玉米麦根酸代谢评价体系的建立及缺铁胁迫相关代谢基因片段的克隆
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摘要
缺铁是世界农业生产中非常普遍的一个产量限制因子,在干旱半干旱的石灰性土壤上,果树、玉米、高粱等作物的缺铁问题尤为严重。解决这一问题有两条途径:一是通过改良土壤营养状况,改变或排除土壤障碍因子以适应植物生长发育,如施肥、添加土壤改良剂和其他技术措施等。二是通过育种手段,改良植物的矿质营养遗传特性,使其活化被土壤吸附固定的养分,变固定态为可利用态,从而被植物吸收利用,即挖掘植物和土壤本身固有的潜力充分利用自然资源。培育铁营养高效新品种是减轻缺铁影响的有效方法,是当前的研究热点之一。
禾本科类植物能够由根系分泌麦根酸类物质(Mugineic acids,简称MAs),也叫植物铁载体(Phytosiderophores,简称PS)来活化土壤中难以被植物吸收利用的铁,使之易于被植物吸收利用,从而解决自身铁营养缺乏的问题。
本研究以玉米自交系为材料,对三种玉米麦根酸类物质的测定方法进行比较,同时分析了不同玉米自交系苗期麦根酸类物质的分泌能力和缺铁敏感性,比较了不同生育时期玉米麦根酸分泌能力的差异,研究了玉米麦根酸分泌特性的遗传规律,建立了玉米苗期缺铁胁迫诱导的cDNA文库,并对该cDNA文库的近1000个克隆进行了测序分析。主要研究结果如下:
1筛选建立了玉米麦根酸类物质测定方法
以16种玉米自交系为材料,根据麦根酸类物质的溶铁特性,建立了麦根酸类物质间接测定新方法——ICP-AES法,改良了脱氧麦根酸(DMA)的UV-HPLC直接测定法,并与传统的比色法测定结果进行比较。结果表明,采用ICP-AES法,以Fe_2O_3粉末作为活化对象,与传统采用Fe(OH)_3的测定结果相比,结果稳定、重复性好,说明ICP-AES法是玉米麦根酸类物质分泌量的一种简单、快速的间接测定方法。改良的UV-HPLC法也具有准确、高效的特点,可应用于禾谷类作物DMA分泌量的直接测定。高效液相色谱法是PS测定的准确方法,但在材料选择时可以利用ICP-AES方法来对材料的麦根酸分泌能力进行测定。
2建立了玉米自交系麦根酸分泌特性的评价体系
以46种玉米自交系为材料,利用高效液相色谱法和等离子体法测定了苗期和穗花期缺铁胁迫前后玉米根系的麦根酸分泌速率,评价了不同材料的麦根酸分泌能力、缺铁敏感性和不同生育时期分泌量的变化。结果得出,在苗期缺铁胁迫前后材料的PS分泌能力之间存在着显著相关关系同时两种测定方法之间存在显著相关关系r (Fe+) = 0.7149~(**);麦根酸分泌特性在不同发育时期存在较大差异,四叶一心期和穗花期的测定结果具有显著的相关性r=0.9701~*,穗花期玉米PS分泌速率的下降程度与苗期玉米PS分泌速率之间具有显著的对数相关关系r=0.892~*;利用苗期材料的麦根酸分泌能力和缺铁敏感性建立了麦根酸分泌特性的评价体系,以缺铁胁迫前PS分泌速率的平均数[2.836 mg/(g~*per plant~*3h)]和缺铁胁迫后的PS分泌速率增加量的平均数[0.209 mg/(g~*per plant~*3h)]作为分界线可以将材料分为具有不同PS分泌能力和缺铁敏感性的四种类型:HH,HL,LH,LL,其中具有LH型低PS分泌速率和高缺铁胁迫敏感性的材料是铁营养高效育种的理想材料。
3分析了玉米麦根酸代谢遗传规律
选用八个具有不同MAs分泌能力的玉米自交系,采用半双列杂交的试验设计,分析了杂交后代的麦根酸分泌能力,结果表明亲本麦根酸分泌特性的测定结果与材料自身GCA的相关系数均达到极显著水平,SCA效应值最高的组合并不一定是麦根酸分泌能力最高的组合,只有组合的SCA和双亲的GCA效应均高,才有可能获得麦根酸分泌能力较高的组合,组合的表现与其TCA效应一致。玉米麦根酸类物质的分泌特性以显性效应占优势,遗传加性效应相对较小,麦根酸分泌能力的遗传模型经检验证明符合加性-显性模型,而其缺铁敏感性的遗传模型则不符合。控制麦根酸分泌能力表现显性的主效基因组不少于一组,显性方向为减效;控制麦根酸缺铁敏感性表现显性的主效基因组数较少,显性方向为增效,二者的主效基因组可能不相同。
4建立了缺铁胁迫诱导的抑制性扣除杂交cDNA文库
以缺铁胁迫前的昌7-2根尖cDNA为driver cDNA,缺铁胁迫后的昌7-2根尖cDNA为Tester cDNA;以缺铁胁迫后的昌7-2根尖cDNA为driver cDNA,缺铁胁迫后的郑58根尖cDNA为Tester cDNA,进行两组抑制性扣除杂交,获得了与玉米缺铁胁迫代谢和麦根酸代谢能力差异表达有关的cDNA PCR产物,建立了相关文库。
5分析了玉米缺铁诱导代谢的相关基因
分析已获得的与玉米缺铁胁迫特异性表达的抑制性扣除杂交的PCR产物建立的SSH文库,其包含的片段大小为200bp-900bp。从文库中挑取了1230个克隆进行测序,除去8%的冗余片段共得到非重复的序列203个,其中77个的Blast结果表明它们可能是新发现的序列,除此之外其中:9个片段(0002,0034,0039,0143,0152, 0214, 0355,0375, 1152)可能与缺铁胁迫后植物适应代谢能量降低的调控有关;3个片段(0183,0232,0350)可能与细胞质膜上铁的还原酶有关;2个片段(0061,0622)可能与植物体内或细胞膜上铁离子的结合蛋白有关;5个片段(0094,0155,0208, 0249, 0985,1055)可能与细胞质膜的转运功能有关,3个片段(0234,0362,0387)与氨基酸的转运功能有关,5个片段(0103,0116,0274, 0382,0681)与植物的抗逆性有关,2个片段(0201,1183)与转录调控有关。同时在所有得到片段中, 0257与DNA的促旋酶有关。可能与遗传规律的减效显性作用有关。其余部分片段可能为未知功能片段。这些基因的差异表达与玉米缺铁胁迫的基因调控的关系有待进一步阐明和分析。
