中国水稻科学 ›› 2016, Vol. 30 ›› Issue (2): 111-120.DOI: 10.16819/j.1001-7216.2016.5138
• • 下一篇
孟姣1, 王海华1,2,3,*(), 向建华1, 蒋丹1, 彭喜旭1,3, 贺欢欢1
收稿日期:
2015-09-16
修回日期:
2015-11-30
出版日期:
2016-03-10
发布日期:
2016-03-10
通讯作者:
王海华
基金资助:
Jiao MENG1, Hai-hua WANG1,2,3,*(), Jian-hua XIANG1, Dan JIANG1, Xi-xu PENG1,3, Huan-huan HE1
Received:
2015-09-16
Revised:
2015-11-30
Online:
2016-03-10
Published:
2016-03-10
Contact:
Hai-hua WANG
摘要:
一氧化氮(nitric oxide, NO)信号分子与WRKY转录因子均参与植物抗逆、发育与代谢等许多生理过程。采用Agilent水稻全基因组cDNA芯片分析了NO处理后1、6和12 h水稻幼苗WRKY转录因子基因的表达谱,鉴定出在1个时间点有两倍或两倍以上表达变化的WRKY基因32个,主要分布在WRKY的Ⅰ和Ⅱ组,其中75%的Ⅱa和45.6%的Ⅱd亚组成员为差异表达基因;鉴定出至少在2个时间点有两倍或两倍以上表达变化的WRKY基因15个,均为早期(1 h)应答,且多数(64.2%)持续上调;基因功能预测分析表明,这些基因主要参与生物学过程中的细胞过程、代谢过程和刺激响应,以及分子功能中的转录调节活性和结合;代谢通路分析表明,WRKY24涉及植物与病原菌相互作用代谢通路。实时荧光定量PCR 验证结果与芯片杂交结果基本一致,印证了芯片杂交结果的有效性。上述发现提示,NO信号可能参与了WRKY转录因子介导的生物学调控功能,并为这些基因的进一步功能分析奠定基础。
中图分类号:
孟姣, 王海华, 向建华, 蒋丹, 彭喜旭, 贺欢欢. 水稻WRKY转录因子基因家族响应外源一氧化氮的表达谱分析[J]. 中国水稻科学, 2016, 30(2): 111-120.
Jiao MENG, Hai-hua WANG, Jian-hua XIANG, Dan JIANG, Xi-xu PENG, Huan-huan HE. Expression Profiles of Rice WRKY Transcription Factor Gene Family Responsive to Exogenous Nitric Oxide Application[J]. Chinese Journal OF Rice Science, 2016, 30(2): 111-120.
基因 Gene | 登录号 Accession No. | 正向引物 Forward primer (5'-3') | 反向引物 Reverse primer (5'-3') |
---|---|---|---|
OsWRKY15 | Os01g0656400 | GTCGTCGCCCTTCGTGTC | ATGAACTCCATGTCGTCTCCC |
OsWRKY42 | Os02g0462800 | ATGGGAGCGTGTCCAACG | GCGGCCATCAGAAGCATAA |
OsWRKY21 | Os01g0821600 | GCTGAGCAACAGAGCAGGTA | GCGTCAGTTATGCGTCAAG |
OsWRKY77 | Os01g0584900 | CCTCGCTGCTGCTTCCTT | TTCAGGTCGCCTTGGTCC |
OsWRKY71 | Os02g0181300 | AGGAATTGATGATTCGCTGTA | CGCCGATGATCGTTGGTT |
OsWRKY24 | Os01g0826400 | TGCTCCTGACCTCCAGTATCTT | TTCCTCTGCTCGTCCTTGC |
OsWRKY69 | Os08g0386200 | TGCTTCGACTTCGACCCG | TCTTCCCTCCTGCCACCA |
Actin 1 | Os03g0718100 | ATCGCCCTGGACTATGAC | GAAACGCTCAGCACCAAT |
表1 候选基因的qRT-PCR引物
Table 1 Primers of the candidate genes for qRT-PCR.
