不同氮肥比例对水稻强弱势籽粒灌浆影响的差异蛋白组学分析
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摘要
水稻是世界上最重要的粮食作物之一,提供给世界人口近60%的粮食需要,水稻籽粒灌浆特性对其最终产量与品质的形成至关重要,因此,水稻籽粒灌浆问题便理所当然地成为当今世界农业科学研究的重要课题。
本文以大穗型水稻“金恢”品系为实验材料,设3种不同的氮肥比例,基蘖蘖肥和穗粒肥比例分别为7∶3,6∶4,5∶5。选取不同的灌浆时期,运用差异蛋白组学从动态上研究这三种氮肥比例对强弱势籽粒灌浆的影响,旨在揭示水稻的强弱势籽粒灌浆差异的分子机理及其与源库流的关系,为有效地挖掘水稻高产潜力、促进粮食增产、提供重要的理论依据。
结果认为大穗型水稻存在着明显的异步灌浆的现象,不同氮肥比例主要影响弱势粒的灌浆,最终影响水稻籽粒的结实率和千粒重,并以生育前后期氮肥比例6∶4,产量最高,氮肥比7∶3,产量最低。进一步采用差异蛋白组学研究强弱势粒对不同氮肥比例的分子响应机制,结果发现在不同的氮肥比例下弱势粒中蛋白出现差异表达的较多,共有26个,强势粒较少,共有9个。经MALDI-TOF/MS分析和数据库检索,弱势粒的有17个差异蛋白质点得到鉴定,强势粒有4个得到鉴定。对这些蛋白质功能的分析发现,后期氮肥的增加对弱势粒灌浆过程中的一些重要的生理活动产生影响,加强了蔗糖的转运能力,促进了淀粉的合成,增加了弱势粒中用于物质代谢的能量,提高了内源激素的含量等。研究还发现后期氮肥的增加对强势粒蛋白质的合成及抗逆能力的加强也有利。此外,有7个在弱势粒中发生差异表达,对籽粒灌浆起重要作用的关键蛋白,发现在强势粒灌浆过程中虽有表达,但表达丰度没有差异,解释了强势粒灌浆的环境稳定性的分子机理。
本研究同时考察了三种不同氮肥比例下大穗型水稻籽粒灌浆期叶片的叶绿素SPAD值的变化,结果可知在总施氮量不变的情况下,适当的加大后期的氮肥比例能够延长叶片叶绿素含量缓降期。进一步采用差异蛋白组学研究了氮肥比例7:3与6:4处理下的叶片蛋白的差异表达情况。结果在这2个氮肥比例处理下的叶片中分别鉴定到24个出现差异表达的蛋白,共有12个蛋白得到鉴定。按照其功能可以分为6类:第一类,参与光合反应:叶绿素a/b结合蛋白,PHY3蛋白,1,5-二磷酸核酮糖羧化酶/加氧酶,核酮糖1,5-二磷酸羧化酶大亚基,第二类:参与抗逆反应:超氧化物歧化酶,依赖谷胱甘肽的脱氢抗坏血酸还原酶I,WRKY转录因子,黄烷酮3-羟化酶,第三类:参与能量与物质代谢:烟酰胺核苷酸腺苷酰转移酶,转氨酶y4uB,第四类:参与物质运输及信号转导:钙调蛋白,v-SNARE家族膜泡运输蛋白。对这些蛋白的功能分析可以发现,籽粒灌浆期叶片受不同氮肥处理的影响主要体现在,加大后期的氮肥施用比例能够延长光合作用的时间,延缓叶片的衰老,增强叶片灌浆后期的能量代谢,促进物质的运输和信号的转导。
在籽粒灌浆的不同时期,叶鞘在不同氮肥比例下共鉴定到20个差异蛋白质点,其中9个得到鉴定。按照其功能可以分为5类:第一类,参与光合作用:核酮糖1,5-二磷酸羧化酶大亚基。第二类,参与物质的运输与合成:DREPP植物质膜多肽家族蛋白,ATP-柠檬酸裂解酶a亚基,推测的内质网膜氧化还原酶5'端。第三类:参与抗逆反应:过氧化物酶,ABC型磷酸盐转运蛋白,肉桂醇脱氢酶,:细胞色素P450家族蛋白。第四类,参与信号转导:磷脂酶D。第五类,参与能量代谢:ATP-柠檬酸裂解酶a亚基。对这些蛋白的功能分析可以发现,在生育后期加大氮肥的比例有利于叶鞘在灌浆的后期光合作用能力的保持,促进物质的转运与信号的转导,加强能量代谢,提高抗逆能力。
本研究表明,水稻强弱势籽粒在灌浆过程中,存在着复杂的蛋白质组调控网络,这为我们进一步揭示水稻灌浆的分子遗传机理奠定了基础。
Rice (Oryza sativa L.), is one of the most important staple food crops for more than 60% of the human population in the world. The grain- filling characteristics of rice plays a crucial role in the yield and quality, so it is the key to study physiological and biochemical characteristics and genetic mechanism during grain filling for high yielding in rice.
