西藏野生大麦铝毒耐性机理研究
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摘要
铝(A1)毒是酸性土壤抑制作物生长和产量形成的主要因子,解决土壤酸害铝毒问题,对充分利用土地和促进全球作物生产具有重大意义。因此,开展作物耐酸铝胁迫的生理生化与分子机理研究,揭示作物酸铝耐性机制,具有重要的理论和实践价值。青藏高原一年生野生大麦(Hordeum vulgare subsp. spontaneum)是我国特有的珍稀大麦种质,遗传多样性丰富,蕴藏有优异的抗逆等位基因。本研究以青藏高原野生大麦为主要研究材料,研究其耐铝的生理与遗传特性,取得的主要结果如下：
1.酸土胁迫对大麦离子组的影响
以90个大麦基因型为材料,测定根和地上部离子组响应酸土胁迫的变化。酸土(pH<4.5)胁迫下,根系铝含量与对照(施石灰)无显著差异,而地上部铝含量显著高于对照；地上部铝含量耐铝品种低于敏感品种。载荷分析表明,地上部铝含量是响应酸土胁迫的主要组分。酸土胁迫显著增加根系磷含量,而降低地上部磷含量,这可能与根表大量形成Al-磷酸盐沉淀有关,阻碍了磷向地上部的转运。酸铝胁迫扰乱地上部硫、钾和锌与其它元素之间的关系,打破地上部组织的离子平衡。
2.西藏野生大麦与栽培大麦酸铝耐性的全基因组关联分析
对110份野生大麦和56份栽培大麦在苗期进行耐铝性鉴定,并结合469个均匀分布于所有大麦染色体上的DArT标记,分别对野生群体、栽培群体和所有供试材料进行耐铝性全基因组关联分析(GWAS)。群体结构和聚类分析表明,野生大麦具有较丰富的遗传多样性,与栽培大麦(11.52cM)相比,野生大麦(9.30cM)的LD衰减较快。因此,利用野生大麦进行关联作图具有相对较高的分辨率。在野生群体中鉴定到两个新的耐铝位点,bpb-9458和bpb-8524,它们与相对最长根伸长率(RLRG)相关联,分别位于2H和7H染色体上,可分别解释12.9%和9.7%的表型变异。此外,在整个群体和栽培群体中鉴定到一个耐铝位点bpb-6949,它与铝胁迫下最长根绝对伸长量(TRG)相关联,与已报道的候选基因HvMATE相距较近(0.8cM)。以上研究结果充分显示野生大麦是大麦耐铝研究和遗传改良的优异种质资源。
3.酸铝胁迫对大麦根系结构、抗氧化系统和有机酸分泌的影响
从根系形态、抗氧化系统和根系有机酸分泌等特性上研究了野生大麦X29(耐铝)、X9(铝敏感)和栽培大麦品种Dayton (耐铝)对铝浓度和处理的响应。结果表明,铝胁迫下,X9的根伸长受到显著抑制,变粗,根表面积减少；而X29和Dayton的根系形态受影响较小。铝胁迫影响抗氧化酶活性,增加丙二醛含量,以X9受影响最大。X29与X9在铝诱导的根系柠檬酸分泌量和(?)HvMATE基因表达量上差异不大,且HvMATE基因表达量均远低于Dayton。与X9相比,铝诱导X29根系分泌较多的草酸和磷酸。结果显示,铝诱导并不显著影响X29的根系柠檬酸分泌,但影响根系草酸和磷酸的分泌,这可以认为是X29的潜在耐铝机制,但需作进一步研究验证。
4.大麦响应酸铝胁迫的代谢组学分析
采用气相色谱-质谱联用技术(GC-MS)测定了X29、X9和Dayton在对照(不加铝)、5μM Al和25μM Al处理6天后根系中79种代谢物的相对含量,并绘制了代谢通路。高铝胁迫下,Dayton通过提高氧自由基清除剂含量,如脯氨酸、阿魏酸、4-羟基-苯甲酸和4-羟基-3-甲基-苯甲酸,减轻膜脂过氧化程度,缓解铝毒造成的氧化胁迫。铝胁迫下,X29能维持较高的糖类代谢中间产物含量,如6-磷酸-葡萄糖和6-磷酸-果糖,这可能是其表现耐铝的生理基础。同时,X29在铝胁迫下根系维持着高水平的磷酸盐和草酸,并向外分泌,螯合根际铝离子,这可能是这一野生大麦材料耐铝的一种主要机制。
Aluminum (Al) toxicity is a main constraint of crop growth and yield formation in acid soils. Overcoming of the problem caused by Al toxicity is quite important for fully utilizing land resource and increasing crop production in the world. Therefore, Understanding of physiological and molecular mechanisms of Al tolerance in crops is imperative. Tibetan annual wild barley(Hordeum vulgare subsp. spontaneum), recognized as one of the ancestors of cultivated barley, is a previous plant germplasm, rich in genetic diversity and elite stress-tolerant alleles. In the present study, the physiological and genetic characteristics of Tibetan wild barley were investigated to make sight into the Al3+-tolerant mechanisms. The main findings were summarized as follows:
1. The effect of acid soil stress on ionomic change in barley
90barley genotypes, including wild and cultivated barleys, were planted in the acid soil with pH below4.5and limed soil, to determine growth and ionomic difference among genotypes under the two soil conditions. There was no significant difference in root aluminum (A1) concentration between the plants on the acid and limed soils. However, the plants grew in the acid soil had higher shoot Al concentration than those grew in the limed soil. Moreover, the genotypes tolerant to acid soil showed lower shoot Al concentration than sensitive ones. Loading analysis indicated that shoot Al concentration is the main factor contributed to shoot ionomic change under Al stress. Root phosphorus (P) concentration was higher for the plants subjected to acid soil than those to limed soil, but opposite was true for shoot P concentration. It may be suggested that Al-phosphate complex was formed in root surface for plants in response to Al stress, preventing P transportation from roots to shoots. Al stress disordered the relationships between S, K, Zn and other elements in shoots, resulting in disturbance of shoot ionic homostesis.