基因 Gene | 登录号 Accession No. | 正向引物 Forward primer (5'-3') | 反向引物 Reverse primer (5'-3') |
---|---|---|---|
OsWRKY15 | Os01g0656400 | GTCGTCGCCCTTCGTGTC | ATGAACTCCATGTCGTCTCCC |
OsWRKY42 | Os02g0462800 | ATGGGAGCGTGTCCAACG | GCGGCCATCAGAAGCATAA |
OsWRKY21 | Os01g0821600 | GCTGAGCAACAGAGCAGGTA | GCGTCAGTTATGCGTCAAG |
OsWRKY77 | Os01g0584900 | CCTCGCTGCTGCTTCCTT | TTCAGGTCGCCTTGGTCC |
OsWRKY71 | Os02g0181300 | AGGAATTGATGATTCGCTGTA | CGCCGATGATCGTTGGTT |
OsWRKY24 | Os01g0826400 | TGCTCCTGACCTCCAGTATCTT | TTCCTCTGCTCGTCCTTGC |
OsWRKY69 | Os08g0386200 | TGCTTCGACTTCGACCCG | TCTTCCCTCCTGCCACCA |
Actin 1 | Os03g0718100 | ATCGCCCTGGACTATGAC | GAAACGCTCAGCACCAAT |
图1 NO处理不同时间后水稻叶片叶绿素含量 T0、T1、T6、T12分别表示NO处理0、1、6和12 h。下同。
Fig. 1. Chlorophyll contents in rice leaves after NO treatment for different time. NO treatments for 0,1,6 and 12 h are indicated as T0, T1, T6 and T12, respectively. The same as below.
基因 Gene | 探针 Probe | 表达倍数 Expression fold change | 组别 Group | ||
---|---|---|---|---|---|
T1/T0 | T6/T0 | T12/T0 | |||
OsWRKY1 | Os01g0246700|mRNA|AY676928|CDS | 1.87 | 2.06 | 1.35 | Ⅱb |
Os01g0246700|mRNA|AK105509|CDS+3'UTR | 2.72 | 1.54 | 2.19 | ||
OsWRKY3 | Os03g0758000|mRNA|AY341859|CDS+3'UTR | 0.33 | 0.38 | 3.70 | Ⅰb |
OsWRKY6 | Os03g0798500|COMBINER_EST|Os03g0798500|8 | 0.59 | 0.57 | 2.25 | Ⅱc |
OsWRKY8 | Os05g0583000|mRNA|AK109568|CDS+3'UTR | 2.95 | 1.13 | 0.44 | Ⅱd |
OsWRKY11 | Os01g0626400|mRNA|AY341856|CDS+3'UTR | 1.91 | 2.88 | 1.60 | Ⅰb |
Os01g0626400|mRNA|AK108745|CDS+3'UTR | 2.23 | 2.57 | 1.63 | ||
OsWRKY12 | Os01g0624700|mRNA|AK111416|CDS+3'UTR | 3.14 | 2.74 | 1.99 | Ⅱd |
OsWRKY13 | Os01g0750100|mRNA|AK060018|CDS+3'UTR | 1.55 | 0.67 | 0.48 | Ⅱd |
OsWRKY15 | Os01g0656400|mRNA|AY676926|CDS | 4.98 | 1.16 | 2.70 | Ⅲa |
OsWRKY17 | Os01g0972800|mRNA|AY341842|CDS+3'UTR | 0.09 | 0.09 | 0.32 | Ⅰb |
OsWRKY21 | Os01g0821600|mRNA|AK108657|CDS+3'UTR | 4.69 | 4.42 | 1.31 | Ⅲb |
OsWRKY23 | Os01g0734000|mRNA|AK108909|CDS+3'UTR | 2.24 | 1.69 | 2.11 | Ⅰb |
OsWRKY24 | Os01g0826400|mRNA|AK107199|CDS+3'UTR | 6.48 | 0.83 | 3.86 | Ⅰa |
Os01g0826400|mRNA|AY341849|CDS+3'UTR | 5.39 | 0.63 | 3.33 | ||
