水稻响应灌浆初期高温的转录组和蛋白质组研究
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摘要
水稻起源于低纬度地区,对高温气候具有一定的忍耐能力,但温度超过水稻最适生长的临界值时,水稻的正常生长便会受到影响。在我国,水稻生产中的高温热害主要发生在长江流域及其以南的双季稻区；通常在7月上中旬,控制这些地区的副高气压脊线往往呈停滞状态,于是在较长时间内维持高温天气。而此时正值双季早稻灌浆期,水稻灌浆期遇高温热害不仅限制稻米品质而且降低籽粒充实度,从而造成水稻品质和产量损失。双季早稻又是我国重要的一茬粮食作物,也是南方各省中不可替代的优势粮食作物。因此,加强水稻耐热的分子机理研究,加速灌浆期耐热的优质早稻品种选育,对保持我国水稻产量及品质的稳定,从而保障国家的粮食安全具有重要意义。
前期研究中本课题组以耐热水稻N22和热敏感水稻协青早B为亲本,通过杂交、回交、自交和定向选育获得了791个回交重组自交株系。本文以该回交重组自交系为材料,
(1)采用表型性状鉴定和基因组多态性分析,鉴定了该重组自交系的灌浆耐热性和部分形态指标以及基因组多态性差异；(2)于水稻抽穗后第10天进行3、6、9、24、48、72、96和120小时的高温处理后,分析了高温热害和常温对照条件下与水稻剑叶光合作用、细胞膜氧化和渗透调节有关的生理指标；(3)于水稻抽穗后第10天进行3、6、9、24、48和72小时的高温处理后,采用cDNA-AFLP基因差异显示技术分析了水稻籽粒中与高温热害相关的差异表达基因；(4)于水稻抽穗后第10天进行24、72和120小时的高温处理后,运用比较蛋白质组学鉴定了水稻籽粒中与灌浆初期高温热害有关的差异表达蛋白质。研究取得如下结果：
(1)从791个重组自交系中鉴定并获得了2个基因组差异较小而灌浆初期耐热性存在极显著差异的2个水稻纯系XN0437T(耐热)和XN0437S(热敏感)。
(2)高温热害降低水稻剑叶叶绿素含量和光合作用效率,加剧了细胞膜氧化作用、增加了细胞膜透性的同时改变了细胞内环境,热敏感水稻纯系XN0437S受影响的程度大于耐热水稻纯系XN0437T。
(3)获得了54个与水稻灌浆期耐热性有关的差异表达基因,其中27个基因在GenBank数据库中有同源序列,另外27个为新发现的与水稻灌浆初期耐热性有关的基因。27个功能已知的差异表达基因中,与逆境胁迫有关的基因表现上调表达,且在耐热水稻纯系XN0437T中的上调表达丰度大于在热敏感水稻纯系XN0437S中的上调表达丰度；与代谢、转录、生物合成和氧化作用有关的基因也表现上调表达,但是在耐热水稻纯系中的上调表达丰度小于在热敏感水稻纯系中的上调表达丰度；与转录、生物合成、光合作用、甲基化和物质转运有关的基因表现下调表达,在热敏感水稻纯系中的下调表达丰度大于在耐热水稻纯系中的下调表达丰度。
(4)获得了23个与水稻灌浆期耐热性有关的差异表达蛋白质,仅9个蛋白在数据库Matrixscience中具有相似蛋白质,其功能分别与氧化作用、生物合成、基因转录、磷酸化和淀粉合成有关；高温热害条件下,与氧化作用和生物合成相关的蛋白表现上调表达,而与基因转录、磷酸化和淀粉合成酶相关的蛋白表现下调表达；另有12个功能未知的蛋白质在高温热害条件下表现上调表达、2个功能未知的蛋白质在高温热害条件下表现下调表达。
研究结果表明：由于灌浆初期高温热害降低了水稻剑叶光合作用效率,加强了细胞膜氧化而造成细胞膜透性增加的同时改变了细胞内环境,最终导致向籽粒运输的光合产物和储藏物质减少是高温热害降低水稻籽粒充实度的生理原因。灌浆初期高温导致水稻籽粒中与储藏物质合成、转录相关的基因和与转录、磷酸化、淀粉合成有关的蛋白质下调表达；与逆境胁迫有关的基因和与氧化作用、(部分)生物合成有关的蛋白质上调表达,是灌浆初期高温造成水稻籽粒充实下降的分子生物学基础。
本文鉴定的54个与水稻响应灌浆初期高温热害有关的差异表达基因(片段)和23个差异表达蛋白质,为进一步研究水稻灌浆期耐热的分子机理奠定了研究基础。
Originating from low latitude areas, rice can survive certain high temperatures, but its normal growth will be impaired once the temperature is higher than the threshold of its growth-optimal temperature. Unfortunately, it is highly likely that crops will have to adjust to even higher temperatures in the near future because of global climate change. In China, high-temperature damage happens mainly in the zone of double-cropping early-season rice at Yangtze River Valley and some areas to its south from early to middle July, because the ridge line of western pacific subtropical anticyclone located these areas at the period, which cause long hot weather in the middle and lower reaches of Yangtze River. This period happens to be the early grain-filling stage of double-cropping early-season rice in the area around the Yangtze River Valley, which often results in low grain quantity and yield loss. Double-cropping early-season rice is an important and irreplaceable food crop in southern China. Therefore, strengthening the researches on molecular mechanisms of heat tolerant characteristic in rice as well as breeding rice cultivars with heat tolerant characteristic at grain-filling stage plays a significant role in maintaining rice yield stability and guaranteeing the human food security.
