西藏野生大麦耐盐种质的发掘及其耐盐机制研究
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摘要
土壤盐化是全球范围内限制作物生产的主要逆境,由于环境污染加剧和农用灌溉水质持续恶化,农田土壤盐化日趋严重。明确作物的耐盐机理是培育耐盐新品种和制订抗盐栽培措施的基础。目前虽已明确渗透调节、离子平衡和抗氧化是植物的主要耐盐机制,但相关的遗传及分子机理尚未完全阐明。
大麦(Hordeum vulgare)是全球第四大粮食作物,适应性广,耐盐性强。以大麦为模式作物,研究其耐盐机制,对揭示禾本科作物的耐盐机理具有重要的理论价值。青藏高原一年生野生大麦(H. Spontaneum和H. agriocrithum)(以下称西藏野生大麦)系我国独有的植物种质资源,具有丰富的遗传变异和抗逆基因资源,发掘其耐盐资源可为大麦及作物耐盐改良提供优异种质。
本研究在评价西藏野生大麦耐盐性的基础上,采用关联分析法鉴定与发掘优异耐盐野生种质,并首次从离子组、膜蛋白组和代谢组三个水平上系统解析野生大麦和栽培大麦对盐胁迫的响应机制及其差异,取得的主要研究结果如下：
1.西藏野生大麦耐盐种质的发掘与遗传关联分析
采用水培试验,大麦三叶期用300mM NaCl胁迫处理3周,以相对干物重为指标,评价了188份西藏野生大麦的耐盐性,发现盐胁迫明显降低根和地上部的干重,降幅为27.6%～73.1%,且以根的降幅较大。西藏野生大麦的耐盐性存在广泛的基因型差异,与普遍认同的耐盐对照品种CM72相比,鉴定到根和地上部相对干重显著大于对照的野生大麦材料。
利用57对多态性芯片技术(DArT)标记分析了西藏野生大麦群体第5染色体的连锁不平衡(LD)结构,发现第5染色体LD的遗传衰减距离为8.9cM(R2<0.1)或者1.5cM (R2<0.2),LD程度较弱。采用测序法分析了野生大麦群体中HvCBF1、HvCBF3、HvCBF4和HvHVA1基因编码区(CDS)序列多态性,与参照序列相比,分别检测到2、15、16和10个单核苷酸多态性(SNP)位点,即平均每100bp碱基中存在0.3、2、2.4和1.6个SNP,对应3、8、13和6个单倍型,基因SNP位点之间存在不同程度的LD关系。
耐盐性状与遗传多态性的关联分析结果表明,标记bPb-4891与地上部和整株的相对干物重显著关联(P<0.05),可分别解释表型变异的2.2%和2.3%；4个基因中,仅HvCBF4的单倍型13与地上部、根和整株的相对干物重显著关联,可分别解释7.44%、2.4%和5.34%的表型变异。标记bPb-4891与HvCBF4基因在大麦遗传图谱上紧密连锁,推测该位点和HvCBF4的单倍型13可能与大麦耐盐性相关。野生大麦种质XZ16和XZ26具有HvCBF4的单倍型13的基因型,具有较强的耐盐能力。
2.大麦离子组对盐胁迫的响应机制
以栽培品种CM72(耐盐)、Gairdner(敏感)和两个野生大麦XZ16(耐盐)和XZ169(敏感)为材料,苗期进行150和300mM NaCl处理0、1、2、3和5周,系统分析了它们的生长和离子组对盐胁迫的响应。结果表明,低盐浓度对大麦生长的抑制不显著,而高盐浓度对4个大麦地上部的抑制率为30～53%,XZ16在两种盐胁迫下均有最大的绝对干物重,低浓度和高浓度下分别是CM72干物重的1.7倍和1.4倍。结果佐证了XZ16是生长快的耐盐材料,而Gairdner和XZ169的耐盐性弱。
根离子组中,盐胁迫下Na含量上升至处理2周后有所降低；P、S、Mg、K、 Mn、Zn和B含量因盐处理而降低,但Mn和Zn在高盐胁迫下降幅较低,Ca、Cu和Fe的含量因盐处理而增加。地上部离子组中,盐处理增加Na含量,降低P、K、Ca、Mg、S、Cu和B含量；Mn和Zn含量在低盐浓度下增加,而在高盐浓度下降低；而Fe含量在低盐下降低,高盐下增加。处理5周后,CM72和XZ16的Na含量和Na/K比显著低于Gairdner和XZ169。以上结果说明维持地上部低Na含量和Na/K比是大麦重要的耐盐机制。此外,大麦通过增加根部Ca、Cu和Fe的含量以及地上部Fe的含量,可增强组织对盐胁迫的耐性。
3.大麦膜蛋白对盐胁迫的响应机制
