空间诱导水稻蛋白变化特征及空间辐射地基模拟诱因分析
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摘要
为了探索空间环境的生物学效应机制,本文以模式植物水稻为材料,经过多次空间飞行,考察了不同水稻品系当代及第2代植株蛋白变化特征;并通过地基模拟重离子辐射基于水稻的蛋白表达谱分析了空间的辐射诱因;利用多蘖矮突变体的蛋白表达差异分析对空间诱导水稻分蘖突变的机制进行了探索。
利用二维荧光差异显示双向电泳(2-D DIGE)技术获取了2次空间飞行当代、第2代和地基模拟辐射后多个水稻品系以及突变体材料分蘖期的99张双向电泳蛋白表达图谱,建立了包含所有品系共同表达的蛋白点以及品系特异表达蛋白点合集的群体,形成了标准图谱;以该标准图谱为统一范围使不同状态下的大量蛋白表达数据存在定性和定量上的可比性。串联质谱鉴定出204个差异表达蛋白,参与的生物功能涵盖代谢、调节、刺激应答、细胞活动和生殖。
为分析空间飞行当代水稻蛋白变化特征及在子代的遗传特性,利用聚类、主成分分析等方法归纳并比较了不同空间飞行当代及第2代材料与地面对照之间的蛋白表达差异。发现卫星搭载与航天飞船的空间飞行能够使当代水稻叶片蛋白表达特征发生改变,但在不同类型的空间飞行中存在变化程度的区别:飞行时间越长,受空间辐射环境影响越大的材料,蛋白表达谱的差异就越大。从差异蛋白分子生物功能分布上分析了空间环境对水稻细胞生理生化功能的影响,空间飞行当代引起易变蛋白的功能主要包括核酸代谢、光反应、蛋白降解、抗真菌、抗胁迫和氨基酸代谢。空间飞行环境对水稻当代蛋白表达特征的改变在相应的第2代植株中可以发生延续和回复,回复与否以及回复的程度与水稻的品系敏感性相关。细胞壁形成、核酸代谢、蛋白降解、逆转录、氨基酸代谢、磷酸戊糖途径和抗胁迫功能的应答易在第2代回复;参与种子成熟、蛋白折叠、糖酵解、脂类合成、糖原合成和三羧酸循环的蛋白分子仅在第2代时产生应答。空间环境的生物学效应能够体现在蛋白表达层面上。
发现了一些对空间环境敏感的蛋白分子,包括核苷二磷酸激酶1、Rieske铁-硫蛋白前体、半胱氨酸蛋白酶、异黄酮还原酶相似蛋白、S-类核糖核酸酶以及参与蛋白折叠的分子伴侣,它们能够作为水稻对空间环境应答的潜在蛋白分子标志物。
空间辐射是一个非常复杂的复合高能粒子环境。为寻找空间环境的离子诱因,在地面模拟了空间2mGy剂量下不同能量的离子辐射,采用传能线密度(LET)分别为13.3keV/μm的12C离子、30keV/μm的12C离子、31keV/μm的20Ne离子、62.2keV/μm的12C离子和500keV/μm的56Fe离子对水稻种子进行处理,并与返回式卫星搭载的水稻蛋白表达模式进行比较。通过聚类、主成分分析和相关性分析后发现:LET-62.2keV/μm的12C离子辐射蛋白表达特征与空间环境最为类似。为进一步分析空间辐射环境的剂量效应,利用与空间效应相近的LET-62.2keV/μm的12C离子进行了不同累积剂量的辐射(剂量分别为2mGy、20mGy、200mGy、2000mGy),对辐射蛋白表达谱与空间材料进行比较后发现2mGy剂量的辐射效应与空间环境最为接近,证实了空间辐射的低剂量效应。通过基于蛋白表达谱的分析建立了新的空间环境辐射诱因效应评价体系。
对空间诱变多蘖矮杆突变体R955及对照植株3个营养生长期叶片总蛋白的分离及定量分析发现,能量代谢、光合作用、蛋白代谢、氮元素同化、氨基酸代谢以及胁迫应答等过程均参与了空间环境诱导的突变分蘖发育。对差异蛋白在不同时期相对表达量的K-均值聚类分析显示不同功能蛋白表现不同表达模式;二元方差分析筛选出S-类核糖核酸酶可能与水稻分蘖突变性状直接关联。相关结果为空间环境诱发水稻产生分蘖突变的机制提供了新的见解。
In order to study the mechanisms of biological effects of space environment, in the present dissertation, the model plant rice was chosen as experimental material. After several seed spaceflights, proteomic expression profiles of different rice variety plants in 1st and 2nd generations were analysed. Space radiation inducement investigation through protein expression profile analysis based on on-ground simulated radiation experiments was also performed. In addition, mechanisms of tillering mutation induced by spaceflight were also explored by differential protein expression analysis of the tillering dwarf mutant.