Iron (Fe) deficiency chlorosis (lime chlorosis) is a world widespread mineral nutrient deficiency in higher plants, especially corn, sorghum and fruit tree, grown on calcareous soils. Soil amendments and foliar sprays of Fe are not viable economically in the long term to correct Fe deficiency. Therefore, to alleviate Fe chlorosis in the long term, alternative methods, for example breeding of more chlorosis resistant genotypes (cultivars) are required. The recent hotspot in agricultural research is to reach the high yield, quality and efficiency target by the using of plant’s stress tolerance. Graminaceous species can enhance iron (Fe) acquisition from sparingly soluble inorganic Fe (Ⅲ) compounds by release of phytosiderophores (PS) which mobilize Fe (Ⅲ) by chelating.
Using corn inbred lines as material, this study had compared three PS amount test method, analized the PS secretion ability and iron sensitivity at seedling stage.While the different of PS secretion ability between seedling and heading stage and the regulation of PS secretion characteristic’s inherit had been studied. Two SSH cDNA library which one was about the Fe deficiency metobolize regulating, another was about the different PS secretion ability had been constructed.Using the SSH cDNA library which one was about the Fe deficiency metobolize regulating, the cDNA which reduced by Fe deficency had been analized.The main results were summarized as follow:
1 The Phytosiderophores (PS)’s test method had been set up.
Using 16 corn inbred lines,according to the Fe(Ⅲ) chelated characteristic of mugineic acid, a new indirect method of inductively coupled plasma-atomic emission spectrometry (ICP-AES) and a modified direct method of ultra-spectra high performance liquid chromatography (UV-HPLC) for the measurement of MAs or deoxymugineic acid (DMA) were established in corn inbred lines. The comparison of these two methods with traditional spectrometry was also made. The results showed that the ICP-AES method, using Fe_2O_3 as the dissoluble object instead of the traditional Fe (OH)_3, was more stable and repeatalble suggesting that ICP-AES was a simple and quick method for evaluation of the MAs content in corn and other graminaceous crops. The modified UV-HPLC was also precise and efficient, and could be used for the direct mesurement of graminaceous’s deoxymugineic acid.