A large-spike rice cultivar Jinhui (indica) was used in this study. The experiment was set in 3 different proportions of nitrogen fertilizer (ratio of tiller fertilizer to panicle-grain fertilizer.7:3,6:4,5:5).The investigation was conducted to address the influence of different proportions of nitrogen fertilizer on superior and inferior grain filling of large-panicle type rice cultivars by the method of differential proteomics. The aim is to reveal the metabolic mechanism of source-sink-stream in large panicles rice during grain filling period and provide important theoretical references to dig the rice high yield potential, and to accelerate grain production increase.
The results showed that there exists obvious asynchronous filling phenomenon in large -panicle type rice, and the change in the N application proportion mainly influenced the inferior grain filling, in turn altered seed setting rate and 1000-grain weight of yield components, indicating the highest grain yield at 6:4 ratio of N supply and the lowest grain yield at 7:3 ratio. Differential proteomic analysis showed much more differentially expressed proteins (26 proteins vs 9 ones in total) detected in the inferior grain than in the superior grains during grain-filling of rice under different N application proportions and 17 differential protein spots from the inferior grains and 4 ones from superior grains have been identified and assigned in functions by using MALDI-TOF/MS analysis and database searching. The result suggested that increasing N application proportion in post growth stage of rice favored in the upregulation of the genes involved in physiological activities of inferior grain filling, including the proteins associated with transport capacity of sucrose, starch biosynthesis , energy accumulation for the material metabolism, and hormone production etc in inferior grains of rice. It was also found that the increases of N application proportion in post growth stage were good for the biosynthesis of proteins and the improvement of stress-resistant ability in superior grains. The 7 key proteins to grain-filling, differentially expressed in the inferior grains mediated by different N application proportions were also detected, but no differences in expression abundance of them were found in the superior grains, which explained the molecular mechanism of environmental stability for the superior grain-filling.
SPAD (Soil and Plant Analyzer Development) values of leaf of large-panicle type rice at 3 different proportions of nitrogen fertilizer applied in pre and post growth stages were investigated. The results showed that the increases of N application proportion in post growth stage were good for delaying the degradation of chlorophyll content. The method of differential proteomics were used to identify the function of differential expression proteins in leaves under 2 different proportions of nitrogen fertilizer(ratio 7:3,6:4),24 proteins were detected, and 12 proteins were identified to be the functional proteins. These proteins were classified into 4 categories according to their function. The fist group, which included chlorophyll a-b binding protein, PHY3 protein, RuBisCO activase small isoform precursor, rubisco large subunit, participated in the photosynthetic reaction. The second group was involved in the defense reactions, such as GSH-dependent dehydroascorbate reductase 1, superoxide dismutase [Mn], WRKY DNA binding domain containing protein, flavanone-3-hydroxylase.The third group was involved in the metabolism of material and energy, which include nicotinate-nucleotide adenylyltransferase family protein, aminotransferase y4uB. The fourth group participated in the material transport and signal transduction. This group included Calmodulin and Vesicle transport v-SNARE family protein. The result suggested that the effect of the increases of N application proportion in post growth stage on the leaves during the grain filling period were as follows: elongate photosynthetic time,delay senescence of leaves, strengthen the energy metabolism of leaves in post growth stage of rice, accelerate matter transportation and signal transduction.
Twenty proteins with obviously different abundances were detected in leaf sheath during grain filling period under 2 different proportions of nitrogen fertilizer(ratio 7:3,6:4), and nine of them were identified in function. These proteins were classified into 5 categories according to their function. The first group involved in the photosynthetic reaction, which include ribulose bisphosphate carboxylase large subunit. The second group, which include DREPP plasma membrane polypeptide family protein, putative ATP citrate lyase a-subunit, putative endoplasmic reticulum oxidoreductin , participated in the transportation and synthesis of material. The third group involved in the signal transduction, which include Phospholipase D. The fourth group participated in the defense reactions. This group included Peroxidase , ABC-type phosphate transport system , Putative cinnamyl-alcohol dehydrogenase, Cytochrome P450 family protein. The fifth group is the Putative ATP citrate lyase a-subunit, which participated in the energy metabolism.These results suggested that increasing N application proportion in post growth stage of rice favored in physiological activities of leaf sheaf, including the photosynthesis, the transportation of material and signal, energy metabolism and resistance.
