechnol Bioeng,1999,62:235-241
[82]Mares D J,Campbell AW.Mapping components of flour colour in Australian wheat[J].Australian Journal of Agricultural Research.2001,52,1297-1309.
[83]张立平,阎俊,夏先春,何中虎,Mark W,Suthedand.普通小麦籽粒黄色素含量的QTL分析[J].作物学报,2006,32(11):41-45
[84]查如璧,国淑惠,赵中民,王德森,陆建中.我国硬粒小麦品质性状及质量分级标准研究[J].中国农业科学,1995,28(02):8-14
[85]W.Zhang and Dubcovsky,Association between allelie variation at the Phytoene synthase 1 gene and yellow pigment content in the wheat grain[J].Theor Appl Genet 2008,116:635-645
[86]Kuehel H,Langridge P,Mosionek L,et al..The genetic control of milling yield,dough rheology and baking quality of wheat[J].Theor Appl Genet,2006,112:1487-1495
[87]Miskelly DM.Flour components aflecting paste and noodle colour[J].J Sci Food Agrie,1984,35:463-471
[88]Baik B K,Czuchajowska Z,Pomeranz Y.Discoloration of dough for oriental noodle[J].Cereal Chem,1995,72:198-205
[89]Parker G D,Chalmers KJ,Rathjen A J,Langridge P.Mapping loci associated with flour colour in wheat(Tdticurn aestivum L.)[J].Theor Appl Genet,1998,97:238-245
[90]杨金,张艳,何中虎,阎俊,王德森,刘建军,王美芳.小麦品质性状与面包和面条品质关系分析[J].作物学报.2004.30(8):739-744
[91]Oliver,J.R.The colour of flour streams as related to ash and pigment contents[J].J.of Cereal Sci.1993.17:169-182
[92]Symons.S.J.Computer analysis of fluorescence for the measurement of flour refinement as determined by flour ash content,flour grade colour,and tristimulus colour measurements[J].Cereal Chem.1991.68:454-460
[93]葛秀秀,何中虎,杨金,张歧军.我国冬小麦品种多酚氧化酶活性的遗传变异及其与品质性状的相关分析[J].作物学报.2003.29:125-131
[94]Oliver,J.R.'Proceedings of the 38th Australian Cereal Chemistry Conference'[C],Royal Australian Chemical Institute,Parkville,Victoria,Australia,1988.216-218
[95]杨朝柱,张磊,司红起,马传喜.小麦面粉白度研究进展麦类作物学报[J].2002,22(3):74-77
[96]查如壁,国淑惠,赵中民,王德森,陆建中.我国硬粒小麦品质性状及质量分级标准研究[J].中国农业科学,1995,28(02):8-14
[97]Ward,S.Millikan,M.and Wootton,M.Examination of carotenoid pigments in wheat[A].Proceedings of the 46thAustralian Cereal Chemistry Conference',Royal Australian Chemical Institute,Parkville,Victoria,Australia,1996.145-149
[98]Ward,S.Millikan,M.and Wootton,M.Quantification of carotenoid compounds in Australian wheats[A].'Proceedings of the 47th Australian Cereal Chemistry Conference',Royal Australian Chemical Institute,Parkville,Vietoria,Australia,1997.294-298
[99]Mares,D.J.Wang,Y.and Cassidy,C.A.Separation,identification and tissue location of compounds responsible for the yellow colour of alkaline noodles[A].'Proceedings of the 47th Australian Cereal Chemistry Conference',Royal Australian Chemical Institute, North Melbourne,Victoria,Australia,1997.114-117
[100]Gallagher CE,Matthews PD,Li F,Wurtzel ET.Gene duplication inthe carotenoid biosynthetic pathway preceded evolution of the grasses[J].Plant Physiology,2004,135:1776-1783.