2 The corn inbred lines PS secretion characteristic had been analized.
The rates of phytosiderophore secretion had been tested by using ultra-spectra high performance liquid chromatography (UV-HPLC) and inductively coupled plasma-atomic emission spectrometry (ICP-AES) with 46 corn inbred lines at seedling stage and heading stage. The iron deficiency sensitivity, the phytosiderophores secretion ability and change in different growth stage had been estimated. There was marked correlation existed in phytosiderophore secretion rate under Fe(+) and Fe(-) condition, and a marked correlation, r(Fe+) = 0.7149~(**),existed in Fe(+) rates of phytosiderophore secretion between two test methods. The rates of phytosiderophore secretion of seedling stage and heading stage had marked correlation, r=0.9701~*. Between the phytosiderophores secretion rate in seedling stage and the decrease degree of phytosiderophores in heading stage, there was a marked logarithm correlation r=0.892~*.The corn inbred-lines could be clustered into 4 types with different phytosiderophores secretion rate and iron difficiency sensitivity using the median of Fe(+) phytosiderophores secretion rate [2.836 mg/(g~*per plant~*3h)] and the increment of phytosiderophores secretion rate after being Fe(-) treatment [0.209 mg/(g~*per plant~*3h)] as standard.The HPLC was an exact method in PS testing, but the ICP-AES could be used in material selection. The phytosiderophores secretion characteristic was different at different growth stage. An estimated system had been constructed using PS secretion ability and iron deficiency sensitivity, and corn inbred lines could been divided into four types. The type with low PS secretion rate and high iron difficiency sensitivity are the expected Fe-efficient nutrition breeding material.
3. Genetic analysis of PS secretion characteristic
8 inbred lines which has different PS secretion characteristic each were used to make 28 hybrids by (1/2) n(n-1) diallel crosses design.The PS secretion ability of the hybrids and parents had been analized. It had been realized that there was a highly significantly correlationship existed between the PS secretion ability and the GCA from same parent.The combination with top SCA maybe not the best PS secretion ability combination .Only when both combination SCA and parent GCA were high ,the combiantion with strong PS secretion ability could be got.The PS secretion characteristic was mainly controlled by dominant effect,and the additive effect was proved to be weaken.The genetic model of PS secretion abilitys were proved to match the additive-dominant model, and was controlled by at least 1 major gene. However the iron sesectivity was not,and it has ralative less major gene.The major gene group seemed not to be the same group, hence the dominant effect they controlled were opposition,which is possitive(iron sensitivity),another is negative(PS secretion ability).
4 suppression substractive hybridization
Using the root apex cDNA from Chang7-2 under normal treatment as driver and Chang7-2 under iron deficient treatment as tester, one suppression subrstractive hybridization had been handled. While another hybridizaiton the iron deficiency treatment root apex cDNA from Chang7-2 as driver and zheng 58 as tester had been done. The PCR product from specific expressed cDNA had been gotten. The secific expressed cDNA was associated with the control of iron deficiency and PS secretion ability.
5 Analysis about the gene which controlled corn iron deficiency metabolize
SSH library had been constructed using the PCR product associated with iron dificiency. The lengh of segments which come from library were about 200 bp-900 bp.1230 clones had been choosen out and sequenced, 203 non-repeated sequence information had been got. Among them, there was 9 fragment could be putatived to be related with lower energy regulate after being iron deficiency treatmeng, 3 fragment was hypothetical iron oxidoreductase, 2 fragment may be associated with iron binding protein, 5 fragment may have the function about the membrane trasport,3 fragment was associated with aminotransferase, 5 could be putatived with plant resistant ability and 2 fragment were putative transcriptional regulator. 0257, a most special fragment within all fragment, was putative to has relationship with helix-turn-helix which was different to the bHLH (helix-loose-helix). The function of 0257 may be the negative control and had some touch with the negative dominateffec of genetic control. Besides, 77 fragments may be the new gene sequence, and the other was unknown function. Further reaseach should be done to explain the fragments’s special expression with corn iron deficiency gene control.