Os01g0826400|mRNA|AY676925|CDS | 5.92 | 0.75 | 3.65 | ||
OsWRKY28 | Os06g0649000|mRNA|AK106282|CDS+3'UTR | 7.87 | 1.32 | 2.28 | Ⅱa |
Os06g0649000|mRNA|AK119644|CDS+3'UTR | 6.77 | 1.05 | 1.91 | ||
OsWRKY29 | Os07g0111400|mRNA|AY341858|CDS+3'UTR | 0.73 | 1.32 | 2.70 | Ⅰb |
OsWRKY36 | Os04g0545000|mRNA|AK073695|CDS+3'UTR | 0.47 | 0.73 | 0.61 | Ⅰb |
OsWRKY39 | Os02g0265200|mRNA|AK066775|CDS+3'UTR | 1.43 | 2.00 | 2.62 | Ⅱd |
OsWRKY42 | Os02g0265200|mRNA|AK119593|CDS+3'UTR | 1.42 | 2.27 | 2.34 | Ⅱc |
Os02g0462800|mRNA|AK110587|CDS+3'UTR | 5.51 | 3.80 | 2.01 | ||
Os02g0462800|mRNA|AY676923|CDS | 5.01 | 3.14 | 2.05 | ||
OsWRKY50 | Os11g0117600|COMBINER_EST|Os11g0117600|8 | 0.65 | 7.31 | 1.11 | Ⅲb |
OsWRKY53 | Os05g0343400|mRNA|AY676929|CDS | 2.10 | 0.89 | 1.02 | Ⅰa |
OsWRKY56 | Os12g0116600|mRNA|AK102093|CDS+3'UTR | 1.31 | 0.42 | 1.03 | Ⅲb |
OsWRKY60 | Os03g0657400|COMBINER_EST|AU057193|6 | 1.01 | 0.58 | 0.44 | Ⅰb |
OsWRKY62 | Os09g0417800|mRNA|AK067834|CDS+3'UTR | 1.48 | 1.65 | 2.03 | Ⅱa |
OsWRKY67 | Os05g0183100|mRNA|AK066252|CDS+3'UTR | 1.13 | 0.66 | 0.28 | Ⅰb |
OsWRKY69 | Os08g0386200|mRNA|AK111606|CDS+3'UTR | 3.05 | 0.98 | 2.77 | Ⅲa |
OsWRKY71 | Os08g0386200|mRNA|AY676931|CDS | 3.33 | 0.88 | 2.80 | Ⅱa |
Os02g0181300|mRNA|AK058773|CDS+3'UTR | 3.79 | 1.72 | 1.67 | ||
Os02g0181300|mRNA|AY676927|CDS | 3.41 | 1.45 | 1.64 | ||
OsWRKY72 | Os11g0490900|mRNA|AK108860|CDS+3'UTR | 2.32 | 0.89 | 1.75 | Ⅰb |
OsWRKY77 | Os01g0584900|mRNA|AK108522|CDS+3'UTR | 1.10 | 1.80 | 2.31 | Ⅰb |
Os01g0584900|mRNA|AY341846|CDS+3'UTR | 1.24 | 1.88 | 2.48 | ||
OsWRKY79 | Os03g0335200|mRNA|AK105244|CDS+3'UTR | 0.66 | 1.83 | 2.53 | Ⅲb |
OsWRKY88 | Os07g0596900|mRNA|AY341847|CDS+3'UTR | 0.94 | 1.53 | 2.13 | Ⅲa |
OsWRKY104 | Os11g0117400|COMBINER_EST|CI444468|6 | 1.40 | 0.32 | 0.93 | Ⅲb |
OsWRKY109 | Os05g0129800|COMBINER_EST|Os05g0129800|8 | 0.73 | 0.82 | 2.02 | Ⅱd |
OsWRKY113 | Os06g0158100|mRNA|AY676932|CDS | 2.35 | 0.96 | 3.71 | Ⅲa |
表2 NO处理下水稻WRKY差异表达基因的芯片分析
Table 2 Microarray analysis of differentially expressed WRKY genes in rice under NO treatment.
