普通小麦—华山新麦草衍生后代的细胞学鉴定和控制大豆种子低镉积累基因的克隆
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摘要
普通小麦作为人类的主要粮食作物,具有广泛的种植面积。尽管其产量和品质在过去儿百年的育种改良进程中得到了显著性地提高,但由于狭窄的遗传背景以及贫乏的遗传多样性,普通小麦也很容易受到生物或非生物的胁迫,严重阻碍了产量以及品质的进一步提升。小麦属的野生近缘属具有丰富的遗传变异,是小麦遗传改良的重要资源。通过远缘杂交将野生近缘属物种中优良的农艺性状或基因转移到普通小麦中,从而丰富普通小麦的遗传多样性以及扩大其遗传变异性,是现今小麦遗传改良的主要途径。本文以普通小麦-华山新麦草杂种衍生后代为研究材料,通过田间农艺性状考察、细胞学、基因组原位杂交和储藏蛋白亚基分析等手段对衍生后代进行了筛选和评价,主要结果如下：
1.596株BC2{[F1×CS]×CS,[F1×CSph26]×CSph26和[F1×J-11]×J-11}和BC1F1{[F1×CS]selfed,[F1×CSph2b]selfed和[F1×J-11]selfed}直株的体细胞染色体数目分布在2n=42-52,其中染色体数目为2n=45植株最多,占到19.6%。由于受到遗传因子的影响,群体染色体数目旱非正态分布。比较群体之间以及组合之间染色体数目的分布发现：染色体数目的分布受到回交亲本基因型的影响。在BC2群体中,CSph2b对染色体数日分布的影响效应最大,而CS和J-11的效应相似。然而在BC1F1群体中,J-11的影响效应大于CS。连续多代的回交能够加快染色体的丢失,不利于外源基因的重组。
2.从BC1F2和BC1F3群体中筛选和鉴定了一系列小麦-华山新麦草外源附加系。5个染色体数目为44以及1个染色体数目为46的植株细胞学上表现稳定。基于染色体配对、GiemsaC-带以及基因组原位杂交的研究,材料编号为156-4、160-12、160-13、173-2以及197-16(2n=44)是附加了一对华山新麦草5Ns染色体的双体附加系。165-2具有46条染色体,分别附加了一对华山新麦草的3Ns和5Ns染色体,是一个双二体附加系。同时也筛选到9个单体附加系,分别附加一条华山新麦草的1Ns、2Ns、4Ns或者5Ns染色体。材料241-2以及241-10是两个罗宾逊易位系。在花粉母细胞减数分裂观察过程中,检测到一些异常的染色体行为,例如：端体、落后端体、姐妹染色端体在减数分裂后期Ⅰ移向同极、末期Ⅱ时的染色体不同步分离。这些结果表明,存在的华山新麦草染色休打乱了原有的减数分裂平衡。通过对这些附加系的农艺性状考察和比较,发现华山新麦草的5Ns染色体上携带有控制小麦芒有无的基因。
3.对9个普通小麦-华山新麦草的异染色体系进行了Giemsa C-带、基因细原位杂交以及条锈病抗性的鉴定。结果表明：附加系163-5、165-1、183-5、240-3和240-3为附加了一条华山新麦草3Ns染色体的单体附加系：183-1和183-20是附加了一对华山新麦草3Ns染色体的双体附加系：165-20是附加了一对3Ns和一对4Ns染色体的双二体附加系;而219-1是分别附加了一对1Ns和3Ns染色体、而5A染色体丢失的一个双二体附加-代换系。这些含有华山新麦草3Ns染色体的附加系表现出对条锈病高抗或者免疫的特性。而含Ns、2Ns、4Ns和5Ns附加系均表现出高感小麦条锈病。这些结果强有力地揭示了华山新麦草3Ns染色体携带有对小麦条锈病高抗或免疫的基因。这些附加系是普通小麦抗病遗传改良新的重要抗性资源供体，将对小麦的条锈病抗性育种具有重要的作用。
4.利用花粉母细胞检测、Giemsa C-带SDS-PAGE以及田间抗病性鉴定从普通小麦-华山新麦草衍生后代BC1F4群体中筛选和鉴定了两个部分双二倍体系列B113(32株)和B21(13株)。15个单株全部是非整倍体,2n=50(8株)、51(6株)和54(1株)。部分植株减数分裂正常。该群体包含了全部的华山新麦草染色体,尽管不是存在于一个单株中。34株具有华山新麦草高分子量谷蛋白亚基以及41株具有华山新麦草低分子量谷蛋白亚基或者一些新的亚基。全部植株对混合条锈病生理小种‘条中30’、‘条中31’、‘水源7’和‘水源14’均表现出高抗(10株)或者免疫(35)。因而,该部分双二倍体群体将是利用华山新麦草基因资源遗传改良普通小麦的重要桥梁材料,也是保护、研究和利用华山新麦草遗传资源的重要材料。
来源于工业废料和农业磷肥的有毒重金属元素镉能够高度富集在土壤中,对各种生物有机体造成严重的毒害。富集在植物体中的镉能够引起植物体细胞的破坏,从而破坏相应的生理代谢进程,以及诱导相应的生理住化应答及其信号转导途径。镉能引起人类多种疾病,例如：肾小管功能障碍和骨质疏松等。不同大豆品种之间种子的镉积累能力不同,大部分的现有栽培大豆品种能够从镉污染的土壤中积累超过标准含量的镉在种子中。大豆长期作为人类的主要食品,能够加工成豆奶、豆腐或者大豆油。人们直接或者间接食用高镉积累的种子能够引起人类潜在的健康问题。在控制大豆种子低镉积累基因定位在大豆基因组连锁群K的Sat_119和SatK176之间的研究基础上,本文以两个大豆品种Westag97(低镉品种)和ACHime(高镉品种)为研究材料,对镉胁迫下的大豆各组织进行了用于量化目的基因表达水平的qRT-PCR内参基因表达稳定性的筛选和评价,以及克隆、表达和功能分析了控制大豆种子低镉积累的基因。主要研究结果如下：
