水稻黄胚乳突变体JMW的基因克隆和功能研究
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摘要
乙醛脱氢酶(ALDH)是生物界中常见的一种酶类。依赖于辅酶NAD(P)+,乙醛脱氢酶负责把代谢中各种醛类氧化成对应的羧酸。基于蛋白序列的差异,所有已知乙醛脱氢酶分为22个家族,不同家族的ALDH基因根据底物的不同,参与各种代谢过程。植物中包含12个ALDH家族,但与人类中的研究相比,植物ALDH基因的研究还很不充分,大多数研究只局限于其表达模式分析。ALDH7在动植物中高度保守,人类中的ALDH7A1基因参与赖氨酸降解途径中α-氨基己二酸半醛(AASA)的氧化过程。植物ALDH7在豌豆,油菜,苹果,藓类,高粱和拟南芥等物种中已得到克隆,它们的表达与果实的成熟和植物所受的渗透压逆境相关,但其具体作用还待进一步研究。近来,水稻的ALDH7基因被证实参与种子老化相关过程,该基因的T-DNA突变体胚乳呈黄色,所积累的色素有可能是美拉德反应(Millard reaction)的中间产物。另外,日本品种Hatsuyamabuki的稻米也是黄色胚乳,含有水稻突变体酸的新色素类化合物,但水稻突变体酸与ALDH7基因参与的生化途径的关系还缺乏系统的研究。本研究以一个黄色胚乳的天然突变体为材料,定位和克隆了OsALDH7基因,研究了该基因的表达模式,并且对该基因参与的具体生化途径做了部分探讨。主要结论如下：
1.从一个水稻低谷蛋白品种(W3660)中,鉴定出一个天然的黄色胚乳突变体,该突变体除了胚乳颜色和种子蛋白含量之外,其它农艺性状与野生型没有显著差别。色素具有水溶性,在各种有机溶剂中溶解性不高。为了和前人通过转基因方法得到的golden rice区别开来,我们将这个突变体命名为jmw(金米王)。突变体色素的积累主要发生在种子发育的后期,并且受发育时期和储存时间的影响。遗传分析表明该表型受单个隐性核基因控制,用图位克隆的方法将基因定位于9号染色体26kb包含6个预测基因的区域内。
2.通过测序和比较,发现定位区间内预测基因(LOC_Os09g26880)的第10外显子的存在单碱基突变,从而造成该基因的转录提前中止。该基因属于乙醛脱氢酶第7家族,命名为OsALDH7.进一步通过互补实验证实了OsALDH7的突变是产生黄色胚乳的主要原因。OsALDH7的表达受干旱,高盐和ABA处理等逆境的诱导,叶片的诱导表达比根部更明显,而且突变体jmw的抗旱能力比野生型W3660要差,显示OsALDH7在植物对渗透压逆境的抵御过程中起重要作用。同时通过检测野生型/突变体中ALDH家族其他基因的表达情况,发现突变体中大多数的被测基因表达有所提高,推测ALDH家族基因之间的表达有一定的补偿效应。
3.通过高效液相色谱(HPLC)检测野生型/突变体中突变体酸A的含量,与前人的结果一致。随着种子的成熟和老化,突变体酸A的积累更加明显,与表型的观察结果相符。进一步检测赖氨酸代谢途径中的相关基因(LOR/SDH, AK, DHPS)的表达和代谢产物(酵母氨酸)在野生型/突变体中的含量,发现有规律性的变化,OsALDH7基因的突变造成赖氨酸降解及其代谢产物酵母氨酸的积累；在过表达OsALDH7基因的酵母中,游离赖氨酸含量大幅下降,而赖氨酸的代谢产物-酵母氨酸及其谷氨酸含量上升。综合上述实验结果,推测OsALDH7很可能参与水稻中赖氨酸的降解途径,缺失OsALDH7,赖氨酸可能通过未知的支路途径转化成突变体酸A等色素物质。
Aldehyde dehydrogenase (ALDH) represents a superfamily of NAD(P)+-dependent enzymes that oxidize aldehydes into their corresponding carboxylic acids. ALDH comprises at least22families according to protein sequence identity, and plays derivate functions based on their substrates. In plant, twelve families were present and a few of them are studied in detail comparing the research in human.Most of the studies of plant ALDH genes were just restricted to the expression patterns. The family ALDH7is highly conserved in whatever animals or plants. ALDH7A1, ALDH7homolog in human, catalyzes the oxidation of α-amino adipic semialdehyde (AASA) in lysine degradation. As yet, the expressions of ALDH7in plants such as garden pea, brassica, apple, tortula ruralis, sorghum and Arabidopsis were found to be associated with seed maturation and osmotic stress, but their exact functions were still remain to be clarified. Recently, ALDH7in rice was found to play a role during seed storage, resulted in an unidentified yellow pigment accumulation of the mutant endosperm, which are probably products of maillard reaction. On the other hand, a new kind of pigment compounds called oryzamutaic acids were detected in var. Hatsuyamabuki. However, no information is yet available for our understanding the production of oryzamutaic acids. In this study, using a natural mutant of yellow endosperm,OsALDH7, the gene responsible for the phenotype, was cloned. Besides its expression pattern, the possible role of OsALDH7in lysine degradation was further investigated. The main conclusions are as follows:
1. A natural mutant of yellow endosperm was identified from a low-glutelin var. W3660, which named as jmw (new golden rice in Chinese) contrast to the artificial produced golden rice. The mutant was identical to the wildtype except for the color and the protein content in endosperm. Yellow pigments could be soluble in water but not in organic solvents. Genetic analysis showed that the yellow endosperm caused by one pair of recessive nuclear gene mutation. Using a F2group, the mutated gene was mapped to an interval of～26-kb on Chromosome9, which included six putative genes.