采用双向电泳(2-DE)和基质辅助激光解离飞行时间质谱系统(MALDI-TOF-TOF-MS)技术,分析了200mM NaCl胁迫处理48h和正常条件下XZ16和CM72大麦根和叶片膜蛋白组的差异,结果发现有41个膜蛋白受盐胁迫诱导差异表达。在根部,ATP合成酶(α亚基)、天冬氨酸转氨酶、单脱氢抗坏血还原酶、第Ⅲ类过氧化物酶和病程相关蛋白10受盐胁迫诱导表达上调,而ATP合成酶(β亚基)、二氢硫辛酰胺脱氢酶家族、丝氨酸蛋白酶、半胱氨酸蛋白酶、钙调蛋白类、NADP-苹果酸酶和第Ⅲ类过氧化物酶受盐胁迫表达下调。叶片中,叶绿体ATP合成酶、Bp2A蛋白、Cp31AHv蛋白、Cp31BHv蛋白、谷氨酰胺合成酶、磷酸甘油酸激酶、二磷酸腺苷葡萄糖焦磷酸酶和叶绿体Oxygen-evolving增强蛋白2受盐胁迫表达下调；而NAD依赖型异构酶、DegP蛋白酶和Harpin结合蛋白受盐胁迫诱导表达上调。
研究结果推测,根部膜蛋白的主要响应机制是保护细胞膜稳定性、清除活性氧自由基(ROS)和参与离子平衡的重建,叶片膜蛋白的主要响应机制是保护光合反应活性。膜蛋白的表达水平基因型之间存在差异,XZ16野生大麦参与渗透调节和离子转运以及保护叶片的光合作用的膜蛋白表达高于栽培大麦CM72,这可能也是前者具有较强耐盐性的分子基础。
4.大麦代谢物组对盐胁迫的响应机制
采用气相色谱-质谱(GC-MS)技术,分析了300mM NaCl胁迫处理3周和正常条件下XZ16和CM72大麦根和叶片代谢组差异,鉴定到82种代谢物响应盐胁迫。与对照相比,CM72和XZ16分别有53种和55种代谢物含量显著变化,根代谢组的响应方式是参与TCA循环和糖转运的代谢物积累增加,而参与糖酵解和氨基酸合成的代谢物积累受到抑制。叶片代谢组中,CM72和XZ16分别有55种和54种代谢物含量显著变化,其响应方式是参与糖酵解、Calvin循环和氨基酸合成的代谢物积累明显增加,而参与TCA循环的代谢物积累受到抑制。
代谢物响应盐胁迫存在组织差异性,根部调节渗透的主要代谢物是脯氨酸、糖类(蔗糖、海藻糖和棉子糖)、甘露醇和纤维醇；而棉子糖和脯氨酸等多种氨基酸是叶片的主要渗透调节物。根部代谢物对盐胁迫的响应机制是积累渗透调节剂含量,增强耐盐性；而叶片代谢物的响应机制除增强组织的渗透调节能力外,还为根部提供足够的糖类和能量,从而调节渗透压和转运Na+。代谢物响应机制存在基因型差异,栽培大麦CM72通过增加代谢物的合成,从而增强渗透调节和离子平衡提供物质和能量；而野生大麦XZ16自身拥有较强的渗透调节能力,维持正常的光合作用和合成代谢,从而保持较高的生长水平。
综上所述,大麦的耐盐性受一个复杂的分子调控网络控制。本研究从西藏野生大麦中鉴定到耐盐性优于CM72的种质,可为大麦耐盐育种提供遗传材料；并从离子组、膜蛋白组和代谢组上分别解析了野生大麦和栽培大麦对盐胁迫的响应及其差异,丰富了作物耐盐机理,可为进一步发掘耐盐遗传资源和培育耐盐作物品种提供理论参考和实践指导。
Soil salinity is a major abiotic stress of restricting crop productivity worldwide, posing a great threat to agricultural sustainability. A thorough understanding of salt-tolerant mechanisms is necessary for alleviating salt injury to agricultural production by improving cultural practices and developing salt-tolerant varieties. Currently, it is well documented that osmotic regulation, ion homeostasis and anti-oxidation are the three aspects of salt tolerance in plants. However, the genetic and molecular mechanisms of salt tolerance have not been fully elucidated.