5.利用0、0.1和1.6μmol/L的镉处理发芽第14天的两个大豆品种Westag97和AC Hime,在处理后的第2、6、12、24h分别收集根、茎和叶,提取其RNA并反转录成cDNA用于qRT-PCR分析。通用的两个内参基因评估软件geNrom和NormFinder均表明所选的10个候选基因中,ACT3、PP2A、ELF IB和F-box在该实验条件下是最稳定的四个内参基因,可以用作后续基因表达的标准量化研究。而G6PD、UBC2、TUB和ELF1A是最不稳定的四个候选基因,不能在该实验条件下用作后续基因表达的研究。
6.对候选基因区域内编码重金属元素转移蛋白的大豆重金属元素结合蛋白(GmHMA3)基因进行了克隆,表达和功能上的分析。GmHMA3a和GmHMA3w的序列一致,除了GmHMA3a在1823by处发生了单碱基的代换。20个现今流行的栽培品种中,所有的高镉品种均具有该单核苷酸多态性。在此基础上,建立了一个HRM标记,该标记与低镉性状紧密连锁且其遗传距离为0.3cM。标记和性状连锁分析表明GmHMA3w属于野生型基因。表达结果表明GmHMA3只在根部表达,且高、低镉品种之间没有表达上的差异。基因功能分析表明GmHMA3w具有转移和运输镉和锌的能力,它将根尖吸附的镉或者锌储藏到根部的液泡膜细胞或者其它根部细胞中,而阻止它们通过木质部向地上组织的转运,从而减少了大豆种了中镉的积累。
7.部分金属元素转运蛋白能够转运多种金属元素。通过本研究发现,GmHMA3w除了转运镉和锌外,对铅、铁和钻都没有转运能力。通过含有GFP的重组载体转化,发现GmHMA3在酵母细胞的内质网上表达,从而解释了为什么含有pYES2.1-GmHMA3w的酵母菌株在镉和锌的培养基中表现出敏感性,而不是耐受性。
Bread wheat (Triticum aestivum L.,2n=6x-42AABBDD) as staple food of the world's population, is the most widely grown crop in the world. Although it has been bred intensively for hundred of years, significant improvements in yield and quality have been achieved. Due to its limited genetic diversity, bread wheat is constantly threatened by various abiotic and biotic factors, which has limited its improvements in yield and quality. The wild relatives of wheat possess a large number of useful genes, which will be the genetic donor of wheat breeding. An efficient strategy to broaden the genetic base of wheat is to introgress some useful genes of its wild relatives in the tertiary gene pool into wheat through distant crosses. In the present study, we selected and characterized some alien chromosome lines derived from wheat-Psathyrostachys huashanica Keng ex Kuo (2n=2x=14, NsNs) by agronomy, cytogenetics, GISH and SDS-PAGE. The results are as follows:
1.596plants of the BC2{[F1×CS]×CS,[F1×CSph2b]×CSph2b and [F1×J-11]×J-11} and BC1F1{[F1×CS] selfed,[F1×CSph2b] selfed and [F1×J-11] selfed} were observed for analyzing the distribution of chromosome numbers. The chromosome numbers of the populations of BC2and BC1F1varied from2n=42to52, and plants with2n=45accounted for the highest frequency (19.6%). The distribution of chromosome numbers in all combinations didn't fit to normal distribution, indicating that the distribution of chromosome numbers was affected by some genetic factors. There were different main ranges of chromosome numbers and reduced chromosome numbers per a hundred of plants in BC2accessions and BC1F1accessions. These results indicated that the distribution of chromosome numbers was affected by the genotype of the backcrossing parents. The effect of CSph2b higher than CS and J-11, and the effect of CS and J-11were similar in BC2accessions. However, the efficiency of J-11was higher than CS in BC1F1accessions. And chromosome eliminated more rapidly by backcrossing than selfing, and selfing may be facilitated to the screening of recombination between wheat and P. huashanica chromosomes.