The present work suggested that there existed a regulative network of complex proteome during grain filling periods and it paved the way for us to further study the molecular genetic mechanism of rice.
本文以大穗型水稻“金恢”品系为实验材料,设3种不同的氮肥比例,基蘖蘖肥和穗粒肥比例分别为7∶3,6∶4,5∶5。选取不同的灌浆时期,运用差异蛋白组学从动态上研究这三种氮肥比例对强弱势籽粒灌浆的影响,旨在揭示水稻的强弱势籽粒灌浆差异的分子机理及其与源库流的关系,为有效地挖掘水稻高产潜力、促进粮食增产、提供重要的理论依据。
结果认为大穗型水稻存在着明显的异步灌浆的现象,不同氮肥比例主要影响弱势粒的灌浆,最终影响水稻籽粒的结实率和千粒重,并以生育前后期氮肥比例6∶4,产量最高,氮肥比7∶3,产量最低。进一步采用差异蛋白组学研究强弱势粒对不同氮肥比例的分子响应机制,结果发现在不同的氮肥比例下弱势粒中蛋白出现差异表达的较多,共有26个,强势粒较少,共有9个。经MALDI-TOF/MS分析和数据库检索,弱势粒的有17个差异蛋白质点得到鉴定,强势粒有4个得到鉴定。对这些蛋白质功能的分析发现,后期氮肥的增加对弱势粒灌浆过程中的一些重要的生理活动产生影响,加强了蔗糖的转运能力,促进了淀粉的合成,增加了弱势粒中用于物质代谢的能量,提高了内源激素的含量等。研究还发现后期氮肥的增加对强势粒蛋白质的合成及抗逆能力的加强也有利。此外,有7个在弱势粒中发生差异表达,对籽粒灌浆起重要作用的关键蛋白,发现在强势粒灌浆过程中虽有表达,但表达丰度没有差异,解释了强势粒灌浆的环境稳定性的分子机理。
本研究同时考察了三种不同氮肥比例下大穗型水稻籽粒灌浆期叶片的叶绿素SPAD值的变化,结果可知在总施氮量不变的情况下,适当的加大后期的氮肥比例能够延长叶片叶绿素含量缓降期。进一步采用差异蛋白组学研究了氮肥比例7:3与6:4处理下的叶片蛋白的差异表达情况。结果在这2个氮肥比例处理下的叶片中分别鉴定到24个出现差异表达的蛋白,共有12个蛋白得到鉴定。按照其功能可以分为6类:第一类,参与光合反应:叶绿素a/b结合蛋白,PHY3蛋白,1,5-二磷酸核酮糖羧化酶/加氧酶,核酮糖1,5-二磷酸羧化酶大亚基,第二类:参与抗逆反应:超氧化物歧化酶,依赖谷胱甘肽的脱氢抗坏血酸还原酶I,WRKY转录因子,黄烷酮3-羟化酶,第三类:参与能量与物质代谢:烟酰胺核苷酸腺苷酰转移酶,转氨酶y4uB,第四类:参与物质运输及信号转导:钙调蛋白,v-SNARE家族膜泡运输蛋白。对这些蛋白的功能分析可以发现,籽粒灌浆期叶片受不同氮肥处理的影响主要体现在,加大后期的氮肥施用比例能够延长光合作用的时间,延缓叶片的衰老,增强叶片灌浆后期的能量代谢,促进物质的运输和信号的转导。
在籽粒灌浆的不同时期,叶鞘在不同氮肥比例下共鉴定到20个差异蛋白质点,其中9个得到鉴定。按照其功能可以分为5类:第一类,参与光合作用:核酮糖1,5-二磷酸羧化酶大亚基。第二类,参与物质的运输与合成:DREPP植物质膜多肽家族蛋白,ATP-柠檬酸裂解酶a亚基,推测的内质网膜氧化还原酶5'端。第三类:参与抗逆反应:过氧化物酶,ABC型磷酸盐转运蛋白,肉桂醇脱氢酶,:细胞色素P450家族蛋白。第四类,参与信号转导:磷脂酶D。第五类,参与能量代谢:ATP-柠檬酸裂解酶a亚基。对这些蛋白的功能分析可以发现,在生育后期加大氮肥的比例有利于叶鞘在灌浆的后期光合作用能力的保持,促进物质的转运与信号的转导,加强能量代谢,提高抗逆能力。
本研究表明,水稻强弱势籽粒在灌浆过程中,存在着复杂的蛋白质组调控网络,这为我们进一步揭示水稻灌浆的分子遗传机理奠定了基础。
Rice (Oryza sativa L.), is one of the most important staple food crops for more than 60% of the human population in the world. The grain- filling characteristics of rice plays a crucial role in the yield and quality, so it is the key to study physiological and biochemical characteristics and genetic mechanism during grain filling for high yielding in rice.