Using two-dimensional fluorescent differential electrophoresis (2-D DIGE) technology, 99 protein expression 2-D patterns were acquired from tillering rice plants of different varieties after two kinds of spaceflights in 1st and 2nd generations, after on-ground simulated radiation treatments and from the mutant variety. Standard gel image pattern including groups of proteins that co-expressed in all rice varieties and particularly expressed in every variety was established. The pattern was used as a normalized standard for comparing the large number of protein expression data in different status qualitively and quantitively. Tandem mass spectrometry identified 204 differentially expressed proteins that involved in biological processes including metabolic process, regulation of biological process, response to stimulus, cellular process and reproduction.
For investigation of protein expression profiles of rice plants after spaceflights and the hereditary capacity in the offspring plants, clustering and principal component analysis were performed to summarize and compare the protein expression variations between the space samples and their corresponding ground controls in different spaceflights and in 1st and 2nd generations. Results indicated that protein expression profiles were changed after seed space environment exposures onboard the satellite or space craft, but there were variations in the degree of difference in different types of flights: the longer the flight duration lasted, which meant that rice seeds were affected by space radiation environment heavier, the greater differences existed in protein expression profiles. Distribution of biological processes of differentially expressed proteins indicated that physiological and biochemical changes of rice cells were induced by space environment. The biological processes impacted in 1st generation plants after flights were nucleic acid metabolic process,light reaction,proteolysis,defense response to fungus,response to stress with amino acid and derivative metabolism. Alterations of protein expression profiles in 1st generation would resume or recover in the corresponding 2nd generation plants, the degree of recovery was associated with the sensitivity of the rice variety to environment. Changes of proteins involving in biological processes such as cell wall organization, nucleic acid metabolic process, proteolysis, RNA-dependent DNA replication, amino acid and derivative metabolism, pentose-phosphate shunt, response to stress tended to recover in 2nd generation plants; while seed maturation, protein folding, glycolysis, lipid biosynthetic process, glycogen biosynthetic process and tricarboxylic acid cycle related proteins only differentially expressed in 2nd generation plants. Therefore,the biological effects of space environment could be reflected in protein expression level.
Some of the space-environment-sensitive proteins were found, including nucleoside diphosphate kinase 1, Rieske (Fe-S) protein precursor, cysteine protease, isoflavone reductase-like protein, S-like RNase and molecular chaperones participating in protein folding. They could be used as potential biomarkers that would answer to the stimulus from space environment.
Spaceflight represents a very complex environmental condition with highly ionizing radiations (HZE). To further investigate the incentives of ion effects in space environment, we performed on-ground simulated HZE particle radiations to rice seeds with the same dose (2mGy) as spaceflight but different liner energy transfer (LET) values (13.3keV/μm-12C, 30keV/μm-12C, 31keV/μm-20Ne, 62.2keV/μm-12C, 500keV/μm-56Fe) then compared the protein expression profiles between radiation and spaceflight plants. By data clustering, principal component analysis and correlation analysis, the results showed that protein expression profile of the LET-62.2keV/μm 12C radiation was most similar to spaceflight effect. The dose effect analysis of spaceflight by simulated radiations was also evaluated: radiations of LET-62.2KeV/μm 12C particles (most similar with spaceflight effect) with different cumulative doses (2mGy, 20mGy, 200mGy, 2000mGy) were further carried out. Comparisons of protein expression profiles between irradiated and spaceflight plants indicated that the 2mGy dose-value radiation shared most similar effect with the spaceflight, which confirmed the low-dose effect of space environment. A new space radiation environment effect incentive evaluation system was established based on the protein expression profile analysis.
Comparative quantitative proteomic analysis between a high-tillering dwarf mutant R955 induced by spaceflight and its ground control was performed at three plant vegetative growth stages. It was found that biological processes including energy pathway, photosynthesis, protein metabolism, nitrogen assimilation, amino acid metabolism and response to stimulus were mainly involved in tillering mutation development. K-means clustering revealed that proteins regulated at different stages tended to be involved in different biological processes. Two-way analysis of variance (Two-way ANOVA) screened out that S-like RNases presented direct correlation with the high-tillering ability. The work might provide new insights for further understanding mechanisms of space induced tillering mutation in rice.