[101]Bailey J E.Toward a science of metabolic engineering[J].Science 1991,252:1668-1675
[102]Hirschberg J,Cohen M,Harker Met al.Molecular genetics of the carotenoid biosynthesis pathway in plants and algae[J].Pure Appl Chem,1997,69:2151-2158
[103]Inagaki,M and M.Tahir.Comparison of haploid production frequencies in wheat varieties crossed with Hordeum bulbosum L.and maize[J].Japan J Breed,1990,40:209-216
[104]Kajiwara S,Fraser P D,Kondo K et al.Expression of an exogenous isopentenyl diphosphate iso-merase gene enhances isoprenoid biosyn2 thesis in Escherichia coli[J].Biochem,1997,324:421-426
[105]Misawa N,Nakagawa M,Kobayashi K et al.Elucidation of the Erwinia uredevora carotenoid biosynthetic pathway by functional analysis of gene products expressed in Escherichia coli[J].Bacteriol,1990,172:6704-6712
[106]Elouafi I,Nachit MM,Martin LM.Identification of a microsatellite on chromosome 7B showing a strong linkage with yellow pigment in durum wheat(Triticum turgidum L.var.durum)[J].Hereditas,2001,135,255-261
[107]Yamano S,Ishii T,Nakagawa M et al.Metabolic engineering for production of b2carotene and lycopene in Saccharomyces cerevisiae[J].Biosci Biotechnol Biochem,1994,58:1112-1114
[108]Pozniak CJ,Knox RE,Clarke FR,et al.,2007.Identification of QTL and association of a Phytoene synthase gene with endosperm colour in durum wheat[J].Theor Appl Genet 114:525-537
[109]Shimada H,Kondo K,Fraser P et al.Increased carotenoid production by the food yeast Candida utilis through metabolic engineering of the isoprenoid pathway[J].Appl Environ Microbiol,1998,64:2676-2680
[110]Alvarez J B,Martin L M,Martin A.Chromosomal localization of genes for carotenoid pigments using addition lines of Hordeum chilense in wheat[J].Plant Breeding,1998.117,287-289
[111]Alvarez JB,Martin LM,Martin A.Genetic variation for carotenoid pigment content in the amphiploid Hordeum chilense×Triticum turgidum conv.durum[J].Plant Breeding,1995,118,187-189
[112]He,Z.H.,Yang J.,Zhang Y.,Quail K.J.and Pena R.J.Pan bread and dry white Chinese noodle quality in Chinese winter wheats[J].Euphytica.2004.139:257-267
[113]X.Y.He,Y.L.Zhang,Z.H.He.Characterization of phytoene synthase 1 gene(Psyl)located on common wheat chromosome 7A and development of a functional marker[J].Theor Appl Genet.2008,116:213-221
[114]杨紫薇,方震,李高潮,罗红炼,赵鹏涛.硬粒小麦面筋和黄色素含量的早代选择[J].麦类作物,1999,19(6):7-9
[115]陈新民,陈孝.小麦×玉米产生单倍体及双单倍体研究进展[J].麦类作物,1998,18(3)1-4
[116]Snape,J.W.,V.Chapman,J.Moss.The cross abilities of wheat varieties with Hordeumbulbousm[J].Heredity,1979,42:291-298
[117]Zenkteler M,Nitzsche W.Wide hybridization experiments in cereals[J].Theor Appl Genet,1984,68:311-315.
[118]Laurie D.A.Wheat×maize hybridization[J].Can.J.Genet.Cytol,1986,28:313-316.
[119]Laurie D.A,Bennett MD.The effect of the crossability loci Kr1 and Kr2 on fertilization frequency in hexaploid wheat×maize crosses[J].Theor.Appl.Genet,1988,76:393-397.
[120]孙敬三,刘辉,路铁刚,等.利用小麦×玉米获得小麦单倍体[J].植物学报,1992,34(11):817-821.
[121]陈新民,徐惠君,周俊芳,等.提高小麦×玉米胚培养植株产生频率的研究[J].中 国农业科学,1996,29(4):29-32.
[122]Campbell A W,Griffi W B,Burdtt D J,Conner A J..The importance of light intensity for pollen tube growth and embryo survival in wheat×maize crosses[J].Annals of Botany,2001,87:517-522
[123]蔡华,马传喜,黄正来,等.小麦×玉米诱导小麦单倍体高效系统的建立[J].核农学报,2006,20(2):90-94
[124]闫林,王成社,王辉.不同处理对小麦×玉米产生小麦单倍体胚频率的影响[J].西北植物学报,2007,27(8):1593-1599
[125]来长凯,王成社,闫林.冬小麦×玉米远缘杂交产生小麦单倍体胚的研究[J].麦类作物学报,2007,27(2):193-196
[126]顾坚,刘琨,李绍祥,等.小麦×玉米杂交诱导小麦单倍体的割穗离体培养研究[J].麦类作物学报,2008,28(1):1-5
[127]Inagaki M.,Tahir M.Production of haploid wheat through intergeneric crosses[J].Hereditas.1992,116:117-120
[128]Inagaki,M.,M.Tahir.Comparison of haploid production frequence in wheat varieties crossed with Hordeum bulbosum L.and Malze[J].Japan J.Breed,1990,40:209-216
[129]Kisan N.S.et al.Production of haploid wheat plants from wheat×maize crosses[J].Plant Breed,1993,110:96-102.
