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作物学报 ›› 2018, Vol. 44 ›› Issue (6): 897-908.doi: 10.3724/SP.J.1006.2018.00897

• 耕作栽培·生理生化 • 上一篇    下一篇

表油菜素内酯影响水稻幼苗响应低温胁迫的蛋白质组学分析

王道平1,2,徐江2,牟永莹2,3,闫文秀2,3,赵梦洁3,马博3,李群1,*(),张丽娜2,3,潘映红2,3,*()   

  1. 1 新疆大学生命科学与技术学院, 新疆乌鲁木齐 830046
    2 中国农业科学院作物科学研究所, 北京 100081
    3 农作物基因资源与基因改良国家重大科学工程, 北京 100081
  • 收稿日期:2017-08-29 接受日期:2018-03-18 出版日期:2018-06-12 网络出版日期:2018-03-19
  • 通讯作者: 李群,潘映红
  • 基金资助:
    本研究由国家自然科学基金项目(31571589);国家重点基础研究发展计划(973计划)项目子课题(2015CB150401);中国农业科学院创新工程项目资助(作物分子标记技术及其应用创新团队)

Proteomic Analysis of the Effect of 2,4-Epibrassinolide on Rice Seedlings Response to Cold Stress

Dao-Ping WANG1,2,Jiang XU2,Yong-Ying MU2,3,Wen-Xiu YAN2,3,Meng-Jie ZHAO3,Bo MA3,Qun LI1,*(),Li-Na ZHANG2,3,Ying-Hong PAN2,3,*()   

  1. 1 College of Life Science and Technology, Xinjiang University, Urumqi 830046, Xinjiang, China
    2 Institute of Crop Sciences, Chinese Academy of Agricultural Sciences, Beijing 100081, China
    3 National Key Facility for Crop Gene Resources and Genetic Improvement, Chinese Academy of Agricultural Sciences, Beijing 100081, China
  • Received:2017-08-29 Accepted:2018-03-18 Published:2018-06-12 Published online:2018-03-19
  • Contact: Qun LI,Ying-Hong PAN
  • Supported by:
    This study was supported by the National Natural Science Foundation of China(31571589);the National Basic Research Program of China (973 Program)(2015CB150401);the Agricultural Science and Technology Innovation Program(作物分子标记技术及其应用创新团队)

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摘要：

表油菜素内酯(2,4-Epibrassinolide, EBR)是一种被广泛研究和应用的油菜素内酯类(brassinosteroids, BRs)植物生长调节剂, 它能有效增强植物对低温的耐受性, 但EBR在蛋白质组水平上对水稻幼苗响应低温胁迫的影响尚不清楚。本研究用0.1 mg L -1 EBR和蒸馏水分别浸泡萌发的日本晴种子, 然后提取4°C胁迫培养和26°C正常培养幼苗的总蛋白, 进行质谱非标(label-free)定量分析和平行反应监测(parallel reaction monitoring, PRM)验证。最终共鉴定出5778个蛋白质, 其中, 在有定量信息的4834个蛋白中, 401个上调和220个下调蛋白与EBR影响水稻幼苗响应低温胁迫有关。功能分析和代谢通路富集分析发现, 上调蛋白主要与RNA结合或水解酶活性等分子功能相关, 并富集在碳代谢、叶酸合成和氨基酸生物合成等途径中; 下调蛋白主要与催化活性和氧化还原酶活性相关, 主要涉及卟啉和叶绿素代谢等代谢途径。PRM验证结果和文献证据显示, 分布在碳代谢和苯丙素代谢通路中的NADP-苹果酸酶、过氧化物酶、3-磷酸甘油酸脱氢酶、烯醇化酶、甘油醛-3-磷酸脱氢酶和丙酮酸激酶参与了EBR对低温胁迫水稻幼苗的调控, 提示BRs可通过多种途径影响水稻幼苗对低温胁迫的响应。

关键词: 表油菜素内酯, 低温胁迫, 蛋白质组学, 水稻

Abstract:

As a plant growth regulator which functionally resembles a kind of plant hormone Brassinosteroids (BRs), 2,4-Epibrassinolide (EBR) has been widely studied and applied in different aspects. EBR can enhance plant’s cold tolerance effectively, but the proteomic characteristics of the effect of EBR on rice seedings response to cold stress are still unclear. In this study, the germinating seeds of rice Nipponbare were treated with 0.1 mg L -1 EBR and distilled water before they were cultivated at 4°C or 26°C, and then the total protein of each group of seedlings was extracted. Finally, proteomes of rice seedings were analyzed by label-free quantitative mass spectrometry, and some important proteins were verified by parallel reaction monitoring technique (PRM). A total of 5778 protein groups were identified by qualitative method and 4834 protein groups were accurately quantitated. Among them, 401 up-regulated and 220 down-regulated proteins were related to the effect of EBR on rice seedings response to cold stress. The up-regulated proteins were mainly related to molecular function of RNA binding and hydrolase activity, and mainly enriched in the pathways of carbon metabolism, folic acid synthesis and amino acid biosynthesis. The down-regulated proteins were mainly related to catalytic activity and oxidoreductase activity, and mainly enriched in the pathways of porphyrin and chlorophyll metabolism and other metabolic pathways. PRM validation and literature analysis showed that NADP-malic acidase, peroxidase, 3-phosphoglycerate dehydrogenase, enolase, glyceraldehyde-3-phosphate dehydrogenase and pyruvate kinase, which are distributed in the pathways of carbon metabolism and phenylpropanol metabolism and others, take part in the regulation of EBR on rice seedlings response to cold stress, suggesting that BRs can affect rice seedlings response to cold stress through a variety of pathways.

Key words: 2, 4-Epibrassinolide, cold stress, proteomics, rice

图1

水稻低温胁迫表型及叶绿素含量 A: 4组样品表型图; B: 4组样品叶绿素a和叶绿素b含量比较; C: 4组样品全株重比较。随机选取10株幼苗称重, 重复3次, 算其平均值作为该组样品的全株鲜重。4B: EBR处理4°C培养样品, 4: 单独4°C培养样品, 26B: EBR处理26°C培养样品, 26: 单独26°C培养样品。B和C中标以不同小字母的同组数值在0.05水平差异显著。"

图2

质谱定性和定量分析结果 A: 定性鉴定蛋白数及样品间共有蛋白数; B: 样品间的皮尔逊相关性分析图; C: 定量鉴定蛋白质信号强度分布表。"

图3

定量蛋白的差异性比较 A: 样品间的差异蛋白数; B: 与EBR影响水稻幼苗响应低温胁迫有关上调蛋白维恩图; C: 与EBR影响水稻幼苗响应低温胁迫有关下调蛋白维恩图。H表示上调, L表示下调。"

图4

影响水稻幼苗响应低温胁迫相关蛋白的本体分析 A: 401个上调蛋白质的本体分析; B: 220个下调蛋白质的本体分析。"

图5

代谢通路分析 A: 401个上调差异蛋白质代谢通路分析; B: 220个下调差异蛋白质的代谢通路分析。"

表1

与EBR影响水稻幼苗响应低温胁迫相关的重要通路及富集的蛋白质"