利用二维荧光差异显示双向电泳(2-D DIGE)技术获取了2次空间飞行当代、第2代和地基模拟辐射后多个水稻品系以及突变体材料分蘖期的99张双向电泳蛋白表达图谱,建立了包含所有品系共同表达的蛋白点以及品系特异表达蛋白点合集的群体,形成了标准图谱;以该标准图谱为统一范围使不同状态下的大量蛋白表达数据存在定性和定量上的可比性。串联质谱鉴定出204个差异表达蛋白,参与的生物功能涵盖代谢、调节、刺激应答、细胞活动和生殖。
为分析空间飞行当代水稻蛋白变化特征及在子代的遗传特性,利用聚类、主成分分析等方法归纳并比较了不同空间飞行当代及第2代材料与地面对照之间的蛋白表达差异。发现卫星搭载与航天飞船的空间飞行能够使当代水稻叶片蛋白表达特征发生改变,但在不同类型的空间飞行中存在变化程度的区别:飞行时间越长,受空间辐射环境影响越大的材料,蛋白表达谱的差异就越大。从差异蛋白分子生物功能分布上分析了空间环境对水稻细胞生理生化功能的影响,空间飞行当代引起易变蛋白的功能主要包括核酸代谢、光反应、蛋白降解、抗真菌、抗胁迫和氨基酸代谢。空间飞行环境对水稻当代蛋白表达特征的改变在相应的第2代植株中可以发生延续和回复,回复与否以及回复的程度与水稻的品系敏感性相关。细胞壁形成、核酸代谢、蛋白降解、逆转录、氨基酸代谢、磷酸戊糖途径和抗胁迫功能的应答易在第2代回复;参与种子成熟、蛋白折叠、糖酵解、脂类合成、糖原合成和三羧酸循环的蛋白分子仅在第2代时产生应答。空间环境的生物学效应能够体现在蛋白表达层面上。
发现了一些对空间环境敏感的蛋白分子,包括核苷二磷酸激酶1、Rieske铁-硫蛋白前体、半胱氨酸蛋白酶、异黄酮还原酶相似蛋白、S-类核糖核酸酶以及参与蛋白折叠的分子伴侣,它们能够作为水稻对空间环境应答的潜在蛋白分子标志物。
空间辐射是一个非常复杂的复合高能粒子环境。为寻找空间环境的离子诱因,在地面模拟了空间2mGy剂量下不同能量的离子辐射,采用传能线密度(LET)分别为13.3keV/μm的12C离子、30keV/μm的12C离子、31keV/μm的20Ne离子、62.2keV/μm的12C离子和500keV/μm的56Fe离子对水稻种子进行处理,并与返回式卫星搭载的水稻蛋白表达模式进行比较。通过聚类、主成分分析和相关性分析后发现:LET-62.2keV/μm的12C离子辐射蛋白表达特征与空间环境最为类似。为进一步分析空间辐射环境的剂量效应,利用与空间效应相近的LET-62.2keV/μm的12C离子进行了不同累积剂量的辐射(剂量分别为2mGy、20mGy、200mGy、2000mGy),对辐射蛋白表达谱与空间材料进行比较后发现2mGy剂量的辐射效应与空间环境最为接近,证实了空间辐射的低剂量效应。通过基于蛋白表达谱的分析建立了新的空间环境辐射诱因效应评价体系。
对空间诱变多蘖矮杆突变体R955及对照植株3个营养生长期叶片总蛋白的分离及定量分析发现,能量代谢、光合作用、蛋白代谢、氮元素同化、氨基酸代谢以及胁迫应答等过程均参与了空间环境诱导的突变分蘖发育。对差异蛋白在不同时期相对表达量的K-均值聚类分析显示不同功能蛋白表现不同表达模式;二元方差分析筛选出S-类核糖核酸酶可能与水稻分蘖突变性状直接关联。相关结果为空间环境诱发水稻产生分蘖突变的机制提供了新的见解。
In order to study the mechanisms of biological effects of space environment, in the present dissertation, the model plant rice was chosen as experimental material. After several seed spaceflights, proteomic expression profiles of different rice variety plants in 1st and 2nd generations were analysed. Space radiation inducement investigation through protein expression profile analysis based on on-ground simulated radiation experiments was also performed. In addition, mechanisms of tillering mutation induced by spaceflight were also explored by differential protein expression analysis of the tillering dwarf mutant.