[130]陈新民.提高小麦×玉米产生单倍体频率的研究.第六届全国作物育种讨论会论文集[D],北京,1997.
[131]蔡华,马传喜,周淼平,等.利用小麦与玉米远缘杂交诱导小麦双单倍体的研究.植物细胞工程与分子育种技术研究[D].北京:中国农业科学技术出版社,2003:68-71
[132]蔡华,马传喜,陆维忠.小麦与玉米远缘杂交诱导小麦单倍体的研究[J].麦类作物学报,2004:24(2):11-14
[133]王广金.小麦与玉米杂交产生单倍体频率的研究[J].麦类作物,1998,18(6):12-14
[134]陈新民,张文祥,崔淑兰,等.小麦×玉米产生小麦单倍体的染色体加倍研究[J].中国农业科学,2002,35(4):447-450
[135]Inagaki M.et al.Use of pollen storage and detached tiller culture in wheat polyploid production though wide crosses[J].Cereal Research Comm.1997,25:7-13.
[136]胡冬梅.利用小麦×玉米属间杂交产生小麦单倍体植株研究初报[J].青海农林科技,1998,3:6-9
[137]孙敬三,路铁刚,辛化伟.利用染色体消除法获得太谷核不育小麦纯合体[J].植物学报,1999,41(1):254-257
[138]蔡华,马传喜,陆维忠.小麦×玉米产生的小麦单倍体植株的染色体加倍研究[J].麦类作物学报,2005:25(3):20-22
[139]李振声,容珊.小麦远缘杂交.[M].科学出版社,北京,1985.
[140]Sharp PJ,2001.Validation of molecular markers for wheat breeding[J].Australian Journal of Agricultural Research,52:1357-1366
[141]Riera-Lizarazu O.Maize mediated wheat polyhaploid production using various crossing methods[J].Cereal Res.Commun.1991,18:339-346.
[142]N.Amrani.小麦×玉米产生单倍体的遗传变异性[J].国外农学-麦类作物,1994,2:2-5.
[143]陈新民,李学渊,陈孝,等.不同杂交技术对小麦×玉米产生单倍体的影响[J].作物学报,1998,24(6):743-746
[144]李大玮,欧阳平,邱纪文,等.染色体消失法在小麦育种中应用的研究[J].农业生物技术学报.2000,8(1):17-21
[145]Mujeeb-Kazia.Polyhaploidy in the Triticeae madited by crosses Triticum species with Zea mays and Tdpsacum datyloides[J].Int.Wheat Genet,1993,8:1119-1123.
[146]陈新民,张文祥等小麦×玉米产生小麦单倍体的染色体加倍研究[J].中国农业科学,2002,35(4):447-450.
[147]孙敬三,路铁刚,辛化伟.硬粒小麦单倍体高效再生体系的建立.中国农业科 学.1996,29(3):61-65
[148]刘辉,陈纯贤.小麦×玉米杂交后代的蛋白质及酯酶同工酶分析[J].植物学报,1999,41(1):55-59.
[149]G..Brazauskas,I.Pasakinskien,A.Jahoor.AFLP analysis indicates no introgression of maize DNA in wheat×maize crosses[J].Plant Breeding 2004,123:117-121.
[150]R(e|¨)mer S,Fraser P D,2005.Recent advances in carotenoid biosynthesis,regulation and manipulation[J].Planta.211:305-308
[151]R(o|¨)mer S,Hugueney P,Bouvier F,et al.,1993.Expression of the genes encoding the early carotenoid biosynthetic enzymes in Capsicum armuum[J].Biochem Biophys Res Commun,196:1414-1421
[152]Schmimidt,Darmert C.Engineering novel carotenoids in microorganisms[J].Curr Opin Biotechnol,2000,11:255-261
[153]Zoi Karvounil,Isaac Johnl,Jane E,et al.,1995.Turnerl and Don Grierson Isolation and eharaetedsation of a melon cDNA clone encoding phytoene synthase[J].Plant Molecular Biology,27:1153-1162
[154]Burkhardt PK,Beyer P,Warm J,et al.,1997.Transgenie rice(Oryza sativa)endosperm expressing daffodil(Narcissus pseudonarcissus) phytoene synthase accumulates phytoene,a key intermediate of provitamin A biosynthesis[J].Plant Physiol,11(5):1071-1078.