Barley (Hordeum vulgare) is the fourth most important cereal crop in the world, characterized by its wide adaptability and high salinity tolerance. Consequently, it is frequently used as a model crop in the attempts to understand salinity tolerance in the cereal crops. Tibetan annual wild barley (H. vulgare ssp. Spontaneum and ssp. agriocrithum)(hereafter referred to as Tibetan wild barley) is considered as one of the ancestors of cultivated barley, and shows wide genetic variations in the tolerance to harsh environments. It is possible for us to explore the elite germplasm in terms of salt tolerance from the wild barley.
In the present study, an association analysis was used to identify and explore salt-tolerant germplasm based on the evaluation of salt tolerance among around200Tibetan wild barley accessions, and the mechanisms of salt tolerance between wild and cultivated barleys were systematically resolved by ionomic, membrane proteomic and metabolomic methods. The main results were summarized as follows:
1. Identification of salt-tolerant germplasm and genetic association analysis within Tibetan wild barleys.
Hundred and eighty-eight accessions of Tibetan wild barley were exposed to salt stress of300mM NaCl in hydroponics at three-leaf stage for3weeks, to compare the difference among these accessions in salt tolerance as indicated by relative dry weights. Salt stress significantly reduced shoot and root dry weight by27.6%to73.1%and the root weight showed greater reduction than the shoot weight. There was a significant difference among the Tibetan wild barley accessions. Compared with CM72, a salt-tolerant cultivated cultivar, some accessions showed higher shoot, root or whole-plant dry weight under salt stress.
The linkage disequilibrium (LD) structures of the chromosome5were evaluated using57diversity arrays technology (DArT) markers over this chromosome. The LD decay of genetic distance was8.9cM (R2<0.1) or1.5cM (R2<0.2). The genetic variation of HvCBF1, HvCBF3, HvCBF4and HvHVA1was studied by sequencing the gene coding regions. There were2,15,16and10single nucleotide polymorphisms (SNP) sites, corresponding to3,8,13and6haplotypes for the four genes, respectively, with0.3,2,2.4and1.6SNPs in each100bp of the coding sequence. Furthermore, the certain LD relationship was detected between SNPs.
Association analysis was conducted between the genetic diversity and the genotypic salt tolerance. It was found that marker bpb-4891was significantly associated with salt tolerance, explaining the2.2%and2.3%of phenotypic variation for the relative shoot and whole-plant dry weight, respectively. The haplotype13of HvCBF4gene exhibited highly significant association with shoot and whole-plant relative dry weight, explaining7.7%and6.4%of the variations, respectively. Based on the distance in the barley genetic map, it may be assumed that the marker bpb-4891was closely linked with the HvCBF4gene. The results suggest that the marker bpb-4891and haplotype13of HvCBF4gene could be related to salinity tolerance. Tibetan wild barley accessions, XZ16and XZ26have haplotype13of HvCBF4genotypes, and could be considered as high salt tolerance.
2. Mechanisms of ionome responding to salt stress in barley
In order to reveal mechanisms of ionome responding to salt stress in barley, two cultivated barleys (CM72, salt-tolerant; Gairdner, salt-sensitive) and two wild barleys (XZ16, salt-tolerant; XZ169, salt-sensitive) differing in salt tolerance were used to investigate shoot biomass and ion changes in response to salt stress of150and300mM NaCl in hydroponics at0,1,2,3and5weeks after treatment. After35days treatment, there was no significant difference in shoot weight between salt stress of150mM NaCl and control for4barley genotypes, but under salt stress of300mM NaCl, the reduction of shoot weight could be found, ranging from30%to53%. Among the four genotypes, XZ16maintained the largest shoot biomass under salt stress, and it had1.7and1.4times larger shoots dry weight than CM72, respectively. In conclusion, the results show that XZ16is a fast-growing and salt-tolerant barley.