2. Genome-wide association analysis of aluminum tolerance in cultivated and Tibetan wild barley
A genome-wide association analysis (GWAS) was performed by using four root parameters reflecting Al tolerance and469DArT markers distributed evenly on all7barley chromosomes after making evaluation of A1tolerance of110Tibetan wild accessions and56cultivated cultivars. Population structure and cluster analysis revealed that a wider genetic diversity was present in Tibetan wild barley. Linkage disequilibrium (LD) decayed more rapidly in Tibetan wild barley (9.30cM) than in cultivated barley (11.52cM), indicating that GWAS may provide higher resolution in Tibetan group. Two novel Tibetan group-specific loci, bpb-9458and bpb-8524were identified. They were associated with relative longest root growth (RLRG), and located on chromosomes2H and7H, and could explain12.9%and9.7%of the phenotypic variation, respectively. Moreover, a common locus bpb-6949, localized at0.8cM away from a candidate gene HvMATE, was detected in both wild and cultivated barleys, and it was significantly associated with total root growth (TRG). The present results showed that Tibetan wild barley could provide elite germplasm and novel genes for barley Al-tolerant improvement.
3. The effect of A1stress on root morphology, antioxidative enzymes and organic acid secretion in barley
The responses of root morphology, antioxidative enzymes and organic acid secretion of wild barley accessions X29, X9and cultivated cultivar Dayton to Al stress were examined. More inhibition of root elongation, larger root diameter and smaller root surface area were observed for X9in comparison with other two genotypes. Under Al stress, the activities of antioxidative enzymes were greatly affected, and malondialdehyde (MDA) content was greatly increased in X9. There was no significant difference in Al3+-induced citrate secretion or HvMATE expression between X29and X9, but Dayton showed much higher HvMATE expression level. The amount of oxalate and phosphate secreted in X29was higher than that in X9under Al stress. It may be suggested that Al3+-induced oxalate and phosphate efflux could be a possible Al3+-resistant mechanism, which need to be proved by more evidences.
4. Metabolites and metabolic pathway in response to Al stress
Using gas chromatography-mass spectrometry (GC-MS), we investigated the metabolites levels of X29, X9and Dayton under Al stress. A total of79key metabolites were identified and the adaptive metabolic pathway was constructed. The concentrations of ROS cleaners, such as proline, Ferulic acid,4-hydroxy-benzoic acid and4-hydroxy-3-methyl benzoic acid, increased greatly in Dayton roots, in response to increased membrane lipid peroxidation. Higher concentrtions of carbohydrate metabolic intermediates, such as glucose-6-P and fructose-6-P in X29could be a physilogical basis for Al tolerance. Maintainance of high level in root phosphate and oxalate concentrations, which are then secreted into rhizophere and cultrue solution, alleviating Al stress, could be a Al3+-tolerant mechanism in X29. But the hypothesis should be confirmed by further study.