2. After sequencing and comparing the intervals between W3660and jmw, a mutant in the exon10of the putative gene LOC_Os09g26880, named OsALDH7, was detected. Complement experiment indicated that OsALDH7was the correct gene responsible for the production of yellow pigment. The expression of OsALDH7in rice may be induced under the treatments of dehydration, salinity and ABA especially in leaf, and the drought resisting was weaker in the mutant, indicating OsALDH7plays an important role in the plant defense to the osmotic stresses. Some ALDH genes express much higher in the mutant than the wild type, indicating the possible compensating pattern in ALDH family.
3. The main pigment in our mutant was confirmed as oryzamutaic acid A, and the contents of oryzamutaic acid A in W3660and jmw were detected by high performance liquid chromatography (HPLC), which could confirm the conclusion. The pigment accumulation was more obvious along with seed ripen or aging, which is consistent with to the phenotype. Comparing the ALDH7function in human, its probable role on lysine catabolism was also investigated. The expression of related gene{LOR/SDH, AK, DHPS) and the contents of intermediate compounds (saccharopine) in lysine metabolism were detected in W3660and jmw. The mutation of OsALDH7induced the accumulation of saccharopine but the degradation of lysine. Moreover, the over-expressing of OsALDH7in yeast could cause great decease of lysine and an increase of glutamate, which is the product of lysine degradation. OsALDH7in rice probably participate in the degradation of lysine, and without OsALDH7, lysine might be transformed to the pigment like oryzamutaic acid A through some unknown processes
1.从一个水稻低谷蛋白品种(W3660)中,鉴定出一个天然的黄色胚乳突变体,该突变体除了胚乳颜色和种子蛋白含量之外,其它农艺性状与野生型没有显著差别。色素具有水溶性,在各种有机溶剂中溶解性不高。为了和前人通过转基因方法得到的golden rice区别开来,我们将这个突变体命名为jmw(金米王)。突变体色素的积累主要发生在种子发育的后期,并且受发育时期和储存时间的影响。遗传分析表明该表型受单个隐性核基因控制,用图位克隆的方法将基因定位于9号染色体26kb包含6个预测基因的区域内。
2.通过测序和比较,发现定位区间内预测基因(LOC_Os09g26880)的第10外显子的存在单碱基突变,从而造成该基因的转录提前中止。该基因属于乙醛脱氢酶第7家族,命名为OsALDH7.进一步通过互补实验证实了OsALDH7的突变是产生黄色胚乳的主要原因。OsALDH7的表达受干旱,高盐和ABA处理等逆境的诱导,叶片的诱导表达比根部更明显,而且突变体jmw的抗旱能力比野生型W3660要差,显示OsALDH7在植物对渗透压逆境的抵御过程中起重要作用。同时通过检测野生型/突变体中ALDH家族其他基因的表达情况,发现突变体中大多数的被测基因表达有所提高,推测ALDH家族基因之间的表达有一定的补偿效应。
3.通过高效液相色谱(HPLC)检测野生型/突变体中突变体酸A的含量,与前人的结果一致。随着种子的成熟和老化,突变体酸A的积累更加明显,与表型的观察结果相符。进一步检测赖氨酸代谢途径中的相关基因(LOR/SDH, AK, DHPS)的表达和代谢产物(酵母氨酸)在野生型/突变体中的含量,发现有规律性的变化,OsALDH7基因的突变造成赖氨酸降解及其代谢产物酵母氨酸的积累；在过表达OsALDH7基因的酵母中,游离赖氨酸含量大幅下降,而赖氨酸的代谢产物-酵母氨酸及其谷氨酸含量上升。综合上述实验结果,推测OsALDH7很可能参与水稻中赖氨酸的降解途径,缺失OsALDH7,赖氨酸可能通过未知的支路途径转化成突变体酸A等色素物质。
Aldehyde dehydrogenase (ALDH) represents a superfamily of NAD(P)+-dependent enzymes that oxidize aldehydes into their corresponding carboxylic acids. ALDH comprises at least22families according to protein sequence identity, and plays derivate functions based on their substrates. In plant, twelve families were present and a few of them are studied in detail comparing the research in human.Most of the studies of plant ALDH genes were just restricted to the expression patterns. The family ALDH7is highly conserved in whatever animals or