A large-spike rice cultivar Jinhui (indica) was used in this study. The experiment was set in 3 different proportions of nitrogen fertilizer (ratio of tiller fertilizer to panicle-grain fertilizer.7:3,6:4,5:5).The investigation was conducted to address the influence of different proportions of nitrogen fertilizer on superior and inferior grain filling of large-panicle type rice cultivars by the method of differential proteomics. The aim is to reveal the metabolic mechanism of source-sink-stream in large panicles rice during grain filling period and provide important theoretical references to dig the rice high yield potential, and to accelerate grain production increase.
The results showed that there exists obvious asynchronous filling phenomenon in large -panicle type rice, and the change in the N application proportion mainly influenced the inferior grain filling, in turn altered seed setting rate and 1000-grain weight of yield components, indicating the highest grain yield at 6:4 ratio of N supply and the lowest grain yield at 7:3 ratio. Differential proteomic analysis showed much more differentially expressed proteins (26 proteins vs 9 ones in total) detected in the inferior grain than in the superior grains during grain-filling of rice under different N application proportions and 17 differential protein spots from the inferior grains and 4 ones from superior grains have been identified and assigned in functions by using MALDI-TOF/MS analysis and database searching. The result suggested that increasing N application proportion in post growth stage of rice favored in the upregulation of the genes involved in physiological activities of inferior grain filling, including the proteins associated with transport capacity of sucrose, starch biosynthesis , energy accumulation for the material metabolism, and hormone production etc in inferior grains of rice. It was also found that the increases of N application proportion in post growth stage were good for the biosynthesis of proteins and the improvement of stress-resistant ability in superior grains. The 7 key proteins to grain-filling, differentially expressed in the inferior grains mediated by different N application proportions were also detected, but no differences in expression abundance of them were found in the superior grains, which explained the molecular mechanism of environmental stability for the superior grain-filling.
SPAD (Soil and Plant Analyzer Development) values of leaf of large-panicle type rice at 3 different proportions of nitrogen fertilizer applied in pre and post growth stages were investigated. The results showed that the increases of N application proportion in post growth stage were good for delaying the degradation of chlorophyll content. The method of differential proteomics were used to identify the function of differential expression proteins in leaves under 2 different proportions of nitrogen fertilizer(ratio 7:3,6:4),24 proteins were detected, and 12 proteins were identified to be the functional proteins. These proteins were classified into 4 categories according to their function. The fist group, which included chlorophyll a-b binding protein, PHY3 protein, RuBisCO activase small isoform precursor, rubisco large subunit, participated in the photosynthetic reaction. The second group was involved in the defense reactions, such as GSH-dependent dehydroascorbate reductase 1, superoxide dismutase [Mn], WRKY DNA binding domain containing protein, flavanone-3-hydroxylase.The third group was involved in the metabolism of material and energy, which include nicotinate-nucleotide adenylyltransferase family protein, aminotransferase y4uB. The fourth group participated in the material transport and signal transduction. This group included Calmodulin and Vesicle transport v-SNARE family protein. The result suggested that the effect of the increases of N application proportion in post growth stage on the leaves during the grain filling period were as follows: elongate photosynthetic time,delay senescence of leaves, strengthen the energy metabolism of leaves in post growth stage of rice, accelerate matter transportation and signal transduction.
Twenty proteins with obviously different abundances were detected in leaf sheath during grain filling period under 2 different proportions of nitrogen fertilizer(ratio 7:3,6:4), and nine of them were identified in function. These proteins were classified into 5 categories according to their function. The first group involved in the photosynthetic reaction, which include ribulose bisphosphate carboxylase large subunit. The second group, which include DREPP plasma membrane polypeptide family protein, putative ATP citrate lyase a-subunit, putative endoplasmic reticulum oxidoreductin , participated in the transportation and synthesis of material. The third group involved in the signal transduction, which include Phospholipase D. The fourth group participated in the defense reactions. This group included Peroxidase , ABC-type phosphate transport system , Putative cinnamyl-alcohol dehydrogenase, Cytochrome P450 family protein. The fifth group is the Putative ATP citrate lyase a-subunit, which participated in the energy metabolism.These results suggested that increasing N application proportion in post growth stage of rice favored in physiological activities of leaf sheaf, including the photosynthesis, the transportation of material and signal, energy metabolism and resistance.
The present work suggested that there existed a regulative network of complex proteome during grain filling periods and it paved the way for us to further study the molecular genetic mechanism of rice.
引文
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