2. Alien chromosome lines were developed and identified from the BC1F2and BC1F3generations. Five lines with2n=44and one line with2n=46showed regular meiosis and were cytologically stable. Based on chromosome pairing, C-banding and GISH analysis, lines156-4,160-12,160-13,173-2and197-16(2n=44) were alien disomic addition lines with the addition of one pair of the P. huashanica5Ns chromosomes. Line165-2, which had each pair of the P. huashanica chromosomes3Ns and5Ns being added, was an alien double addition line. Nine lines with the P. huashanica chromosome1Ns,2Ns,4Ns or5Ns were alien monosomic addition lines. Robertsonian translocation was observed in lines241-2and241-10. Some chromosomal abnormalities were observed, such as telosomics, lagging of telosomics, and telosomic sister chromatids that moved to one pole at anaphase I and asynchronous chromosome separation at telophase Ⅱ. These results indicated that the presence of alien chromosomes from P. huashanica had influenced the meiosis. Meanwhile, by comparing the series of wheat-P. huashanica chromosome addition lines, the gene(s) for awns were mapped to the P. huashanica5Ns chromosome.
3. Nine wheat-P.huashanica addition lines were characterized by Giemsa C-banding, genomic in situ hybridization (GISH) and disease resistance evaluation. Giemsa C-banding and GISH demonstrated that lines163-5,165-1,183-5,240-3and240-4are P. huashanica3Ns chromosome monosomic addition lines; lines183-1and183-20are P. huashanica3Ns chromosome disomic addition lines; line165-20is P. huashanica3Ns and4Ns chromosome double disomic addition lines; and line219-1is P. huashanica INs and3Ns/5A chromosome double disomic addition-substitution lines. All of these addition lines with P. huashanica3Ns chromosome(s) expressed high resistance or immunity to stripe rust. By comparing the series of whea1-P. huashanica chromosomes addition lines, we concluded that the P. huashanica3Ns chromosome carries the gene(s) for resistance or immunity to stripe rust. These addition lines can be used as a donor source of novel stripe rust resistance to wheat breeding programs.
