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作物学报 ›› 2018, Vol. 44 ›› Issue (11): 1733-1742.doi: 10.3724/SP.J.1006.2018.01733

• 研究简报 • 上一篇    

玉米种胚HSP20基因对人工老化处理的响应

邢芦蔓,吕伟增,雷薇,梁雨欢,卢洋,陈军营()   

  1. 河南农业大学农学院, 河南郑州 450046
  • 收稿日期:2018-03-20 接受日期:2018-07-20 出版日期:2018-11-12 网络出版日期:2018-07-30
  • 通讯作者: 陈军营
  • 基金资助:
    本研究由国家自然科学基金项目(31571761)

Response of HSP20 Genes to Artificial Aging Treatment in Maize Embryo

Lu-Man XING,Wei-Zeng LYU,Wei LEI,Yu-Huan LIANG,Yang LU,Jun-Ying CHEN()   

  1. College of Agronomy, Henan Agricultural University, Zhengzhou 450046, Henan, China
  • Received:2018-03-20 Accepted:2018-07-20 Published:2018-11-12 Published online:2018-07-30
  • Contact: Jun-Ying CHEN
  • Supported by:
    This study was supported by the National Natural Science Foundation of China(31571761)

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摘要：

以玉米杂交种“郑单958”为材料, 采用高温(45°C)高湿(100%相对湿度)对玉米种子进行人工老化处理, 并用转录组技术, 研究植物HSP20基因对种子人工老化处理的响应, 旨在为揭示种子衰老的分子机制提供依据。结果表明, 随着老化时间的延长, 种子活力和种胚内过氧化氢酶的活性均表现下降趋势; 过氧化氢含量在老化第3天达到最大值, 随后下降; 丙二醛含量逐渐升高; 转录组检测表明, 种子老化过程中差异显著的HSP20基因有25个, 这些基因编码的HSP20蛋白主要被定位在细胞核、线粒体、以及叶绿体上, 其序列中均含有ACD保守序列(RVDWRETPDAHEIVVDVP GMRREDLRIEVEDNRVLRVSGERRRAEERKGDH WHREERSYGRFWRRFRLPENADLDSVAASLDSGVL TVRFRK)。该序列中含有较多的Arg (11.2%)、Lys (7.2%)、Pro (4.2%)、Thr (3.9%)等氨基酸, 老化过程中积累的ROS可能氧化这些氨基酸, 导致HSP20结构破坏、功能丧失。利用qRT-PCR技术对挑选的编码细胞质、叶绿体和线粒体HSP20的基因的表达模式分析显示, 随着老化程度的加深, 2个编码细胞质HSP20的基因上调表达, 另外两个编码叶绿体和线粒体HSP20基因的表达量在老化第3天达最大值, 随后下降。推测HSP20基因对种子老化有重要作用, HSP20蛋白的ACD结构域中Arg、Lys等氨基酸的靶向氧化可能是种子衰老的主要原因之一。

关键词: 玉米, 种胚, 种子老化, 差异表达基因, HSP20

Abstract:

Maize (Zea mays L.) cultivar ‘Zhengdan 958’ seeds were treated by artificial aging (45°C, 100% relative humidity) and RNA-seq was used to study the response of HSP20 genes to artificial aging treatment of seeds, so as to provide evidences for uncovering the molecular mechanism of seed aging. In the study, the seed vigor decreased dramatically with increasing aging time. The activity of catalase in seed embryo showed a decreasing trend. The content of hydrogen peroxide increased to the maximum at the third day of aging and then decreased. The content of malonaldehyde increased, which indicated that the damage to the membrane system increased. Twenty-five HSP20 genes were identified. These genes encoded HSP20 and mainly distributed in nuclei, mitochondria and chloroplasts. There was a conserved ACD sequence (RVDWRETPDAHEIVVDVPGMRREDLRIEVE DNRVLRVSGERRRAEERKGDHWHREERSYGRFW RRFRLPENADLDSVAASLDSGVLTVRFRK) in HSP20 protein which contains more amino acids, such as Arg (11.2%), Lys (7.2%), Pro (4.2%), and Thr (3.9%). In the aging process these amino acids might be oxidized by ROS accumulated in the embryo, leading to protein structure damaged and loss of functions. The expression patterns of HSP20 genes in cytoplasm, chloroplasts and mitochondria were validated by qRT-PCR, showing that the expressions of cytoplasmic HSP20 genes were up-regulated and those of chloroplast and mitochondrial HSP20 genes reached peak at the third day of aging treatment, and then declined. These suggested that the HSP20 genes play an important role in seeds aging, and the targeted oxidation of Arg, Lys and other amino acids in the ACD structural domain may be an essential cause leading to seed deterioration.