In roots, Na content reached the maximum at2weeks after treatments, and then tended to decrease. The contents of P, S, Mg, K, Mn, Zn and B decreased steadily during salt treatment, while Mn and Zn contents decreased greatly under salt stress of300mM NaCl. By contrast, the contents of Ca, Cu and Fe increased under salt stress. In shoots, Na content increased and the contents of P, K, Ca, Mg, S, Cu and B decreased due to salt treatments. The contents of Mn and Zn increased under salt stress of150mM NaCl, and increased under salt stress of300mM NaCl. In contrast, Fe content decreased under salt stress of150mM NaCl, but increased under salt stress of300mM NaCl. These results suggest that maintenance of low Na content and Na/K ratio in shoots is a key important for salt tolerance in both cultivated and wild barleys. In addition, the increased contents of Ca, Cu and Fe in roots and Fe in shoots may enhance tissues tolerance to salt stress.
3. Mechanisms of membrane proteins responding to salt stress in barley.
In order to reveal mechanisms of membrane proteins responding to salt stress in barley, wild barley (XZ16) and cultivated barley (CM72) were used to investigate differential expression of membrane proteomes in roots and leaves under normal condition and salt stress of200mM NaCl after48hours, using two-dimensional electrophoresis (2-DE) and matrix-assisted laser desorption/ionization time of flight mass spectrometry (MALDI-TOF-TOF-MS) techniques.
In roots, salt stress induced6proteins up-expressed, being more than1.5times than those in the control, and these proteins included ATP synthase (subunit a), aspartate aminotransferase, lactoylglutathione lyase, class III peroxidase and pathogenesis related protein No.10. On the other hand,7proteins were down-expressed under salt stress, including ATP synthase (subunit β), dihydrolipoamide dehydrogenase family protein, serine protease, cysteine proteinase, Calmodulin-like protein, NADP-malic enzyme and class Ⅲ peroxidase. In leaves, salt stress induced eight proteins down-expressed, and these proteins included ATP synthase, Bp2A protein, Cp31AHv protein, Cp31BHv protein, glutamine synthetase, germin-like protein phosphoglycerate kinase, and oxygen-evolving enhancer protein No.2. In contrast, NAD-dependent epimerase, DegP protease and harpin binding protein No.1were up-expressed under salt stress.
Based on the membrane proteome analysis, it can be suggested that the main mechanisms of membrane proteins for salt tolerance in roots were protection of membrane stability, scavenging reactive oxygen species (ROS) and function of ion homeostasis, while the mechanisms of membrane proteins in leaves were protection of photosynthetic reaction activity. Furthermore, there was genotypic difference in expression levels of membrane proteins, with a wild barley, XZ16having higher expression of proteins involved in osmotic regulation, ion transport and protection of photosynthesis than CM72, which could be considered as the molecular basis of high salt tolerance for XZ16.
4. Mechanisms of metabolome responding to salt stress in barley.
In order to reveal adaptive metabolic pathways in barley under salt stress, CM72and XZ16were used to investigate metabolome changes in response to salt stress of300mM NaCl after three weeks using gas chromatography-mass spectrometry (GC-MS). A total of82key metabolites were identified and their concentrations were affected by salt stress. Compared with the control, salt stress caused the significant changes in the contents of53and55metabolites for CM72and XZ16, respectively. The response of metabolites in roots was enhanced TCA cycle and sugar accumulations, and inhibited glycolysis and amino acid synthesis. In contrast, Calvin cycle, glycolysis and amino acid synthesis were enhanced, while TCA cycle was inhibited in leaves exposed to salt stress.
There was an obvious difference in metabolites in response to salt stress between roots and leaves. The results showed that the most important compatible solutes are proline, sugars (sucrose, raffinose and trehalose), mannitol and inositol for roots, and raffinose, proline and some amino acids for leaves. Osmotic adjustment is a basic mechanism for salt tolerance in roots. In leaves, a dramatic enhancement of proline, sugars and amino acid synthesis was found, which is obviously favorable for osmotic adjustment, and may provide sufficient carbohydrates and energy for roots. In addition, metabolites in response to salt stress differ between cultivated and wild barleys. CM72accumulated more metabolites associated with photosynthesis and TCA cycle in leaves, but less amino acids and organic acids, suggesting that the cultivated barley enhances its salt tolerance through increasing photosynthesis and energy consumption. While the contents of fructose-6-P, glucose-6-P and PEP were not affected by salt stress in the leaves of XZ16. Wild barley has higher contents of compatible solutes, more active metabolite synthesis and rapid growth than cultivated barley under salt stress.
In summary, XZ16, a Tibetan wild barley accession was identified in this study, being higher salt tolerance than CM72, a standard cultivated cultivar. Salt tolerance of barley is involved in a complex molecular network, and to our understanding it is first attempt that we systematically investigated the different mechanisms of salt tolerance between wild barleys and cultivated barleys using ionomic, membrane proteomic and metabolomic methods. The results surely enrich the knowledge of crop salt tolerance and can provide a theoretical reference and practical guidance for exploring genetic resources and breeding crop cultivars with salt stress.