plants. ALDH7A1, ALDH7homolog in human, catalyzes the oxidation of α-amino adipic semialdehyde (AASA) in lysine degradation. As yet, the expressions of ALDH7in plants such as garden pea, brassica, apple, tortula ruralis, sorghum and Arabidopsis were found to be associated with seed maturation and osmotic stress, but their exact functions were still remain to be clarified. Recently, ALDH7in rice was found to play a role during seed storage, resulted in an unidentified yellow pigment accumulation of the mutant endosperm, which are probably products of maillard reaction. On the other hand, a new kind of pigment compounds called oryzamutaic acids were detected in var. Hatsuyamabuki. However, no information is yet available for our understanding the production of oryzamutaic acids. In this study, using a natural mutant of yellow endosperm,OsALDH7, the gene responsible for the phenotype, was cloned. Besides its expression pattern, the possible role of OsALDH7in lysine degradation was further investigated. The main conclusions are as follows:
1. A natural mutant of yellow endosperm was identified from a low-glutelin var. W3660, which named as jmw (new golden rice in Chinese) contrast to the artificial produced golden rice. The mutant was identical to the wildtype except for the color and the protein content in endosperm. Yellow pigments could be soluble in water but not in organic solvents. Genetic analysis showed that the yellow endosperm caused by one pair of recessive nuclear gene mutation. Using a F2group, the mutated gene was mapped to an interval of～26-kb on Chromosome9, which included six putative genes.
2. After sequencing and comparing the intervals between W3660and jmw, a mutant in the exon10of the putative gene LOC_Os09g26880, named OsALDH7, was detected. Complement experiment indicated that OsALDH7was the correct gene responsible for the production of yellow pigment. The expression of OsALDH7in rice may be induced under the treatments of dehydration, salinity and ABA especially in leaf, and the drought resisting was weaker in the mutant, indicating OsALDH7plays an important role in the plant defense to the osmotic stresses. Some ALDH genes express much higher in the mutant than the wild type, indicating the possible compensating pattern in ALDH family.
3. The main pigment in our mutant was confirmed as oryzamutaic acid A, and the contents of oryzamutaic acid A in W3660and jmw were detected by high performance liquid chromatography (HPLC), which could confirm the conclusion. The pigment accumulation was more obvious along with seed ripen or aging, which is consistent with to the phenotype. Comparing the ALDH7function in human, its probable role on lysine catabolism was also investigated. The expression of related gene{LOR/SDH, AK, DHPS) and the contents of intermediate compounds (saccharopine) in lysine metabolism were detected in W3660and jmw. The mutation of OsALDH7induced the accumulation of saccharopine but the degradation of lysine. Moreover, the over-expressing of OsALDH7in yeast could cause great decease of lysine and an increase of glutamate, which is the product of lysine degradation. OsALDH7in rice probably participate in the degradation of lysine, and without OsALDH7, lysine might be transformed to the pigment like oryzamutaic acid A through some unknown processes
引文
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