4. Two partial amphiploid lines, B113(32plants) and B21(13plants) which were BC1F4, were characterized by Giemsa C-banding and SDS-PAGE and evaluated for stripe rust resistance. All15partial amphiploid plants analysed were aneuploids with either50(8plants),51(6plants) or54(I plant) chromosomes. Some of them showed regular meiosis and all the P. huashanica chromosomes were included, although not in a single plant. SDS-PAGE analysis of the45plants showed that34expressed some specific bands representing high molecular weight glutenin subunit (HMW-GS) and41had bands representing P. huashanica low molecular weight glutenin subunit (LMW-GS), including two new subunits. All45plants were highly resistant (10) or immune (35) to stripe rust mixed races CYR-30, CYR-31, Shuiyuan7and Shuiyuan14. These amphiploid plants could be useful germplasm for enhancing stripe rust resistance and might improve wheat grain quality.
Cadmium (Cd) is a toxic trace pollutant for all living organisms that can accumulate to high concentrations in soil from industrial processes and phosphate fertilizers. It can accumulate in plants causing cellular damage, leading to a disruption of physiological processes, and them inducing the responses at the biochemical-physiological level and on signal transduction pathways. Cd has been recognized as a contributing factor to human diseases, such as renal proximal tubular dysfunction and the bone disease called itai-itai. There are cultivar differences in seed Cd accumulation in soybean [Glycine Max (L.) Merr] and many cultivars accumulate high Cd concentrations in seed when grown on Cd-contaminated soils. These cultivars can exceed the proposed Codex upper limit. Soybean has long been a staple food for humans, especially as soymilk, tofu and oil. Consumption, either directly or indirectly, of seed with high levels of Cd could be a human health concern. In the present study, based on a major gene or QTL controlling seed Cd accumulation in soybean has been located on linkage group K between the SSR markers Sat119and SatK176, two cultivars, Westag97(a low Cd accumulator) and AC Hime (a high Cd accumulator), were used to evaluate reference genes for normalization of qRT-PCR analysis of differentially expressed genes in soybean exposed to Cd, and clone, expressed and functionally analyze GmHMA3w limiting Cd translocation in soybean. The results are as follows:
5. Due to no study has concentrated on identifying multiple reference genes for normalization of qRT-PCR data in soybean under exposure to heavy metal stress treatments, we evaluated the expression stability of ten candidate housekeeping genes in leaves, roots and stems of two soybean cultivars exposed to increased Cd concentrations. Under evaluation of geNorm' and NormFinder, ACT3, PP2A, ELF1B and F-box were the four best reference genes, based on geNorm and NormFinder analysis, for these gene expression studies. These four genes also may be suitable for normalization of gene expression in soybean under exposure to other heavy metals which have similar properties and effects as Cd. Both geNorm and NormFinder analyses revealed that G6PD, UBC2, TUB, and ELF1A could not be used as reference genes for normalization in these experimental conditions.
6. A candidate gene, Glycine max heavy metal associated protein3(GmHMA3), which encodes a heavy metal transporter belonging to the P1B-type ATPase family, was selected for analysis. Sequence analysis of20diverse soybean cultivars detected a single nucleotide change in this gene in all high seed Cd accumulators. A single nucleotide polymorphism (SNP) marker was developed and used to genotype166F8RILs by high resolution melt (HRM) analysis. Segregation analysis mapped the GmHMA3to0.3cM away from the Cdal gene previously tagged by the SSR markers, SatK147, SacK149and Saatk150. Marker-trait association analysis also indicated that the wild type allele, GmHMA3w, is tightly associated with low seed Cd concentration. Expression level studies revealed that GmHMA3is only expressed in root. Functional assay of the GmHMA3gene in yeast indicated that GmHMA3w encodes a Cd/Zn transporter, which sequesters Cd and Zn into vacuoles or other place in roots, thereby limiting the Cd and Zn accumulations in soybean seed.