大麦(Hordeum vulgare)是全球第四大粮食作物,适应性广,耐盐性强。以大麦为模式作物,研究其耐盐机制,对揭示禾本科作物的耐盐机理具有重要的理论价值。青藏高原一年生野生大麦(H. Spontaneum和H. agriocrithum)(以下称西藏野生大麦)系我国独有的植物种质资源,具有丰富的遗传变异和抗逆基因资源,发掘其耐盐资源可为大麦及作物耐盐改良提供优异种质。
本研究在评价西藏野生大麦耐盐性的基础上,采用关联分析法鉴定与发掘优异耐盐野生种质,并首次从离子组、膜蛋白组和代谢组三个水平上系统解析野生大麦和栽培大麦对盐胁迫的响应机制及其差异,取得的主要研究结果如下：
1.西藏野生大麦耐盐种质的发掘与遗传关联分析
采用水培试验,大麦三叶期用300mM NaCl胁迫处理3周,以相对干物重为指标,评价了188份西藏野生大麦的耐盐性,发现盐胁迫明显降低根和地上部的干重,降幅为27.6%～73.1%,且以根的降幅较大。西藏野生大麦的耐盐性存在广泛的基因型差异,与普遍认同的耐盐对照品种CM72相比,鉴定到根和地上部相对干重显著大于对照的野生大麦材料。
利用57对多态性芯片技术(DArT)标记分析了西藏野生大麦群体第5染色体的连锁不平衡(LD)结构,发现第5染色体LD的遗传衰减距离为8.9cM(R2<0.1)或者1.5cM (R2<0.2),LD程度较弱。采用测序法分析了野生大麦群体中HvCBF1、HvCBF3、HvCBF4和HvHVA1基因编码区(CDS)序列多态性,与参照序列相比,分别检测到2、15、16和10个单核苷酸多态性(SNP)位点,即平均每100bp碱基中存在0.3、2、2.4和1.6个SNP,对应3、8、13和6个单倍型,基因SNP位点之间存在不同程度的LD关系。
耐盐性状与遗传多态性的关联分析结果表明,标记bPb-4891与地上部和整株的相对干物重显著关联(P<0.05),可分别解释表型变异的2.2%和2.3%；4个基因中,仅HvCBF4的单倍型13与地上部、根和整株的相对干物重显著关联,可分别解释7.44%、2.4%和5.34%的表型变异。标记bPb-4891与HvCBF4基因在大麦遗传图谱上紧密连锁,推测该位点和HvCBF4的单倍型13可能与大麦耐盐性相关。野生大麦种质XZ16和XZ26具有HvCBF4的单倍型13的基因型,具有较强的耐盐能力。
2.大麦离子组对盐胁迫的响应机制
以栽培品种CM72(耐盐)、Gairdner(敏感)和两个野生大麦XZ16(耐盐)和XZ169(敏感)为材料,苗期进行150和300mM NaCl处理0、1、2、3和5周,系统分析了它们的生长和离子组对盐胁迫的响应。结果表明,低盐浓度对大麦生长的抑制不显著,而高盐浓度对4个大麦地上部的抑制率为30～53%,XZ16在两种盐胁迫下均有最大的绝对干物重,低浓度和高浓度下分别是CM72干物重的1.7倍和1.4倍。结果佐证了XZ16是生长快的耐盐材料,而Gairdner和XZ169的耐盐性弱。
根离子组中,盐胁迫下Na含量上升至处理2周后有所降低；P、S、Mg、K、 Mn、Zn和B含量因盐处理而降低,但Mn和Zn在高盐胁迫下降幅较低,Ca、Cu和Fe的含量因盐处理而增加。地上部离子组中,盐处理增加Na含量,降低P、K、Ca、Mg、S、Cu和B含量；Mn和Zn含量在低盐浓度下增加,而在高盐浓度下降低；而Fe含量在低盐下降低,高盐下增加。处理5周后,CM72和XZ16的Na含量和Na/K比显著低于Gairdner和XZ169。以上结果说明维持地上部低Na含量和Na/K比是大麦重要的耐盐机制。此外,大麦通过增加根部Ca、Cu和Fe的含量以及地上部Fe的含量,可增强组织对盐胁迫的耐性。
3.大麦膜蛋白对盐胁迫的响应机制