禾本科类植物能够由根系分泌麦根酸类物质(Mugineic acids,简称MAs),也叫植物铁载体(Phytosiderophores,简称PS)来活化土壤中难以被植物吸收利用的铁,使之易于被植物吸收利用,从而解决自身铁营养缺乏的问题。
本研究以玉米自交系为材料,对三种玉米麦根酸类物质的测定方法进行比较,同时分析了不同玉米自交系苗期麦根酸类物质的分泌能力和缺铁敏感性,比较了不同生育时期玉米麦根酸分泌能力的差异,研究了玉米麦根酸分泌特性的遗传规律,建立了玉米苗期缺铁胁迫诱导的cDNA文库,并对该cDNA文库的近1000个克隆进行了测序分析。主要研究结果如下:
1筛选建立了玉米麦根酸类物质测定方法
以16种玉米自交系为材料,根据麦根酸类物质的溶铁特性,建立了麦根酸类物质间接测定新方法——ICP-AES法,改良了脱氧麦根酸(DMA)的UV-HPLC直接测定法,并与传统的比色法测定结果进行比较。结果表明,采用ICP-AES法,以Fe_2O_3粉末作为活化对象,与传统采用Fe(OH)_3的测定结果相比,结果稳定、重复性好,说明ICP-AES法是玉米麦根酸类物质分泌量的一种简单、快速的间接测定方法。改良的UV-HPLC法也具有准确、高效的特点,可应用于禾谷类作物DMA分泌量的直接测定。高效液相色谱法是PS测定的准确方法,但在材料选择时可以利用ICP-AES方法来对材料的麦根酸分泌能力进行测定。
2建立了玉米自交系麦根酸分泌特性的评价体系
以46种玉米自交系为材料,利用高效液相色谱法和等离子体法测定了苗期和穗花期缺铁胁迫前后玉米根系的麦根酸分泌速率,评价了不同材料的麦根酸分泌能力、缺铁敏感性和不同生育时期分泌量的变化。结果得出,在苗期缺铁胁迫前后材料的PS分泌能力之间存在着显著相关关系同时两种测定方法之间存在显著相关关系r (Fe+) = 0.7149~(**);麦根酸分泌特性在不同发育时期存在较大差异,四叶一心期和穗花期的测定结果具有显著的相关性r=0.9701~*,穗花期玉米PS分泌速率的下降程度与苗期玉米PS分泌速率之间具有显著的对数相关关系r=0.892~*;利用苗期材料的麦根酸分泌能力和缺铁敏感性建立了麦根酸分泌特性的评价体系,以缺铁胁迫前PS分泌速率的平均数[2.836 mg/(g~*per plant~*3h)]和缺铁胁迫后的PS分泌速率增加量的平均数[0.209 mg/(g~*per plant~*3h)]作为分界线可以将材料分为具有不同PS分泌能力和缺铁敏感性的四种类型:HH,HL,LH,LL,其中具有LH型低PS分泌速率和高缺铁胁迫敏感性的材料是铁营养高效育种的理想材料。
3分析了玉米麦根酸代谢遗传规律
选用八个具有不同MAs分泌能力的玉米自交系,采用半双列杂交的试验设计,分析了杂交后代的麦根酸分泌能力,结果表明亲本麦根酸分泌特性的测定结果与材料自身GCA的相关系数均达到极显著水平,SCA效应值最高的组合并不一定是麦根酸分泌能力最高的组合,只有组合的SCA和双亲的GCA效应均高,才有可能获得麦根酸分泌能力较高的组合,组合的表现与其TCA效应一致。玉米麦根酸类物质的分泌特性以显性效应占优势,遗传加性效应相对较小,麦根酸分泌能力的遗传模型经检验证明符合加性-显性模型,而其缺铁敏感性的遗传模型则不符合。控制麦根酸分泌能力表现显性的主效基因组不少于一组,显性方向为减效;控制麦根酸缺铁敏感性表现显性的主效基因组数较少,显性方向为增效,二者的主效基因组可能不相同。
4建立了缺铁胁迫诱导的抑制性扣除杂交cDNA文库
以缺铁胁迫前的昌7-2根尖cDNA为driver cDNA,缺铁胁迫后的昌7-2根尖cDNA为Tester cDNA;以缺铁胁迫后的昌7-2根尖cDNA为driver cDNA,缺铁胁迫后的郑58根尖cDNA为Tester cDNA,进行两组抑制性扣除杂交,获得了与玉米缺铁胁迫代谢和麦根酸代谢能力差异表达有关的cDNA PCR产物,建立了相关文库。
5分析了玉米缺铁诱导代谢的相关基因
分析已获得的与玉米缺铁胁迫特异性表达的抑制性扣除杂交的PCR产物建立的SSH文库,其包含的片段大小为200bp-900bp。从文库中挑取了1230个克隆进行测序,除去8%的冗余片段共得到非重复的序列203个,其中77个的Blast结果表明它们可能是新发现的序列,除此之外其中:9个片段(0002,0034,0039,0143,0152, 0214, 0355,0375, 1152)可能与缺铁胁迫后植物适应代谢能量降低的调控有关;3个片段(0183,0232,0350)可能与细胞质膜上铁的还原酶有关;2个片段(0061,0622)可能与植物体内或细胞膜上铁离子的结合蛋白有关;5个片段(0094,0155,0208, 0249, 0985,1055)可能与细胞质膜的转运功能有关,3个片段(0234,0362,0387)与氨基酸的转运功能有关,5个片段(0103,0116,0274, 0382,0681)与植物的抗逆性有关,2个片段(0201,1183)与转录调控有关。同时在所有得到片段中, 0257与DNA的促旋酶有关。可能与遗传规律的减效显性作用有关。其余部分片段可能为未知功能片段。这些基因的差异表达与玉米缺铁胁迫的基因调控的关系有待进一步阐明和分析。