1.酸土胁迫对大麦离子组的影响
以90个大麦基因型为材料,测定根和地上部离子组响应酸土胁迫的变化。酸土(pH<4.5)胁迫下,根系铝含量与对照(施石灰)无显著差异,而地上部铝含量显著高于对照；地上部铝含量耐铝品种低于敏感品种。载荷分析表明,地上部铝含量是响应酸土胁迫的主要组分。酸土胁迫显著增加根系磷含量,而降低地上部磷含量,这可能与根表大量形成Al-磷酸盐沉淀有关,阻碍了磷向地上部的转运。酸铝胁迫扰乱地上部硫、钾和锌与其它元素之间的关系,打破地上部组织的离子平衡。
2.西藏野生大麦与栽培大麦酸铝耐性的全基因组关联分析
对110份野生大麦和56份栽培大麦在苗期进行耐铝性鉴定,并结合469个均匀分布于所有大麦染色体上的DArT标记,分别对野生群体、栽培群体和所有供试材料进行耐铝性全基因组关联分析(GWAS)。群体结构和聚类分析表明,野生大麦具有较丰富的遗传多样性,与栽培大麦(11.52cM)相比,野生大麦(9.30cM)的LD衰减较快。因此,利用野生大麦进行关联作图具有相对较高的分辨率。在野生群体中鉴定到两个新的耐铝位点,bpb-9458和bpb-8524,它们与相对最长根伸长率(RLRG)相关联,分别位于2H和7H染色体上,可分别解释12.9%和9.7%的表型变异。此外,在整个群体和栽培群体中鉴定到一个耐铝位点bpb-6949,它与铝胁迫下最长根绝对伸长量(TRG)相关联,与已报道的候选基因HvMATE相距较近(0.8cM)。以上研究结果充分显示野生大麦是大麦耐铝研究和遗传改良的优异种质资源。
3.酸铝胁迫对大麦根系结构、抗氧化系统和有机酸分泌的影响
从根系形态、抗氧化系统和根系有机酸分泌等特性上研究了野生大麦X29(耐铝)、X9(铝敏感)和栽培大麦品种Dayton (耐铝)对铝浓度和处理的响应。结果表明,铝胁迫下,X9的根伸长受到显著抑制,变粗,根表面积减少；而X29和Dayton的根系形态受影响较小。铝胁迫影响抗氧化酶活性,增加丙二醛含量,以X9受影响最大。X29与X9在铝诱导的根系柠檬酸分泌量和(?)HvMATE基因表达量上差异不大,且HvMATE基因表达量均远低于Dayton。与X9相比,铝诱导X29根系分泌较多的草酸和磷酸。结果显示,铝诱导并不显著影响X29的根系柠檬酸分泌,但影响根系草酸和磷酸的分泌,这可以认为是X29的潜在耐铝机制,但需作进一步研究验证。
4.大麦响应酸铝胁迫的代谢组学分析
采用气相色谱-质谱联用技术(GC-MS)测定了X29、X9和Dayton在对照(不加铝)、5μM Al和25μM Al处理6天后根系中79种代谢物的相对含量,并绘制了代谢通路。高铝胁迫下,Dayton通过提高氧自由基清除剂含量,如脯氨酸、阿魏酸、4-羟基-苯甲酸和4-羟基-3-甲基-苯甲酸,减轻膜脂过氧化程度,缓解铝毒造成的氧化胁迫。铝胁迫下,X29能维持较高的糖类代谢中间产物含量,如6-磷酸-葡萄糖和6-磷酸-果糖,这可能是其表现耐铝的生理基础。同时,X29在铝胁迫下根系维持着高水平的磷酸盐和草酸,并向外分泌,螯合根际铝离子,这可能是这一野生大麦材料耐铝的一种主要机制。
Aluminum (Al) toxicity is a main constraint of crop growth and yield formation in acid soils. Overcoming of the problem caused by Al toxicity is quite important for fully utilizing land resource and increasing crop production in the world. Therefore, Understanding of physiological and molecular mechanisms of Al tolerance in crops is imperative. Tibetan annual wild barley(Hordeum vulgare subsp. spontaneum), recognized as one of the ancestors of cultivated barley, is a previous plant germplasm, rich in genetic diversity and elite stress-tolerant alleles. In the present study, the physiological and genetic characteristics of Tibetan wild barley were investigated to make sight into the Al3+-tolerant mechanisms. The main findings were summarized as follows:
1. The effect of acid soil stress on ionomic change in barley
90barley genotypes, including wild and cultivated barleys, were planted in the acid soil with pH below4.5and limed soil, to determine growth and ionomic difference among genotypes under the two soil conditions. There was no significant difference in root aluminum (A1) concentration between the plants on the acid and limed soils. However, the plants grew in the acid soil had higher shoot Al concentration than those grew in the limed soil. Moreover, the genotypes tolerant to acid soil showed lower shoot Al concentration than sensitive ones. Loading analysis indicated that shoot Al concentration is the main factor contributed to shoot ionomic change under Al stress. Root phosphorus (P) concentration was higher for the plants subjected to acid soil than those to limed soil, but opposite was true for shoot P concentration. It may be suggested that Al-phosphate complex was formed in root surface for plants in response to Al stress, preventing P transportation from roots to shoots. Al stress disordered the relationships between S, K, Zn and other elements in shoots, resulting in disturbance of shoot ionic homostesis.