Key words: maize, seed embryo, seed aging, differential expressed genes, HSP20

附表1

HAP20S基因表达分析的引物"

基因ID号
Gene ID		 引物序列
Primer sequence (5'-3')		 退火温度
Annealing temperature (℃)		 产物长度
Product length (bp)
GRMZM2G049767 F GCTGTACTACAGGTAGTATAAC 59 105
R GCACAACACTTCCATACA 59
GRMZM2G306679 F TAGTGATTATCCGTTATGTACTC 59 141
R ACAGAACAGATTGTATTAGGTT 59
GRMZM2G479260 F GTCTTCGTCTTAGTGTTGAT 58.9 109
R TACTCAAGACAGCCATGA 59
GRMZM2G375517 F AGATGTGAATGTGATCTGAC 59 116
R AGTACATATTAACTCACGCAA 58.9
GRMZM5G858128 F TCTAACAGAACTCGCTGA 59 179
R CATCATCATACCACCGATC 59.1
GRMZM2G346839 F CATCATCATACCACCGATC 59.1 179
R TCTAACAGAACTCGCTGA 59
GRMZM2G335242 F GTACGTCTGAATTCTGGTC 59.1 98
R CTCACTCGGATTACTTGC 58.9
GRMZM2G080724 F GATTCATCAGGACGATGAG 59 101
R AAGAAGAGTGGTAGCTAGAT 58.9
GRMZM2G012455 F AGACCATCGAGATTAAGGT 59 179
R ACACATCAGGTAGGATGAT 59
GRMZM2G311710 F ACACATCAGGTAGGATGAT 59 179
R AGACCATCGAGATTAAGGT 59
GRMZM2G333635 F GAGGACAAGAACGACAAG 59.1 154
R TCTTCACCTCCGTCTTAG 59.1
GRMZM2G158232 F TGAGGTGAAGGCTATTGA 59 89
R ATATCGTAGAAGACACAGGA 59
GRMZM2G331701 F ATGACTCAATGACGGAGA 59.1 90
R GCATATACGCTGTCACAA 58.9
GRMZM5G849535 F ATATCGTAGAAGACACAGGA 59 110
R GATATCTGGTTGAGCATCC 59
GRMZM2G034157 F GGTATCACCAATCATCCTATC 59.2 83
R GTTACAAGAGACATGTAGAGAT 58.9
GRMZM2G083810 F ACCAATTCCTATCTGATGTATC 59 112
R GTCAATGAAGACTACGGATTA 59
GRMZM2G332512 F GCTTGTTGTGTTGGTTTC 58.9 117
R CCTTCCAGAAGAACAGAAG 59
GRMZM2G404274 F ACACATCAGGTAGGATGAT 59 179
R AGACCATCGAGATTAAGGT 59
GRMZM2G085934 F CGATGGACATAGTGGAGA 59.2 113
R CTTCATCACTAGCATCCG 58.7
GRMZM2G149647 F GGTCTTTGTGTAGCACTAG 59.1 109
R ATAGCAGAATTGAAGCGTT 59
GRMZM2G098167 F AGAGTAGCAGCAGTAGAAT 59 154
R CAGAGACACGATTGAATAATTC 59
GRMZM2G046382 F GTCGTAATGTCGTAGGATG 59 85
R ATCCATACATAGATTCAGACAC 59
AC208204.3_FGT006 F CAACGTCCTTCAGATCAG 58.9 175
R CACAGTGACTGTAAGCAC 59.4
NM-001155179.1(Actin1) F GTATGAGCAAGGAGATCAC 58.9 194
R TTAGAAGCACTTCATGTGG 59.0

表1

人工老化处理对种子不同活力指标的影响"

处理时间
Treatment time		 发芽率
Germination rate (%)		 发芽指数
Germination index		 活力指数
Vigor index		 平均发芽日数
Mean days of germination (d)
0 d 100±0.00 a 9.44±0.25 a 124.14±5.57 a 2.17±0.08 b
1 d 100±0.00 a 8.22±0.54 b 100.07±4.93 b 2.53±0.16 b
2 d 100±0.00 a 8.11±0.25 b 101.32±1.17 b 2.56±0.08 b
3 d 86.67±0.06 b 6.29±0.32 c 75.73±10.45 c 2.86±0.03 b
4 d 66.67±0.08 c 3.96±0.50 d 41.63±5.39 d 3.48±0.11 ab
5 d 21.67±0.10 d 1.11±0.42 e 8.95±1.84 e 4.11±1.02 a

图1

人工老化对种子活力的影响"

表2

人工老化处理对种胚中不同生理指标的影响"

处理时间
Treatment time		 过氧化氢酶活性
CAT activity (U g-1 FW min-1)		 过氧化氢含量
H2O2 content (μmol g-1 FW)		 丙二醛含量
MDA content (μmol g-1 FW)
0 d 0.39±0.11 a 0.38±0.03 b 0.16±0.11 c
1 d 0.26±0.13 a 0.40±0.02 b 0.18±0.03 c
2 d 0.18±0.12 ab 0.42±0.02 b 0.33±0.10 c
3 d 0.14±0.07 b 0.51±0.06 a 0.43±0.19 c
4 d 0.14±0.03 b 0.43±0.02 ab 0.87±0.26 b
5 d 0.06±0.02 b 0.29±0.03 c 1.955±0.21 a

表3

人工老化处理条件下HSP20相关基因及其注释"