基因 Gene | 探针 Probe | 表达倍数 Expression fold change | 组别 Group | ||
---|---|---|---|---|---|
T1/T0 | T6/T0 | T12/T0 | |||
OsWRKY1 | Os01g0246700|mRNA|AY676928|CDS | 1.87 | 2.06 | 1.35 | Ⅱb |
Os01g0246700|mRNA|AK105509|CDS+3'UTR | 2.72 | 1.54 | 2.19 | ||
OsWRKY3 | Os03g0758000|mRNA|AY341859|CDS+3'UTR | 0.33 | 0.38 | 3.70 | Ⅰb |
OsWRKY6 | Os03g0798500|COMBINER_EST|Os03g0798500|8 | 0.59 | 0.57 | 2.25 | Ⅱc |
OsWRKY8 | Os05g0583000|mRNA|AK109568|CDS+3'UTR | 2.95 | 1.13 | 0.44 | Ⅱd |
OsWRKY11 | Os01g0626400|mRNA|AY341856|CDS+3'UTR | 1.91 | 2.88 | 1.60 | Ⅰb |
Os01g0626400|mRNA|AK108745|CDS+3'UTR | 2.23 | 2.57 | 1.63 | ||
OsWRKY12 | Os01g0624700|mRNA|AK111416|CDS+3'UTR | 3.14 | 2.74 | 1.99 | Ⅱd |
OsWRKY13 | Os01g0750100|mRNA|AK060018|CDS+3'UTR | 1.55 | 0.67 | 0.48 | Ⅱd |
OsWRKY15 | Os01g0656400|mRNA|AY676926|CDS | 4.98 | 1.16 | 2.70 | Ⅲa |
OsWRKY17 | Os01g0972800|mRNA|AY341842|CDS+3'UTR | 0.09 | 0.09 | 0.32 | Ⅰb |
OsWRKY21 | Os01g0821600|mRNA|AK108657|CDS+3'UTR | 4.69 | 4.42 | 1.31 | Ⅲb |
OsWRKY23 | Os01g0734000|mRNA|AK108909|CDS+3'UTR | 2.24 | 1.69 | 2.11 | Ⅰb |
OsWRKY24 | Os01g0826400|mRNA|AK107199|CDS+3'UTR | 6.48 | 0.83 | 3.86 | Ⅰa |
Os01g0826400|mRNA|AY341849|CDS+3'UTR | 5.39 | 0.63 | 3.33 | ||
Os01g0826400|mRNA|AY676925|CDS | 5.92 | 0.75 | 3.65 | ||
OsWRKY28 | Os06g0649000|mRNA|AK106282|CDS+3'UTR | 7.87 | 1.32 | 2.28 | Ⅱa |
Os06g0649000|mRNA|AK119644|CDS+3'UTR | 6.77 | 1.05 | 1.91 | ||
OsWRKY29 | Os07g0111400|mRNA|AY341858|CDS+3'UTR | 0.73 | 1.32 | 2.70 | Ⅰb |
OsWRKY36 | Os04g0545000|mRNA|AK073695|CDS+3'UTR | 0.47 | 0.73 | 0.61 | Ⅰb |
OsWRKY39 | Os02g0265200|mRNA|AK066775|CDS+3'UTR | 1.43 | 2.00 | 2.62 | Ⅱd |
OsWRKY42 | Os02g0265200|mRNA|AK119593|CDS+3'UTR | 1.42 | 2.27 | 2.34 | Ⅱc |
Os02g0462800|mRNA|AK110587|CDS+3'UTR | 5.51 | 3.80 | 2.01 | ||
Os02g0462800|mRNA|AY676923|CDS | 5.01 | 3.14 | 2.05 | ||
OsWRKY50 | Os11g0117600|COMBINER_EST|Os11g0117600|8 | 0.65 | 7.31 | 1.11 | Ⅲb |
OsWRKY53 | Os05g0343400|mRNA|AY676929|CDS | 2.10 | 0.89 | 1.02 | Ⅰa |
OsWRKY56 | Os12g0116600|mRNA|AK102093|CDS+3'UTR | 1.31 | 0.42 | 1.03 | Ⅲb |
OsWRKY60 | Os03g0657400|COMBINER_EST|AU057193|6 | 1.01 | 0.58 | 0.44 | Ⅰb |
OsWRKY62 | Os09g0417800|mRNA|AK067834|CDS+3'UTR | 1.48 | 1.65 | 2.03 | Ⅱa |
OsWRKY67 | Os05g0183100|mRNA|AK066252|CDS+3'UTR | 1.13 | 0.66 | 0.28 | Ⅰb |