7. Some metal transporters of plant can transport multi-metals. Although GmHMA3w is a Cd and Zn transporter, it can not transport Fe, Pb and Co. Due to GmHMA3was mis-localized on endoplasmic reticulum in yeast, GmHMA3w increased the sensitive of Cd and Zn.
1.596株BC2{[F1×CS]×CS,[F1×CSph26]×CSph26和[F1×J-11]×J-11}和BC1F1{[F1×CS]selfed,[F1×CSph2b]selfed和[F1×J-11]selfed}直株的体细胞染色体数目分布在2n=42-52,其中染色体数目为2n=45植株最多,占到19.6%。由于受到遗传因子的影响,群体染色体数目旱非正态分布。比较群体之间以及组合之间染色体数目的分布发现：染色体数目的分布受到回交亲本基因型的影响。在BC2群体中,CSph2b对染色体数日分布的影响效应最大,而CS和J-11的效应相似。然而在BC1F1群体中,J-11的影响效应大于CS。连续多代的回交能够加快染色体的丢失,不利于外源基因的重组。
2.从BC1F2和BC1F3群体中筛选和鉴定了一系列小麦-华山新麦草外源附加系。5个染色体数目为44以及1个染色体数目为46的植株细胞学上表现稳定。基于染色体配对、GiemsaC-带以及基因组原位杂交的研究,材料编号为156-4、160-12、160-13、173-2以及197-16(2n=44)是附加了一对华山新麦草5Ns染色体的双体附加系。165-2具有46条染色体,分别附加了一对华山新麦草的3Ns和5Ns染色体,是一个双二体附加系。同时也筛选到9个单体附加系,分别附加一条华山新麦草的1Ns、2Ns、4Ns或者5Ns染色体。材料241-2以及241-10是两个罗宾逊易位系。在花粉母细胞减数分裂观察过程中,检测到一些异常的染色体行为,例如：端体、落后端体、姐妹染色端体在减数分裂后期Ⅰ移向同极、末期Ⅱ时的染色体不同步分离。这些结果表明,存在的华山新麦草染色休打乱了原有的减数分裂平衡。通过对这些附加系的农艺性状考察和比较,发现华山新麦草的5Ns染色体上携带有控制小麦芒有无的基因。
3.对9个普通小麦-华山新麦草的异染色体系进行了Giemsa C-带、基因细原位杂交以及条锈病抗性的鉴定。结果表明：附加系163-5、165-1、183-5、240-3和240-3为附加了一条华山新麦草3Ns染色体的单体附加系：183-1和183-20是附加了一对华山新麦草3Ns染色体的双体附加系：165-20是附加了一对3Ns和一对4Ns染色体的双二体附加系;而219-1是分别附加了一对1Ns和3Ns染色体、而5A染色体丢失的一个双二体附加-代换系。这些含有华山新麦草3Ns染色体的附加系表现出对条锈病高抗或者免疫的特性。而含Ns、2Ns、4Ns和5Ns附加系均表现出高感小麦条锈病。这些结果强有力地揭示了华山新麦草3Ns染色体携带有对小麦条锈病高抗或免疫的基因。这些附加系是普通小麦抗病遗传改良新的重要抗性资源供体，将对小麦的条锈病抗性育种具有重要的作用。
4.利用花粉母细胞检测、Giemsa C-带SDS-PAGE以及田间抗病性鉴定从普通小麦-华山新麦草衍生后代BC1F4群体中筛选和鉴定了两个部分双二倍体系列B113(32株)和B21(13株)。15个单株全部是非整倍体,2n=50(8株)、51(6株)和54(1株)。部分植株减数分裂正常。该群体包含了全部的华山新麦草染色体,尽管不是存在于一个单株中。34株具有华山新麦草高分子量谷蛋白亚基以及41株具有华山新麦草低分子量谷蛋白亚基或者一些新的亚基。全部植株对混合条锈病生理小种‘条中30’、‘条中31’、‘水源7’和‘水源14’均表现出高抗(10株)或者免疫(35)。因而,该部分双二倍体群体将是利用华山新麦草基因资源遗传改良普通小麦的重要桥梁材料,也是保护、研究和利用华山新麦草遗传资源的重要材料。
来源于工业废料和农业磷肥的有毒重金属元素镉能够高度富集在土壤中,对各种生物有机体造成严重的毒害。富集在植物体中的镉能够引起植物体细胞的破坏,从而破坏相应的生理代谢进程,以及诱导相应的生理住化应答及其信号转导途径。镉能引起人类多种疾病,例如：肾小管功能障碍和骨质疏松等。不同大豆品种之间种子的镉积累能力不同,大部分的现有栽培大豆品种能够从镉污染的土壤中积累超过标准含量的镉在种子中。大豆长期作为人类的主要食品,能够加工成豆奶、豆腐或者大豆油。人们直接或者间接食用高镉积累的种子能够引起人类潜在的健康问题。在控制大豆种子低镉积累基因定位在大豆基因组连锁群K的Sat_119和SatK176之间的研究基础上,本文以两个大豆品种Westag97(低镉品种)和ACHime(高镉品种)为研究材料,对镉胁迫下的大豆各组织进行了用于量化目的基因表达水平的qRT-PCR内参基因表达稳定性的筛选和评价,以及克隆、表达和功能分析了控制大豆种子低镉积累的基因。主要研究结果如下：
5.利用0、0.1和1.6μmol/L的镉处理发芽第14天的两个大豆品种Westag97和AC Hime,在处理后的第2、6、12、24h分别收集根、茎和叶,提取其RNA并反转录成cDNA用于qRT-PCR分析。通用的两个内参基因评估软件geNrom和NormFinder均表明所选的10个候选基因中,ACT3、PP2A、ELF IB和F-box在该实验条件下是最稳定的四个内参基因,可以用作后续基因表达的标准量化研究。而G6PD、UBC2、TUB和ELF1A是最不稳定的四个候选基因,不能在该实验条件下用作后续基因表达的研究。