采用双向电泳(2-DE)和基质辅助激光解离飞行时间质谱系统(MALDI-TOF-TOF-MS)技术,分析了200mM NaCl胁迫处理48h和正常条件下XZ16和CM72大麦根和叶片膜蛋白组的差异,结果发现有41个膜蛋白受盐胁迫诱导差异表达。在根部,ATP合成酶(α亚基)、天冬氨酸转氨酶、单脱氢抗坏血还原酶、第Ⅲ类过氧化物酶和病程相关蛋白10受盐胁迫诱导表达上调,而ATP合成酶(β亚基)、二氢硫辛酰胺脱氢酶家族、丝氨酸蛋白酶、半胱氨酸蛋白酶、钙调蛋白类、NADP-苹果酸酶和第Ⅲ类过氧化物酶受盐胁迫表达下调。叶片中,叶绿体ATP合成酶、Bp2A蛋白、Cp31AHv蛋白、Cp31BHv蛋白、谷氨酰胺合成酶、磷酸甘油酸激酶、二磷酸腺苷葡萄糖焦磷酸酶和叶绿体Oxygen-evolving增强蛋白2受盐胁迫表达下调；而NAD依赖型异构酶、DegP蛋白酶和Harpin结合蛋白受盐胁迫诱导表达上调。
研究结果推测,根部膜蛋白的主要响应机制是保护细胞膜稳定性、清除活性氧自由基(ROS)和参与离子平衡的重建,叶片膜蛋白的主要响应机制是保护光合反应活性。膜蛋白的表达水平基因型之间存在差异,XZ16野生大麦参与渗透调节和离子转运以及保护叶片的光合作用的膜蛋白表达高于栽培大麦CM72,这可能也是前者具有较强耐盐性的分子基础。
4.大麦代谢物组对盐胁迫的响应机制
采用气相色谱-质谱(GC-MS)技术,分析了300mM NaCl胁迫处理3周和正常条件下XZ16和CM72大麦根和叶片代谢组差异,鉴定到82种代谢物响应盐胁迫。与对照相比,CM72和XZ16分别有53种和55种代谢物含量显著变化,根代谢组的响应方式是参与TCA循环和糖转运的代谢物积累增加,而参与糖酵解和氨基酸合成的代谢物积累受到抑制。叶片代谢组中,CM72和XZ16分别有55种和54种代谢物含量显著变化,其响应方式是参与糖酵解、Calvin循环和氨基酸合成的代谢物积累明显增加,而参与TCA循环的代谢物积累受到抑制。
代谢物响应盐胁迫存在组织差异性,根部调节渗透的主要代谢物是脯氨酸、糖类(蔗糖、海藻糖和棉子糖)、甘露醇和纤维醇；而棉子糖和脯氨酸等多种氨基酸是叶片的主要渗透调节物。根部代谢物对盐胁迫的响应机制是积累渗透调节剂含量,增强耐盐性；而叶片代谢物的响应机制除增强组织的渗透调节能力外,还为根部提供足够的糖类和能量,从而调节渗透压和转运Na+。代谢物响应机制存在基因型差异,栽培大麦CM72通过增加代谢物的合成,从而增强渗透调节和离子平衡提供物质和能量；而野生大麦XZ16自身拥有较强的渗透调节能力,维持正常的光合作用和合成代谢,从而保持较高的生长水平。
综上所述,大麦的耐盐性受一个复杂的分子调控网络控制。本研究从西藏野生大麦中鉴定到耐盐性优于CM72的种质,可为大麦耐盐育种提供遗传材料；并从离子组、膜蛋白组和代谢组上分别解析了野生大麦和栽培大麦对盐胁迫的响应及其差异,丰富了作物耐盐机理,可为进一步发掘耐盐遗传资源和培育耐盐作物品种提供理论参考和实践指导。
Soil salinity is a major abiotic stress of restricting crop productivity worldwide, posing a great threat to agricultural sustainability. A thorough understanding of salt-tolerant mechanisms is necessary for alleviating salt injury to agricultural production by improving cultural practices and developing salt-tolerant varieties. Currently, it is well documented that osmotic regulation, ion homeostasis and anti-oxidation are the three aspects of salt tolerance in plants. However, the genetic and molecular mechanisms of salt tolerance have not been fully elucidated.