OsWRKY69 | Os08g0386200|mRNA|AK111606|CDS+3'UTR | 3.05 | 0.98 | 2.77 | Ⅲa |
OsWRKY71 | Os08g0386200|mRNA|AY676931|CDS | 3.33 | 0.88 | 2.80 | Ⅱa |
Os02g0181300|mRNA|AK058773|CDS+3'UTR | 3.79 | 1.72 | 1.67 | ||
Os02g0181300|mRNA|AY676927|CDS | 3.41 | 1.45 | 1.64 | ||
OsWRKY72 | Os11g0490900|mRNA|AK108860|CDS+3'UTR | 2.32 | 0.89 | 1.75 | Ⅰb |
OsWRKY77 | Os01g0584900|mRNA|AK108522|CDS+3'UTR | 1.10 | 1.80 | 2.31 | Ⅰb |
Os01g0584900|mRNA|AY341846|CDS+3'UTR | 1.24 | 1.88 | 2.48 | ||
OsWRKY79 | Os03g0335200|mRNA|AK105244|CDS+3'UTR | 0.66 | 1.83 | 2.53 | Ⅲb |
OsWRKY88 | Os07g0596900|mRNA|AY341847|CDS+3'UTR | 0.94 | 1.53 | 2.13 | Ⅲa |
OsWRKY104 | Os11g0117400|COMBINER_EST|CI444468|6 | 1.40 | 0.32 | 0.93 | Ⅲb |
OsWRKY109 | Os05g0129800|COMBINER_EST|Os05g0129800|8 | 0.73 | 0.82 | 2.02 | Ⅱd |
OsWRKY113 | Os06g0158100|mRNA|AY676932|CDS | 2.35 | 0.96 | 3.71 | Ⅲa |
组/亚组 Group/ subgroup | 差异表达基因数 Number of differentially expressed genes | 组/亚组的基因总数 Total number of genes in group/subgroup | 比例 Ratio /% |
---|---|---|---|
Ⅰ | 12 | 34 | 35.3 |
Ⅱ | 11 | 30 | 36.7 |
a | 3 | 4 | 75.0 |
b | 1 | 8 | 12.5 |
c | 2 | 7 | 28.6 |
d | 5 | 11 | 45.6 |
Ⅲ | 9 | 36 | 25.0 |
表3 NO处理下水稻WRKY差异表达基因在各组中的分布
Table 3 Group distribution of differentially expressed WRKY genes of rice responsive to nitric oxide.
组/亚组 Group/ subgroup | 差异表达基因数 Number of differentially expressed genes | 组/亚组的基因总数 Total number of genes in group/subgroup | 比例 Ratio /% |
---|---|---|---|
Ⅰ | 12 | 34 | 35.3 |
Ⅱ | 11 | 30 | 36.7 |
a | 3 | 4 | 75.0 |
b | 1 | 8 | 12.5 |
c | 2 | 7 | 28.6 |
d | 5 | 11 | 45.6 |
Ⅲ | 9 | 36 | 25.0 |
图3 NO响应的水稻WRKY基因的维恩图(A)和聚类热图(B) 比色刻度尺在上边。红色表示上调,绿色表示下调。
Fig.3. Venny analysis (A) and heatmap clustering (B) of NO-responsive rice WRKY gene. The color scale is shown on the top. Red, Up-regulated; Green, Down-regulated.
图5 NO响应的水稻WRKY基因的表达模式 Log2FC-Log2处理后的基因表达变化倍数。
Fig.5. Expression patterns of the NO-responsive rice WRKY genes. Log2FC indicates Log2 transformed expression fold change.
图6 NO响应的水稻WRKY基因表达水平的qRT-PCR验证 Log2FC-Log2处理后的基因表达变化倍数。
Fig. 6. Expression verification of NO-responsive rice WRKY genes by qRT-PCR. Log2FC indicates Log2 transformed expression fold change.
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