6.对候选基因区域内编码重金属元素转移蛋白的大豆重金属元素结合蛋白(GmHMA3)基因进行了克隆,表达和功能上的分析。GmHMA3a和GmHMA3w的序列一致,除了GmHMA3a在1823by处发生了单碱基的代换。20个现今流行的栽培品种中,所有的高镉品种均具有该单核苷酸多态性。在此基础上,建立了一个HRM标记,该标记与低镉性状紧密连锁且其遗传距离为0.3cM。标记和性状连锁分析表明GmHMA3w属于野生型基因。表达结果表明GmHMA3只在根部表达,且高、低镉品种之间没有表达上的差异。基因功能分析表明GmHMA3w具有转移和运输镉和锌的能力,它将根尖吸附的镉或者锌储藏到根部的液泡膜细胞或者其它根部细胞中,而阻止它们通过木质部向地上组织的转运,从而减少了大豆种了中镉的积累。
7.部分金属元素转运蛋白能够转运多种金属元素。通过本研究发现,GmHMA3w除了转运镉和锌外,对铅、铁和钻都没有转运能力。通过含有GFP的重组载体转化,发现GmHMA3在酵母细胞的内质网上表达,从而解释了为什么含有pYES2.1-GmHMA3w的酵母菌株在镉和锌的培养基中表现出敏感性,而不是耐受性。
Bread wheat (Triticum aestivum L.,2n=6x-42AABBDD) as staple food of the world's population, is the most widely grown crop in the world. Although it has been bred intensively for hundred of years, significant improvements in yield and quality have been achieved. Due to its limited genetic diversity, bread wheat is constantly threatened by various abiotic and biotic factors, which has limited its improvements in yield and quality. The wild relatives of wheat possess a large number of useful genes, which will be the genetic donor of wheat breeding. An efficient strategy to broaden the genetic base of wheat is to introgress some useful genes of its wild relatives in the tertiary gene pool into wheat through distant crosses. In the present study, we selected and characterized some alien chromosome lines derived from wheat-Psathyrostachys huashanica Keng ex Kuo (2n=2x=14, NsNs) by agronomy, cytogenetics, GISH and SDS-PAGE. The results are as follows:
1.596plants of the BC2{[F1×CS]×CS,[F1×CSph2b]×CSph2b and [F1×J-11]×J-11} and BC1F1{[F1×CS] selfed,[F1×CSph2b] selfed and [F1×J-11] selfed} were observed for analyzing the distribution of chromosome numbers. The chromosome numbers of the populations of BC2and BC1F1varied from2n=42to52, and plants with2n=45accounted for the highest frequency (19.6%). The distribution of chromosome numbers in all combinations didn't fit to normal distribution, indicating that the distribution of chromosome numbers was affected by some genetic factors. There were different main ranges of chromosome numbers and reduced chromosome numbers per a hundred of plants in BC2accessions and BC1F1accessions. These results indicated that the distribution of chromosome numbers was affected by the genotype of the backcrossing parents. The effect of CSph2b higher than CS and J-11, and the effect of CS and J-11were similar in BC2accessions. However, the efficiency of J-11was higher than CS in BC1F1accessions. And chromosome eliminated more rapidly by backcrossing than selfing, and selfing may be facilitated to the screening of recombination between wheat and P. huashanica chromosomes.