Iron (Fe) deficiency chlorosis (lime chlorosis) is a world widespread mineral nutrient deficiency in higher plants, especially corn, sorghum and fruit tree, grown on calcareous soils. Soil amendments and foliar sprays of Fe are not viable economically in the long term to correct Fe deficiency. Therefore, to alleviate Fe chlorosis in the long term, alternative methods, for example breeding of more chlorosis resistant genotypes (cultivars) are required. The recent hotspot in agricultural research is to reach the high yield, quality and efficiency target by the using of plant’s stress tolerance. Graminaceous species can enhance iron (Fe) acquisition from sparingly soluble inorganic Fe (Ⅲ) compounds by release of phytosiderophores (PS) which mobilize Fe (Ⅲ) by chelating.
Using corn inbred lines as material, this study had compared three PS amount test method, analized the PS secretion ability and iron sensitivity at seedling stage.While the different of PS secretion ability between seedling and heading stage and the regulation of PS secretion characteristic’s inherit had been studied. Two SSH cDNA library which one was about the Fe deficiency metobolize regulating, another was about the different PS secretion ability had been constructed.Using the SSH cDNA library which one was about the Fe deficiency metobolize regulating, the cDNA which reduced by Fe deficency had been analized.The main results were summarized as follow:
1 The Phytosiderophores (PS)’s test method had been set up.
Using 16 corn inbred lines,according to the Fe(Ⅲ) chelated characteristic of mugineic acid, a new indirect method of inductively coupled plasma-atomic emission spectrometry (ICP-AES) and a modified direct method of ultra-spectra high performance liquid chromatography (UV-HPLC) for the measurement of MAs or deoxymugineic acid (DMA) were established in corn inbred lines. The comparison of these two methods with traditional spectrometry was also made. The results showed that the ICP-AES method, using Fe_2O_3 as the dissoluble object instead of the traditional Fe (OH)_3, was more stable and repeatalble suggesting that ICP-AES was a simple and quick method for evaluation of the MAs content in corn and other graminaceous crops. The modified UV-HPLC was also precise and efficient, and could be used for the direct mesurement of graminaceous’s deoxymugineic acid.