Using two-dimensional fluorescent differential electrophoresis (2-D DIGE) technology, 99 protein expression 2-D patterns were acquired from tillering rice plants of different varieties after two kinds of spaceflights in 1st and 2nd generations, after on-ground simulated radiation treatments and from the mutant variety. Standard gel image pattern including groups of proteins that co-expressed in all rice varieties and particularly expressed in every variety was established. The pattern was used as a normalized standard for comparing the large number of protein expression data in different status qualitively and quantitively. Tandem mass spectrometry identified 204 differentially expressed proteins that involved in biological processes including metabolic process, regulation of biological process, response to stimulus, cellular process and reproduction.
For investigation of protein expression profiles of rice plants after spaceflights and the hereditary capacity in the offspring plants, clustering and principal component analysis were performed to summarize and compare the protein expression variations between the space samples and their corresponding ground controls in different spaceflights and in 1st and 2nd generations. Results indicated that protein expression profiles were changed after seed space environment exposures onboard the satellite or space craft, but there were variations in the degree of difference in different types of flights: the longer the flight duration lasted, which meant that rice seeds were affected by space radiation environment heavier, the greater differences existed in protein expression profiles. Distribution of biological processes of differentially expressed proteins indicated that physiological and biochemical changes of rice cells were induced by space environment. The biological processes impacted in 1st generation plants after flights were nucleic acid metabolic process,light reaction,proteolysis,defense response to fungus,response to stress with amino acid and derivative metabolism. Alterations of protein expression profiles in 1st generation would resume or recover in the corresponding 2nd generation plants, the degree of recovery was associated with the sensitivity of the rice variety to environment. Changes of proteins involving in biological processes such as cell wall organization, nucleic acid metabolic process, proteolysis, RNA-dependent DNA replication, amino acid and derivative metabolism, pentose-phosphate shunt, response to stress tended to recover in 2nd generation plants; while seed maturation, protein folding, glycolysis, lipid biosynthetic process, glycogen biosynthetic process and tricarboxylic acid cycle related proteins only differentially expressed in 2nd generation plants. Therefore,the biological effects of space environment could be reflected in protein expression level.
Some of the space-environment-sensitive proteins were found, including nucleoside diphosphate kinase 1, Rieske (Fe-S) protein precursor, cysteine protease, isoflavone reductase-like protein, S-like RNase and molecular chaperones participating in protein folding. They could be used as potential biomarkers that would answer to the stimulus from space environment.
Spaceflight represents a very complex environmental condition with highly ionizing radiations (HZE). To further investigate the incentives of ion effects in space environment, we performed on-ground simulated HZE particle radiations to rice seeds with the same dose (2mGy) as spaceflight but different liner energy transfer (LET) values (13.3keV/μm-12C, 30keV/μm-12C, 31keV/μm-20Ne, 62.2keV/μm-12C, 500keV/μm-56Fe) then compared the protein expression profiles between radiation and spaceflight plants. By data clustering, principal component analysis and correlation analysis, the results showed that protein expression profile of the LET-62.2keV/μm 12C radiation was most similar to spaceflight effect. The dose effect analysis of spaceflight by simulated radiations was also evaluated: radiations of LET-62.2KeV/μm 12C particles (most similar with spaceflight effect) with different cumulative doses (2mGy, 20mGy, 200mGy, 2000mGy) were further carried out. Comparisons of protein expression profiles between irradiated and spaceflight plants indicated that the 2mGy dose-value radiation shared most similar effect with the spaceflight, which confirmed the low-dose effect of space environment. A new space radiation environment effect incentive evaluation system was established based on the protein expression profile analysis.
Comparative quantitative proteomic analysis between a high-tillering dwarf mutant R955 induced by spaceflight and its ground control was performed at three plant vegetative growth stages. It was found that biological processes including energy pathway, photosynthesis, protein metabolism, nitrogen assimilation, amino acid metabolism and response to stimulus were mainly involved in tillering mutation development. K-means clustering revealed that proteins regulated at different stages tended to be involved in different biological processes. Two-way analysis of variance (Two-way ANOVA) screened out that S-like RNases presented direct correlation with the high-tillering ability. The work might provide new insights for further understanding mechanisms of space induced tillering mutation in rice.
引文
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