In our previous work, a recombinant inbred line (RIL) related to heat-tolerance at rice grain-filling stage and derived from Xieqingzao B/N22were inbred. In present study,791rice lines of the RIL were used as plant materials;1. The heat-tolerant and physiological characteristics of rice at grain-filling stage were determined and the genomic polymorphism were analyzed by SSR molecular markers, among the791RILs;2. The physiological indexes of flag leaves related to photosynthesis, oxidation of cell membrane and osmotic adjustment were analyzed and compared between treatment and control of both XN0437S and XN0437T rice lines;3. differentially expressed genes related to heat-tolerant characteristic of rice caryopsis were analyzed and compared between treatment and control of both XN0437S and XN0437T rice lines by cDNA-AFLP;4. differentially expressed proteins related to heat-tolerant characteristic of rice caryopsis were analyzed and compared between treatment and control of both XN0437S and XN0437T rice lines by proteomics based on two-dimensional electrophoresis (2-DE). The results are as follow.
1. Two rice lines. XN0437T (heat-tolerant) and XN0437S (heat-sensitive) with similar genome but significant difference in heat-tolerant trait at grain-filling stage, were obtained from the791RILs.
2. Chlorophyll content and photosynthesis rate of rice flag leaves were decreased, while the matters content of cell membrane oxidation and permeability of plasma membrane were increased, under high temperature stress, which resulted in a change of intracellular environment, but the influence of the heat sensitive rice was more severe than that of the heat tolerant rice.
3. Fifty-four differentially expressed gene fragments from rice caryopsis were obtained by cDNA-AFLP. Among them,27genes had homologous genes in the GenBank database and another27gene fragments were newly identified. Further analysis with RT-PCR indicated that genes involved in adverse stress were showed up-regulation and the up-regulated degree in the heat-tolerant rice line XN0437T was higher than that of the heat-sensitive rice line XN0437S, while the other up-regulated genes were involved in metabolism, transcription, biosynthesis and oxidation, and the up-regulated degree of the heat-tolerant rice line XN0437T was lower than that of the heat-sensitive rice line XN0437S. Genes involved in transcription, biosynthesis, photosynthesis, methylation and substance transportation were down-regulated, and the down-regulated degree in the heat-sensitive rice line XN0437T was higher than that of the heat-tolerant rice line XN0437S.
4. Twenty-three differentially expressed proteins from rice caryopsis were obtained after a proteomics study, but only9proteins had homologous proteins in the protein database of Matrixscience. Functions of the homologous proteins included oxidation, biosynthesis, gene transcription, phosphorylation and starch synthesis. Proteins involved in oxidation and biosynthesis showed up-regulation, while proteins involved in transcription, phosphorylation and starch synthesis showed down-regulation. There are12and2unknown proteins that showed up and down regulation respectively under high temperature stress.