2 The corn inbred lines PS secretion characteristic had been analized.
The rates of phytosiderophore secretion had been tested by using ultra-spectra high performance liquid chromatography (UV-HPLC) and inductively coupled plasma-atomic emission spectrometry (ICP-AES) with 46 corn inbred lines at seedling stage and heading stage. The iron deficiency sensitivity, the phytosiderophores secretion ability and change in different growth stage had been estimated. There was marked correlation existed in phytosiderophore secretion rate under Fe(+) and Fe(-) condition, and a marked correlation, r(Fe+) = 0.7149~(**),existed in Fe(+) rates of phytosiderophore secretion between two test methods. The rates of phytosiderophore secretion of seedling stage and heading stage had marked correlation, r=0.9701~*. Between the phytosiderophores secretion rate in seedling stage and the decrease degree of phytosiderophores in heading stage, there was a marked logarithm correlation r=0.892~*.The corn inbred-lines could be clustered into 4 types with different phytosiderophores secretion rate and iron difficiency sensitivity using the median of Fe(+) phytosiderophores secretion rate [2.836 mg/(g~*per plant~*3h)] and the increment of phytosiderophores secretion rate after being Fe(-) treatment [0.209 mg/(g~*per plant~*3h)] as standard.The HPLC was an exact method in PS testing, but the ICP-AES could be used in material selection. The phytosiderophores secretion characteristic was different at different growth stage. An estimated system had been constructed using PS secretion ability and iron deficiency sensitivity, and corn inbred lines could been divided into four types. The type with low PS secretion rate and high iron difficiency sensitivity are the expected Fe-efficient nutrition breeding material.
3. Genetic analysis of PS secretion characteristic
8 inbred lines which has different PS secretion characteristic each were used to make 28 hybrids by (1/2) n(n-1) diallel crosses design.The PS secretion ability of the hybrids and parents had been analized. It had been realized that there was a highly significantly correlationship existed between the PS secretion ability and the GCA from same parent.The combination with top SCA maybe not the best PS secretion ability combination .Only when both combination SCA and parent GCA were high ,the combiantion with strong PS secretion ability could be got.The PS secretion characteristic was mainly controlled by dominant effect,and the additive effect was proved to be weaken.The genetic model of PS secretion abilitys were proved to match the additive-dominant model, and was controlled by at least 1 major gene. However the iron sesectivity was not,and it has ralative less major gene.The major gene group seemed not to be the same group, hence the dominant effect they controlled were opposition,which is possitive(iron sensitivity),another is negative(PS secretion ability).
4 suppression substractive hybridization
Using the root apex cDNA from Chang7-2 under normal treatment as driver and Chang7-2 under iron deficient treatment as tester, one suppression subrstractive hybridization had been handled. While another hybridizaiton the iron deficiency treatment root apex cDNA from Chang7-2 as driver and zheng 58 as tester had been done. The PCR product from specific expressed cDNA had been gotten. The secific expressed cDNA was associated with the control of iron deficiency and PS secretion ability.
5 Analysis about the gene which controlled corn iron deficiency metabolize
SSH library had been constructed using the PCR product associated with iron dificiency. The lengh of segments which come from library were about 200 bp-900 bp.1230 clones had been choosen out and sequenced, 203 non-repeated sequence information had been got. Among them, there was 9 fragment could be putatived to be related with lower energy regulate after being iron deficiency treatmeng, 3 fragment was hypothetical iron oxidoreductase, 2 fragment may be associated with iron binding protein, 5 fragment may have the function about the membrane trasport,3 fragment was associated with aminotransferase, 5 could be putatived with plant resistant ability and 2 fragment were putative transcriptional regulator. 0257, a most special fragment within all fragment, was putative to has relationship with helix-turn-helix which was different to the bHLH (helix-loose-helix). The function of 0257 may be the negative control and had some touch with the negative dominateffec of genetic control. Besides, 77 fragments may be the new gene sequence, and the other was unknown function. Further reaseach should be done to explain the fragments’s special expression with corn iron deficiency gene control.
引文
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