Barley (Hordeum vulgare) is the fourth most important cereal crop in the world, characterized by its wide adaptability and high salinity tolerance. Consequently, it is frequently used as a model crop in the attempts to understand salinity tolerance in the cereal crops. Tibetan annual wild barley (H. vulgare ssp. Spontaneum and ssp. agriocrithum)(hereafter referred to as Tibetan wild barley) is considered as one of the ancestors of cultivated barley, and shows wide genetic variations in the tolerance to harsh environments. It is possible for us to explore the elite germplasm in terms of salt tolerance from the wild barley.
In the present study, an association analysis was used to identify and explore salt-tolerant germplasm based on the evaluation of salt tolerance among around200Tibetan wild barley accessions, and the mechanisms of salt tolerance between wild and cultivated barleys were systematically resolved by ionomic, membrane proteomic and metabolomic methods. The main results were summarized as follows:
1. Identification of salt-tolerant germplasm and genetic association analysis within Tibetan wild barleys.
Hundred and eighty-eight accessions of Tibetan wild barley were exposed to salt stress of300mM NaCl in hydroponics at three-leaf stage for3weeks, to compare the difference among these accessions in salt tolerance as indicated by relative dry weights. Salt stress significantly reduced shoot and root dry weight by27.6%to73.1%and the root weight showed greater reduction than the shoot weight. There was a significant difference among the Tibetan wild barley accessions. Compared with CM72, a salt-tolerant cultivated cultivar, some accessions showed higher shoot, root or whole-plant dry weight under salt stress.
The linkage disequilibrium (LD) structures of the chromosome5were evaluated using57diversity arrays technology (DArT) markers over this chromosome. The LD decay of genetic distance was8.9cM (R2<0.1) or1.5cM (R2<0.2). The genetic variation of HvCBF1, HvCBF3, HvCBF4and HvHVA1was studied by sequencing the gene coding regions. There were2,15,16and10single nucleotide polymorphisms (SNP) sites, corresponding to3,8,13and6haplotypes for the four genes, respectively, with0.3,2,2.4and1.6SNPs in each100bp of the coding sequence. Furthermore, the certain LD relationship was detected between SNPs.
Association analysis was conducted between the genetic diversity and the genotypic salt tolerance. It was found that marker bpb-4891was significantly associated with salt tolerance, explaining the2.2%and2.3%of phenotypic variation for the relative shoot and whole-plant dry weight, respectively. The haplotype13of HvCBF4gene exhibited highly significant association with shoot and whole-plant relative dry weight, explaining7.7%and6.4%of the variations, respectively. Based on the distance in the barley genetic map, it may be assumed that the marker bpb-4891was closely linked with the HvCBF4gene. The results suggest that the marker bpb-4891and haplotype13of HvCBF4gene could be related to salinity tolerance. Tibetan wild barley accessions, XZ16and XZ26have haplotype13of HvCBF4genotypes, and could be considered as high salt tolerance.
2. Mechanisms of ionome responding to salt stress in barley
In order to reveal mechanisms of ionome responding to salt stress in barley, two cultivated barleys (CM72, salt-tolerant; Gairdner, salt-sensitive) and two wild barleys (XZ16, salt-tolerant; XZ169, salt-sensitive) differing in salt tolerance were used to investigate shoot biomass and ion changes in response to salt stress of150and300mM NaCl in hydroponics at0,1,2,3and5weeks after treatment. After35days treatment, there was no significant difference in shoot weight between salt stress of150mM NaCl and control for4barley genotypes, but under salt stress of300mM NaCl, the reduction of shoot weight could be found, ranging from30%to53%. Among the four genotypes, XZ16maintained the largest shoot biomass under salt stress, and it had1.7and1.4times larger shoots dry weight than CM72, respectively. In conclusion, the results show that XZ16is a fast-growing and salt-tolerant barley.
In roots, Na content reached the maximum at2weeks after treatments, and then tended to decrease. The contents of P, S, Mg, K, Mn, Zn and B decreased steadily during salt treatment, while Mn and Zn contents decreased greatly under salt stress of300mM NaCl. By contrast, the contents of Ca, Cu and Fe increased under salt stress. In shoots, Na content increased and the contents of P, K, Ca, Mg, S, Cu and B decreased due to salt treatments. The contents of Mn and Zn increased under salt stress of150mM NaCl, and increased under salt stress of300mM NaCl. In contrast, Fe content decreased under salt stress of150mM NaCl, but increased under salt stress of300mM NaCl. These results suggest that maintenance of low Na content and Na/K ratio in shoots is a key important for salt tolerance in both cultivated and wild barleys. In addition, the increased contents of Ca, Cu and Fe in roots and Fe in shoots may enhance tissues tolerance to salt stress.