The research results indicated that physiological cause of rice plumpness decrease was that photo synthetic products and storage matters transported to plumpness were reduced, due to the decrease of photosynthesis rate and the increase of cell membrane oxidation and permeability of plasma membrane. Molecular biological cause of rice plumpness decreased was that genes involved in biosynthesis of storage products, and genes involved in transcription, phosphorylation and starch biosynthesis were down-regulated, while that genes involved in adverse stress and proteins encoded by genes involved in oxidation and biosynthesis were up-regulated.
The54candidate genes and23candidate proteins identified in the present study will be helpful for elucidating the molecular mechanism on rice adapting high temperature stress.
前期研究中本课题组以耐热水稻N22和热敏感水稻协青早B为亲本,通过杂交、回交、自交和定向选育获得了791个回交重组自交株系。本文以该回交重组自交系为材料,
(1)采用表型性状鉴定和基因组多态性分析,鉴定了该重组自交系的灌浆耐热性和部分形态指标以及基因组多态性差异；(2)于水稻抽穗后第10天进行3、6、9、24、48、72、96和120小时的高温处理后,分析了高温热害和常温对照条件下与水稻剑叶光合作用、细胞膜氧化和渗透调节有关的生理指标；(3)于水稻抽穗后第10天进行3、6、9、24、48和72小时的高温处理后,采用cDNA-AFLP基因差异显示技术分析了水稻籽粒中与高温热害相关的差异表达基因；(4)于水稻抽穗后第10天进行24、72和120小时的高温处理后,运用比较蛋白质组学鉴定了水稻籽粒中与灌浆初期高温热害有关的差异表达蛋白质。研究取得如下结果：
(1)从791个重组自交系中鉴定并获得了2个基因组差异较小而灌浆初期耐热性存在极显著差异的2个水稻纯系XN0437T(耐热)和XN0437S(热敏感)。
(2)高温热害降低水稻剑叶叶绿素含量和光合作用效率,加剧了细胞膜氧化作用、增加了细胞膜透性的同时改变了细胞内环境,热敏感水稻纯系XN0437S受影响的程度大于耐热水稻纯系XN0437T。
(3)获得了54个与水稻灌浆期耐热性有关的差异表达基因,其中27个基因在GenBank数据库中有同源序列,另外27个为新发现的与水稻灌浆初期耐热性有关的基因。27个功能已知的差异表达基因中,与逆境胁迫有关的基因表现上调表达,且在耐热水稻纯系XN0437T中的上调表达丰度大于在热敏感水稻纯系XN0437S中的上调表达丰度；与代谢、转录、生物合成和氧化作用有关的基因也表现上调表达,但是在耐热水稻纯系中的上调表达丰度小于在热敏感水稻纯系中的上调表达丰度；与转录、生物合成、光合作用、甲基化和物质转运有关的基因表现下调表达,在热敏感水稻纯系中的下调表达丰度大于在耐热水稻纯系中的下调表达丰度。
(4)获得了23个与水稻灌浆期耐热性有关的差异表达蛋白质,仅9个蛋白在数据库Matrixscience中具有相似蛋白质,其功能分别与氧化作用、生物合成、基因转录、磷酸化和淀粉合成有关；高温热害条件下,与氧化作用和生物合成相关的蛋白表现上调表达,而与基因转录、磷酸化和淀粉合成酶相关的蛋白表现下调表达；另有12个功能未知的蛋白质在高温热害条件下表现上调表达、2个功能未知的蛋白质在高温热害条件下表现下调表达。
研究结果表明：由于灌浆初期高温热害降低了水稻剑叶光合作用效率,加强了细胞膜氧化而造成细胞膜透性增加的同时改变了细胞内环境,最终导致向籽粒运输的光合产物和储藏物质减少是高温热害降低水稻籽粒充实度的生理原因。灌浆初期高温导致水稻籽粒中与储藏物质合成、转录相关的基因和与转录、磷酸化、淀粉合成有关的蛋白质下调表达；与逆境胁迫有关的基因和与氧化作用、(部分)生物合成有关的蛋白质上调表达,是灌浆初期高温造成水稻籽粒充实下降的分子生物学基础。
本文鉴定的54个与水稻响应灌浆初期高温热害有关的差异表达基因(片段)和23个差异表达蛋白质,为进一步研究水稻灌浆期耐热的分子机理奠定了研究基础。
Originating from low latitude areas, rice can survive certain high temperatures, but its normal growth will be impaired once the temperature is higher than the threshold of its growth-optimal temperature. Unfortunately, it is highly likely that crops will have to adjust to even higher temperatures in the near future because of global climate change. In China, high-temperature damage happens mainly in the zone of double-cropping early-season rice at Yangtze River Valley and some areas to its south from early to middle July, because the ridge line of western pacific subtropical anticyclone located these areas at the period, which cause long hot weather in the middle and lower reaches of Yangtze River. This period happens to be the early grain-filling stage of double-cropping early-season rice in the area around the Yangtze River Valley, which often results in low grain quantity and yield loss. Double-cropping early-season rice is an important and irreplaceable food crop in southern China. Therefore, strengthening the researches on molecular mechanisms of heat tolerant characteristic in rice as well as breeding rice cultivars with heat tolerant characteristic at grain-filling stage plays a significant role in maintaining rice yield stability and guaranteeing the human food security.