2. Alien chromosome lines were developed and identified from the BC1F2and BC1F3generations. Five lines with2n=44and one line with2n=46showed regular meiosis and were cytologically stable. Based on chromosome pairing, C-banding and GISH analysis, lines156-4,160-12,160-13,173-2and197-16(2n=44) were alien disomic addition lines with the addition of one pair of the P. huashanica5Ns chromosomes. Line165-2, which had each pair of the P. huashanica chromosomes3Ns and5Ns being added, was an alien double addition line. Nine lines with the P. huashanica chromosome1Ns,2Ns,4Ns or5Ns were alien monosomic addition lines. Robertsonian translocation was observed in lines241-2and241-10. Some chromosomal abnormalities were observed, such as telosomics, lagging of telosomics, and telosomic sister chromatids that moved to one pole at anaphase I and asynchronous chromosome separation at telophase Ⅱ. These results indicated that the presence of alien chromosomes from P. huashanica had influenced the meiosis. Meanwhile, by comparing the series of wheat-P. huashanica chromosome addition lines, the gene(s) for awns were mapped to the P. huashanica5Ns chromosome.
3. Nine wheat-P.huashanica addition lines were characterized by Giemsa C-banding, genomic in situ hybridization (GISH) and disease resistance evaluation. Giemsa C-banding and GISH demonstrated that lines163-5,165-1,183-5,240-3and240-4are P. huashanica3Ns chromosome monosomic addition lines; lines183-1and183-20are P. huashanica3Ns chromosome disomic addition lines; line165-20is P. huashanica3Ns and4Ns chromosome double disomic addition lines; and line219-1is P. huashanica INs and3Ns/5A chromosome double disomic addition-substitution lines. All of these addition lines with P. huashanica3Ns chromosome(s) expressed high resistance or immunity to stripe rust. By comparing the series of whea1-P. huashanica chromosomes addition lines, we concluded that the P. huashanica3Ns chromosome carries the gene(s) for resistance or immunity to stripe rust. These addition lines can be used as a donor source of novel stripe rust resistance to wheat breeding programs.