2. Genome-wide association analysis of aluminum tolerance in cultivated and Tibetan wild barley
A genome-wide association analysis (GWAS) was performed by using four root parameters reflecting Al tolerance and469DArT markers distributed evenly on all7barley chromosomes after making evaluation of A1tolerance of110Tibetan wild accessions and56cultivated cultivars. Population structure and cluster analysis revealed that a wider genetic diversity was present in Tibetan wild barley. Linkage disequilibrium (LD) decayed more rapidly in Tibetan wild barley (9.30cM) than in cultivated barley (11.52cM), indicating that GWAS may provide higher resolution in Tibetan group. Two novel Tibetan group-specific loci, bpb-9458and bpb-8524were identified. They were associated with relative longest root growth (RLRG), and located on chromosomes2H and7H, and could explain12.9%and9.7%of the phenotypic variation, respectively. Moreover, a common locus bpb-6949, localized at0.8cM away from a candidate gene HvMATE, was detected in both wild and cultivated barleys, and it was significantly associated with total root growth (TRG). The present results showed that Tibetan wild barley could provide elite germplasm and novel genes for barley Al-tolerant improvement.
3. The effect of A1stress on root morphology, antioxidative enzymes and organic acid secretion in barley
The responses of root morphology, antioxidative enzymes and organic acid secretion of wild barley accessions X29, X9and cultivated cultivar Dayton to Al stress were examined. More inhibition of root elongation, larger root diameter and smaller root surface area were observed for X9in comparison with other two genotypes. Under Al stress, the activities of antioxidative enzymes were greatly affected, and malondialdehyde (MDA) content was greatly increased in X9. There was no significant difference in Al3+-induced citrate secretion or HvMATE expression between X29and X9, but Dayton showed much higher HvMATE expression level. The amount of oxalate and phosphate secreted in X29was higher than that in X9under Al stress. It may be suggested that Al3+-induced oxalate and phosphate efflux could be a possible Al3+-resistant mechanism, which need to be proved by more evidences.
4. Metabolites and metabolic pathway in response to Al stress
Using gas chromatography-mass spectrometry (GC-MS), we investigated the metabolites levels of X29, X9and Dayton under Al stress. A total of79key metabolites were identified and the adaptive metabolic pathway was constructed. The concentrations of ROS cleaners, such as proline, Ferulic acid,4-hydroxy-benzoic acid and4-hydroxy-3-methyl benzoic acid, increased greatly in Dayton roots, in response to increased membrane lipid peroxidation. Higher concentrtions of carbohydrate metabolic intermediates, such as glucose-6-P and fructose-6-P in X29could be a physilogical basis for Al tolerance. Maintainance of high level in root phosphate and oxalate concentrations, which are then secreted into rhizophere and cultrue solution, alleviating Al stress, could be a Al3+-tolerant mechanism in X29. But the hypothesis should be confirmed by further study.
引文
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