[155]Faqiang Li,Ratnakar Vallabhaneni,and Eleanore T.PSY3,a New Member of the Phytoene Synthase Gene Family Conserved in the Poaceae and Regulator of Abiotic Stress-Induced Root Carotenogenesis[J].Plant Physiol,2008,146(3):1333-1345
[156]Li Y Y,Ma C Z,Fu T D,et al..Construction of a molecular functional map ofrapeseed(Brassiea napus L.) using differentially expressed genesbetween hybrid and its parents[J].Euphytiea,2006,152:25-39.
[157]Ronen G,Carmel,Goren L,Zamir D,et al.,2000.An alternative pathway to β-carotene formation in plant ehromoplasts discovered by map-based cloning of beta and old-gold color mutations in tomato[J].Proceedings of the National Academy of Sciences of the United States of America,97:11102-11107.
[158]Wong J C,Lambert R J and Wurtzel E T,2004.QTL and candidate genes phytoene synthase and carotene desaturase associated with the accumulation of carotenoids in maize[J].Theor Appl Genet,108:349-359
[159]Gale MD,Devos KM Comparative genetics in the grasses[J].Proc Natl Acad Sci USA,1998,95:1971-1974
[160]孙道杰,何中虎,王辉.小麦面粉黄色素相关基因研究[J].西北植物学报,2006,26(4):0655-0660
[161]AACC(1995) Approved Methods of the American Association of Cereal Chemists,9th end.St.Paul,MN,USA
[162]Guo M,Rupe M A,Zinselmeier C,et al.Allelic variation of gene expression in maize hybrids[J].Plant Cell,2004,16(7):1707-1716.
[163]Knott DR.Translocations involving Triticum chromosomes and Agropyron chromosomes carrying rust resistance[J].Canadian Journal of Genetics and Cytology,1968,10:695-696.
[164]Riera-Lizarazu O,Rines H W,Phillips R L.Maize chromosome retention,sexual transimission,and introgression in plants generated by oat×maize crosses[J].In:Abstract of the second international conference on the plant genome.San Diego:California,1994.178
[165]Cenci A,Somma S,Chantret N,Dubcovsky J,Blanco A.PCR identification of durum wheat BAC clones containing genes coding for carotenoid biosynthesis enzymes and their chromosome localization[J].Genome,2004,47,911-917
[166]Ruther A,Misawa N,BEger P et al.Production of zeaxanthin in Eschedchia coli transformed with different carotenogenic plasmids[J].Appl Microbiol Biotechnol,1997,48:162-167
[167]Atienza SG,Avila CM,Ramirez MC,Martin A.Application of near infrared reflectance spectroscopy to the determination of carotenoid content in tritordeum for breeding purposes[J].Australian Journal of Agricultural Research,2005,56,85-89
[168]Atienza SG,Ballesteros J,Martin A,Hornero-M'endez D.Genetic variability of carotenoid concentration and degree of esterification among tritordeum(×Tritordeum Ascherson et Graebner) and durum wheat accessions.Journal of Agricultural and Food Chemistry[J],2007,55,4244-4251
[169]李懋学,陈瑞阳.关于植物核型分析的标准化问题[J].武汉植物学研究,1985,3(4):297-302.
[170]陈瑞阳,宋文芹,李秀兰,等.中国主要经济植物基因组染色体图谱(第3册)[J].北京:科学出版社,2003:1-152.
[171]安海龙,卫志明,黄健秋.小麦幼胚培养高效成株系统的建立[J].植物生理学报,2000,26(6):532-538.
[172]宋国琦,王成社,何培茹.小麦幼胚培养技术及其应用的研究进展[J].西安联合大学学报.2003,6(2):22-27
[173]陈耀锋,曹团武,丁云姣,等.普通小麦基因转化中良好受体系统的建立[J].西北农林科技大学学报(自然科学版).2004,32(9):1-3
[174]王子宁,张艳敏,郭北海,等.利用单倍体育种技术快速培育糯性小麦新品系[J].华北农学报,2001,16(1):1-6
[175]Comeau A,Nadeau P,Ploured A.Media for the in ovule culture of proembryos of wheat and wheat-derived interspecific hybrids or haploids[J].Plant Sci,1992,81:117-125
[176]陈纯贤,孙敬三,朱立煌.由小麦×玉米获得的普通小麦加倍单倍体后代的RFLP变异[J].植物学报,1999,41(1):55-59
[177]李振声.用于小麦染色体工程的蓝粒小麦单体系列材料的创制[J].遗传,2001,23(1):42-42
© 2004-2018 中国地质图书馆版权所有 京ICP备05064691号 京公网安备11010802017129号 地址：北京市海淀区学院路29号 邮编：100083 电话：办公室：(+86 10)66554848；文献借阅、咨询服务、科技查新：66554700 |