3. Mechanisms of membrane proteins responding to salt stress in barley.
In order to reveal mechanisms of membrane proteins responding to salt stress in barley, wild barley (XZ16) and cultivated barley (CM72) were used to investigate differential expression of membrane proteomes in roots and leaves under normal condition and salt stress of200mM NaCl after48hours, using two-dimensional electrophoresis (2-DE) and matrix-assisted laser desorption/ionization time of flight mass spectrometry (MALDI-TOF-TOF-MS) techniques.
In roots, salt stress induced6proteins up-expressed, being more than1.5times than those in the control, and these proteins included ATP synthase (subunit a), aspartate aminotransferase, lactoylglutathione lyase, class III peroxidase and pathogenesis related protein No.10. On the other hand,7proteins were down-expressed under salt stress, including ATP synthase (subunit β), dihydrolipoamide dehydrogenase family protein, serine protease, cysteine proteinase, Calmodulin-like protein, NADP-malic enzyme and class Ⅲ peroxidase. In leaves, salt stress induced eight proteins down-expressed, and these proteins included ATP synthase, Bp2A protein, Cp31AHv protein, Cp31BHv protein, glutamine synthetase, germin-like protein phosphoglycerate kinase, and oxygen-evolving enhancer protein No.2. In contrast, NAD-dependent epimerase, DegP protease and harpin binding protein No.1were up-expressed under salt stress.
Based on the membrane proteome analysis, it can be suggested that the main mechanisms of membrane proteins for salt tolerance in roots were protection of membrane stability, scavenging reactive oxygen species (ROS) and function of ion homeostasis, while the mechanisms of membrane proteins in leaves were protection of photosynthetic reaction activity. Furthermore, there was genotypic difference in expression levels of membrane proteins, with a wild barley, XZ16having higher expression of proteins involved in osmotic regulation, ion transport and protection of photosynthesis than CM72, which could be considered as the molecular basis of high salt tolerance for XZ16.
4. Mechanisms of metabolome responding to salt stress in barley.
In order to reveal adaptive metabolic pathways in barley under salt stress, CM72and XZ16were used to investigate metabolome changes in response to salt stress of300mM NaCl after three weeks using gas chromatography-mass spectrometry (GC-MS). A total of82key metabolites were identified and their concentrations were affected by salt stress. Compared with the control, salt stress caused the significant changes in the contents of53and55metabolites for CM72and XZ16, respectively. The response of metabolites in roots was enhanced TCA cycle and sugar accumulations, and inhibited glycolysis and amino acid synthesis. In contrast, Calvin cycle, glycolysis and amino acid synthesis were enhanced, while TCA cycle was inhibited in leaves exposed to salt stress.
There was an obvious difference in metabolites in response to salt stress between roots and leaves. The results showed that the most important compatible solutes are proline, sugars (sucrose, raffinose and trehalose), mannitol and inositol for roots, and raffinose, proline and some amino acids for leaves. Osmotic adjustment is a basic mechanism for salt tolerance in roots. In leaves, a dramatic enhancement of proline, sugars and amino acid synthesis was found, which is obviously favorable for osmotic adjustment, and may provide sufficient carbohydrates and energy for roots. In addition, metabolites in response to salt stress differ between cultivated and wild barleys. CM72accumulated more metabolites associated with photosynthesis and TCA cycle in leaves, but less amino acids and organic acids, suggesting that the cultivated barley enhances its salt tolerance through increasing photosynthesis and energy consumption. While the contents of fructose-6-P, glucose-6-P and PEP were not affected by salt stress in the leaves of XZ16. Wild barley has higher contents of compatible solutes, more active metabolite synthesis and rapid growth than cultivated barley under salt stress.
In summary, XZ16, a Tibetan wild barley accession was identified in this study, being higher salt tolerance than CM72, a standard cultivated cultivar. Salt tolerance of barley is involved in a complex molecular network, and to our understanding it is first attempt that we systematically investigated the different mechanisms of salt tolerance between wild barleys and cultivated barleys using ionomic, membrane proteomic and metabolomic methods. The results surely enrich the knowledge of crop salt tolerance and can provide a theoretical reference and practical guidance for exploring genetic resources and breeding crop cultivars with salt stress.
引文
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