通路和蛋白
Pathway name and protein IDs		 注解
Annotation		 信号强度Intensity
26 26B 4 4B
碳代谢 Carbon metabolism
LOC_Os01g54030.1 Nadp-dependent malic enzyme 0.00E+00 0.00E+00 0.00E+00 1.78E+07
LOC_Os02g38200.1 Dehydrogenase 3.97E+08 1.78E+08 9.81E+07 2.64E+08
LOC_Os03g15050.2 Phosphoenolpyruvate carboxykinase 1.22E+08 5.70E+07 4.21E+07 1.26E+08
LOC_Os04g24140.1 Ribose-5-phosphate isomerase a 1.10E+08 8.97E+06 1.79E+07 7.07E+07
LOC_Os06g04510.1 Enolase 5.68E+07 1.08E+07 0.00E+00 2.00E+07
LOC_Os06g05700.1 Cysteine synthase 0.00E+00 0.00E+00 0.00E+00 8.73E+06
LOC_Os06g45590.1 Glyceraldehyde-3-phosphate dehydrogenase 0.00E+00 0.00E+00 0.00E+00 3.28E+07
LOC_Os07g09890.1 Hexokinase 2.82E+07 0.00E+00 0.00E+00 2.09E+07
LOC_Os08g02700.1 Fructose-bisphospate aldolase isozyme 3.73E+07 0.00E+00 0.00E+00 8.83E+06
LOC_Os09g24910.2 Phosphofructokinase 0.00E+00 0.00E+00 0.00E+00 2.38E+07
LOC_Os11g10980.1 Pyruvate kinase 0.00E+00 0.00E+00 0.00E+00 1.06E+07
LOC_Os11g41160.3 Phosphoserine phosphatase 2.13E+07 1.00E+07 0.00E+00 2.83E+07
LOC_Os12g05110.1 Pyruvate kinase 2.37E+08 1.29E+08 5.51E+07 2.95E+08
LOC_Os06g35540.1 Aminotransferase 5.93E+07 1.63E+08 1.61E+08 7.89E+07
LOC_Os06g44460.1 D-3-phosphoglycerate dehydrogenase 9.76E+06 7.49E+07 2.85E+07 3.37E+07
LOC_Os05g49760.1 Dehydrogenase 0.00E+00 1.20E+08 1.00E+08 3.93E+07
苯丙素生物合成 Phenylpropanoid biosynthesis
LOC_Os10g17650.1 Os10bglu34—beta-glucosidase homologue 1.02E+08 0.00E+00 7.90E+07 4.76E+08
LOC_Os01g32364.1 Os1bglu1—beta-mannosidase/glucosidase homologue 0.00E+00 0.00E+00 0.00E+00 8.51E+06
LOC_Os01g73200.1 Peroxidase 0.00E+00 0.00E+00 0.00E+00 2.12E+07
LOC_Os02g41680.1 Phenylalanine ammonia-lyase 7.94E+06 0.00E+00 0.00E+00 6.33E+06
LOC_Os04g56180.1 Peroxidase 1.25E+08 2.86E+07 5.14E+07 7.05E+07
LOC_Os03g11420.1
 Os3bglu6—beta-glucosidase/beta-fucosidase/beta-galactosidase 0.00E+00
 2.25E+07
 3.03E+07
 6.87E+06
LOC_Os01g22249.1 Peroxidase 1.24E+07 5.91E+07 2.05E+08 0.00E+00
LOC_Os05g04500.1 Peroxidase 8.07E+06 1.92E+08 5.61E+07 0.00E+00
LOC_Os07g01410.1 Peroxidase 2.01E+07 5.70E+06 8.45E+07 0.00E+00
LOC_Os08g34280.1 Cinnamoyl-coa reductase 0.00E+00 6.70E+06 6.25E+07 0.00E+00
LOC_Os09g33680.1 Os9bglu31—beta-glucosidase, dhurrinase 0.00E+00 4.08E+07 1.92E+07 1.48E+07
卟啉和叶绿素代谢 Porphyrin and chlorophyll metabolism
LOC_Os03g22780.1 DVR 8.72E+07 0.00E+00 4.27E+07 9.48E+07
LOC_Os01g16520.1 Glutamyl-tRNA synthetase 0.00E+00 1.59E+07 1.40E+07 0.00E+00
LOC_Os01g57460.1 Frataxin, putative, expressed 4.04E+07 9.91E+07 1.15E+08 0.00E+00
LOC_Os10g37210.1
 FAD dependent oxidoreductase domain containing
Protein		 2.01E+07
 3.51E+06
 5.45E+07
 0.00E+00
LOC_Os04g41260.1 Amine oxidase 0.00E+00 2.87E+06 1.70E+07 0.00E+00
叶酸生物合成 Folate biosynthesis
LOC_Os11g29390.1
 Bifunctional dihydrofolate reductase-thymidylate
synthase		 4.47E+06
 1.31E+07
 3.66E+06
 0.00E+00
LOC_Os09g38759.1 Dihydroneopterin aldolase 2.42E+07 0.00E+00 5.66E+06 2.25E+07
LOC_Os04g38950.1 Class I glutamine amidotransferase 6.99E+07 0.00E+00 2.89E+07 2.37E+07
LOC_Os03g02030.2 Folylpolyglutamate synthase 0.00E+00 0.00E+00 0.00E+00 2.04E+07
LOC_Os02g35200.1 Vp15 0.00E+00 0.00E+00 0.00E+00 7.22E+06
通路和蛋白
Pathway name and protein IDs		 注解
Annotation		 信号强度Intensity
26 26B 4 4B
不饱和脂肪酸生物合成 Biosynthesis of unsaturated fatty acids
LOC_Os01g65830.1 Acyl-desaturase 6.12E+07 0.00E+00 0.00E+00 8.06E+06
LOC_Os02g48560.6 Fatty acid desaturase 4.69E+08 7.57E+07 8.14E+07 3.83E+08
LOC_Os08g10010.1 Acyl-desaturase 1.38E+08 2.72E+07 4.92E+07 6.52E+07
LOC_Os11g39220.2 Acyl-coenzyme A oxidase 0.00E+00 4.07E+07 3.18E+07 0.00E+00
脂肪酸生物合成 Fatty acid biosynthesis
LOC_Os01g65830.1 Acyl-desaturase 6.12E+07 0.00E+00 0.00E+00 8.06E+06
LOC_Os08g10010.1 Acyl-desaturase 1.38E+08 2.72E+07 4.92E+07 6.52E+07
LOC_Os03g28420.1 3-oxoacyl-synthase 8.02E+07 3.53E+07 0.00E+00 1.01E+08
LOC_Os01g48910.2 Long-chain acyl-coa synthetase 3.27E+07 2.61E+07 4.38E+06 5.34E+07
LOC_Os12g04990.3 Long-chain acyl-coa synthetase 1.80E+07 2.67E+07 6.15E+07 9.72E+06

图6

部分差异丰度蛋白PRM验证 A: 3-磷酸甘油酸脱氢酶; B: 烯醇化酶; C: NADP-苹果酸酶; D: 甘油醛-3-磷酸脱氢酶; E: 丙酮酸激酶; F: 过氧化物酶。"

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