4. Two partial amphiploid lines, B113(32plants) and B21(13plants) which were BC1F4, were characterized by Giemsa C-banding and SDS-PAGE and evaluated for stripe rust resistance. All15partial amphiploid plants analysed were aneuploids with either50(8plants),51(6plants) or54(I plant) chromosomes. Some of them showed regular meiosis and all the P. huashanica chromosomes were included, although not in a single plant. SDS-PAGE analysis of the45plants showed that34expressed some specific bands representing high molecular weight glutenin subunit (HMW-GS) and41had bands representing P. huashanica low molecular weight glutenin subunit (LMW-GS), including two new subunits. All45plants were highly resistant (10) or immune (35) to stripe rust mixed races CYR-30, CYR-31, Shuiyuan7and Shuiyuan14. These amphiploid plants could be useful germplasm for enhancing stripe rust resistance and might improve wheat grain quality.
Cadmium (Cd) is a toxic trace pollutant for all living organisms that can accumulate to high concentrations in soil from industrial processes and phosphate fertilizers. It can accumulate in plants causing cellular damage, leading to a disruption of physiological processes, and them inducing the responses at the biochemical-physiological level and on signal transduction pathways. Cd has been recognized as a contributing factor to human diseases, such as renal proximal tubular dysfunction and the bone disease called itai-itai. There are cultivar differences in seed Cd accumulation in soybean [Glycine Max (L.) Merr] and many cultivars accumulate high Cd concentrations in seed when grown on Cd-contaminated soils. These cultivars can exceed the proposed Codex upper limit. Soybean has long been a staple food for humans, especially as soymilk, tofu and oil. Consumption, either directly or indirectly, of seed with high levels of Cd could be a human health concern. In the present study, based on a major gene or QTL controlling seed Cd accumulation in soybean has been located on linkage group K between the SSR markers Sat119and SatK176, two cultivars, Westag97(a low Cd accumulator) and AC Hime (a high Cd accumulator), were used to evaluate reference genes for normalization of qRT-PCR analysis of differentially expressed genes in soybean exposed to Cd, and clone, expressed and functionally analyze GmHMA3w limiting Cd translocation in soybean. The results are as follows:
5. Due to no study has concentrated on identifying multiple reference genes for normalization of qRT-PCR data in soybean under exposure to heavy metal stress treatments, we evaluated the expression stability of ten candidate housekeeping genes in leaves, roots and stems of two soybean cultivars exposed to increased Cd concentrations. Under evaluation of geNorm' and NormFinder, ACT3, PP2A, ELF1B and F-box were the four best reference genes, based on geNorm and NormFinder analysis, for these gene expression studies. These four genes also may be suitable for normalization of gene expression in soybean under exposure to other heavy metals which have similar properties and effects as Cd. Both geNorm and NormFinder analyses revealed that G6PD, UBC2, TUB, and ELF1A could not be used as reference genes for normalization in these experimental conditions.
6. A candidate gene, Glycine max heavy metal associated protein3(GmHMA3), which encodes a heavy metal transporter belonging to the P1B-type ATPase family, was selected for analysis. Sequence analysis of20diverse soybean cultivars detected a single nucleotide change in this gene in all high seed Cd accumulators. A single nucleotide polymorphism (SNP) marker was developed and used to genotype166F8RILs by high resolution melt (HRM) analysis. Segregation analysis mapped the GmHMA3to0.3cM away from the Cdal gene previously tagged by the SSR markers, SatK147, SacK149and Saatk150. Marker-trait association analysis also indicated that the wild type allele, GmHMA3w, is tightly associated with low seed Cd concentration. Expression level studies revealed that GmHMA3is only expressed in root. Functional assay of the GmHMA3gene in yeast indicated that GmHMA3w encodes a Cd/Zn transporter, which sequesters Cd and Zn into vacuoles or other place in roots, thereby limiting the Cd and Zn accumulations in soybean seed.
7. Some metal transporters of plant can transport multi-metals. Although GmHMA3w is a Cd and Zn transporter, it can not transport Fe, Pb and Co. Due to GmHMA3was mis-localized on endoplasmic reticulum in yeast, GmHMA3w increased the sensitive of Cd and Zn.
引文
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