基因ID
Gene ID		 log2比值
log2 Ratio		 P值
P-value		 错误发现率
False discovery rate		 基因注释
Gene annotation		 亚细胞定位
Subcellular localization
GRMZM2G481605 8.84 1.89E-259 5.63E-257 17.9 kD class I heat shock protein Nuclear
GRMZM2G049767 8.19 1.07E-164 2.04E-162 17.9 kD class I heat shock protein Nuclear
GRMZM2G306679 7.37 0 0 17.9 kD class I heat shock protein Nuclear
GRMZM2G479260 6.71 0 0 17.9 kD class I heat shock protein Nuclear
GRMZM2G375517 6.70 0 0 17.8 kD class I heat shock protein Cytoplasmic
GRMZM5G858128 6.53 6.93E-222 1.83E-219 22.0 kD class IV heat shock protein Mitochondria
GRMZM2G306714 6.28 0 0 Class I heat shock protein 3 Nuclear
GRMZM2G346839 5.97 7.04E-66 5.89E-64 18.3 kD class I heat shock protein Cytoplasmic
GRMZM2G335242 5.96 0 0 15.7 kD class I heat shock protein Cytoplasmic
GRMZM2G080724 5.61 0 0 Retrotransposon protein [Zea mays] Mitochondrial
GRMZM2G012455 5.19 8.52E-135 1.34E-132 17.6 kD class II heat shock protein Cytoplasmic
GRMZM2G311710 4.98 0 0 17.0 kD class II heat shock protein Nuclear
GRMZM2G333635 4.48 0 0 17.9 kD class I heat shock protein Nuclear
GRMZM2G158232 4.33 0 0 17.9 kD class I heat shock protein Nuclear
GRMZM2G331701 4.31 0 0 18.3 kD class I heat shock Cytoplasmic
GRMZM5G849535 4.30 0 0 16.9 kD class I heat shock protein 1-like Mitochondrial
GRMZM2G034157 4.27 2.84E-22 8.22E-21 17.8 kD class II heat shock protein Cytoplasmic
GRMZM2G083810 4.18 0 0 17.8 kD class II heat shock protein Cytoplasmic
GRMZM2G332512 3.94 1.36E-70 1.22E-68 Heat shock protein18c (hsp18c) Nuclear
GRMZM2G085934 3.88 3.28E-10 4.36E-09 18.6 kD class III heat shock protein Cytoplasmic
GRMZM2G404274 3.88 1.34E-197 3.16E-195 18.0 kD heat shock protein 18a Nuclear
GRMZM2G149647 3.52 0 0 Hsp26 - heat shock protein 26 Chloroplast
GRMZM2G098167 3.49 0 0 17.0 kD class II heat shock protein Cytoplasmic
GRMZM2G046382 3.22 0 0 17.9 kD class I heat shock protein Nuclear
AC208204.3_FG006 3.10 0 0 17.4 kD class I heat shock protein Nuclear

图2

HSP20保守结构域"

表4

ACD结构域序列分析"

基因ID
Gene ID		 ACD氨基酸数
Amino acid number		 个数/百分比Number/percentage (%)
精氨酸 Arg (R) 赖氨酸 Lys (K) 脯氨酸 Pro (P) 苏氨酸 Thr (T)
GRMZM2G481605 109 12/11.0 3/2.7 3/2.7 4/3.7
GRMZM2G049767 96 14/14.6 5/5.2 4/4.2 3/3.1
GRMZM2G306679 92 9/9.8 10/10.9 4/4.3 3/3.3
GRMZM2G479260 92 8/8.7 9/9.8 3/3.3 8/8.7
GRMZM2G375517 87 16/18.4 4/4.6 4/4.6 3/3.4
GRMZM2G346839 93 17/18.3 2/2.3 3/3.2 2/2.2
GRMZM2G335242 97 6/6.2 8/8.2 7/7.2 4/4.1
GRMZM2G080724 97 10/10.3 4/4.1 6/6.2 3/3.1
GRMZM2G012455 97 11/11.3 5/5.2 2/2.1 3/3.1
GRMZM2G333635 92 8/8.7 11/12 4/4.3 3/3.3
GRMZM2G158232 92 8/8.7 11/12 4/4.3 4/4.3
GRMZM2G331701 92 19/20.7 2/2.2 3/3.3 2/2.3
GRMZM2G034157 98 10/10.2 5/5.1 3/3.1 2/2.0
GRMZM2G083810 87 10/11.5 6/6.9 3/3.4 2/2.3
GRMZM2G085934 78 9/11.5 5/6.4 4/5.1 1/1.3
GRMZM2G149647 107 6/5.6 11/10.3 3/2.8 3/2.8
GRMZM2G098167 113 11/9.7 10/8.8 7/6.2 6/5.3
GRMZM2G046382 92 8/8.7 9/9.8 4/4.3 7/7.6
AC208204.3_FG006 92 8/8.7 9/9.8 4/4.3 7/7.6
平均值Mean 95 11/11.2 7/7.2 4 /4.2 4/3.9

附图1

HSP20s基因在不同老化时间内的表达图中红色字体为文中挑选的4个基因。"

图3

HSP20基因在不同老化时间内的表达误差线上不同小写英文字母表示在0.05水平上的差异显著。"

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