In our previous work, a recombinant inbred line (RIL) related to heat-tolerance at rice grain-filling stage and derived from Xieqingzao B/N22were inbred. In present study,791rice lines of the RIL were used as plant materials;1. The heat-tolerant and physiological characteristics of rice at grain-filling stage were determined and the genomic polymorphism were analyzed by SSR molecular markers, among the791RILs;2. The physiological indexes of flag leaves related to photosynthesis, oxidation of cell membrane and osmotic adjustment were analyzed and compared between treatment and control of both XN0437S and XN0437T rice lines;3. differentially expressed genes related to heat-tolerant characteristic of rice caryopsis were analyzed and compared between treatment and control of both XN0437S and XN0437T rice lines by cDNA-AFLP;4. differentially expressed proteins related to heat-tolerant characteristic of rice caryopsis were analyzed and compared between treatment and control of both XN0437S and XN0437T rice lines by proteomics based on two-dimensional electrophoresis (2-DE). The results are as follow.
1. Two rice lines. XN0437T (heat-tolerant) and XN0437S (heat-sensitive) with similar genome but significant difference in heat-tolerant trait at grain-filling stage, were obtained from the791RILs.
2. Chlorophyll content and photosynthesis rate of rice flag leaves were decreased, while the matters content of cell membrane oxidation and permeability of plasma membrane were increased, under high temperature stress, which resulted in a change of intracellular environment, but the influence of the heat sensitive rice was more severe than that of the heat tolerant rice.
3. Fifty-four differentially expressed gene fragments from rice caryopsis were obtained by cDNA-AFLP. Among them,27genes had homologous genes in the GenBank database and another27gene fragments were newly identified. Further analysis with RT-PCR indicated that genes involved in adverse stress were showed up-regulation and the up-regulated degree in the heat-tolerant rice line XN0437T was higher than that of the heat-sensitive rice line XN0437S, while the other up-regulated genes were involved in metabolism, transcription, biosynthesis and oxidation, and the up-regulated degree of the heat-tolerant rice line XN0437T was lower than that of the heat-sensitive rice line XN0437S. Genes involved in transcription, biosynthesis, photosynthesis, methylation and substance transportation were down-regulated, and the down-regulated degree in the heat-sensitive rice line XN0437T was higher than that of the heat-tolerant rice line XN0437S.
4. Twenty-three differentially expressed proteins from rice caryopsis were obtained after a proteomics study, but only9proteins had homologous proteins in the protein database of Matrixscience. Functions of the homologous proteins included oxidation, biosynthesis, gene transcription, phosphorylation and starch synthesis. Proteins involved in oxidation and biosynthesis showed up-regulation, while proteins involved in transcription, phosphorylation and starch synthesis showed down-regulation. There are12and2unknown proteins that showed up and down regulation respectively under high temperature stress.
The research results indicated that physiological cause of rice plumpness decrease was that photo synthetic products and storage matters transported to plumpness were reduced, due to the decrease of photosynthesis rate and the increase of cell membrane oxidation and permeability of plasma membrane. Molecular biological cause of rice plumpness decreased was that genes involved in biosynthesis of storage products, and genes involved in transcription, phosphorylation and starch biosynthesis were down-regulated, while that genes involved in adverse stress and proteins encoded by genes involved in oxidation and biosynthesis were up-regulated.
The54candidate genes and23candidate proteins identified in the present study will be helpful for elucidating the molecular mechanism on rice adapting high temperature stress.
引文
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