草业学报 ›› 2023, Vol. 32 ›› Issue (3): 91-106.DOI: 10.11686/cyxb2022094
李雯1,2(), 赵丽蓉1,2, 张建平1,2(), 刘自刚1(), 齐燕妮2, 李闻娟2, 谢亚萍2
收稿日期:
2022-02-23
修回日期:
2022-04-06
出版日期:
2023-03-20
发布日期:
2022-12-30
通讯作者:
张建平,刘自刚
作者简介:
E-mail: 739015868@qq.com基金资助:
Wen LI1,2(), Li-rong ZHAO1,2, Jian-ping ZHANG1,2(), Zi-gang LIU1(), Yan-ni QI2, Wen-juan LI2, Ya-ping XIE2
Received:
2022-02-23
Revised:
2022-04-06
Online:
2023-03-20
Published:
2022-12-30
Contact:
Jian-ping ZHANG,Zi-gang LIU
摘要:
DMP家族成员是植物特有的膜蛋白,在植物生殖发育及衰老过程中发挥重要作用。亚麻是我国重要的油料及经济作物,目前关于DMP基因家族在亚麻中的分布及功能研究尚未有报道。本研究利用生物信息学方法对亚麻DMP基因家族进行鉴定,对其序列特征、系统进化、复制事件进行了分析,并利用转录组数据和qRT-PCR分析其在不同组织器官及胁迫处理下的表达模式。本研究以陇亚10号基因组为参照,利用同源性分析共鉴定到17个LuDMP成员,命名为LuDMP-1~LuDMP-17,不均匀分布在9条染色体上。序列特征分析结果表明,除LuDMP-9、LuDMP-16和LuDMP-17 外,其余成员均不含内含子;基因启动子区含有大量激素与逆境胁迫相关响应元件;编码的蛋白氨基酸长度为195~240 aa,分子量为20946~25830 Da,等电点为4.76~9.71;所有蛋白均为疏水性蛋白,含有2~5个跨膜结构域及4~8个motif,其中有4个motif存在于所有LuDMP蛋白;绝大多数蛋白定位于细胞膜。LuDMP家族共有3对串联复制事件和8对片段复制事件,进化过程中经历了强烈的纯化选择。转录组分析结果表明,大多数LuDMP成员在果实和茎中呈现特异性表达模式。系统进化分析将 DMP蛋白分为5个亚族,第III亚族DMP数量最多,含有11个LuDMP成员,各亚族中DMP蛋白具有单子叶或双子叶特异性聚类模式。第IV亚族中的LuDMP-1和LuDMP-7与拟南芥及玉米单倍体诱导基因具有很近的亲缘关系,位于同一进化分支,序列同源性达到67%以上,而且该基因在成熟花药中的表达量明显高于其他组织,推测其为亚麻单倍体诱导的候选基因。本研究在陇亚10号中克隆到了LuDMP-1,全长为699 bp。LuDMP-1/7启动子区域含有激素响应及逆境胁迫相关作用元件,吲哚乙酸、赤霉素、萘乙酸、聚乙二醇、高温及低温处理下, LuDMP-1/7基因表达量均有不同程度的提高,其中对低温和高温的响应最明显,分别较对照提高了约260和600倍。本研究为阐明亚麻DMP蛋白的功能奠定了基础,并为亚麻单倍体育种提供了基因资源。
李雯, 赵丽蓉, 张建平, 刘自刚, 齐燕妮, 李闻娟, 谢亚萍. 亚麻DMP基因家族的全基因组鉴定与分析[J]. 草业学报, 2023, 32(3): 91-106.
Wen LI, Li-rong ZHAO, Jian-ping ZHANG, Zi-gang LIU, Yan-ni QI, Wen-juan LI, Ya-ping XIE. Genome-wide identification and analysis of the DMP gene family in flax (Linum usitatissimum)[J]. Acta Prataculturae Sinica, 2023, 32(3): 91-106.
引物名称Primer name | 序列Primer sequence (5′-3′) | 用途Usage |
---|---|---|
DMP-F | ATGGAGGAAGACGACTATGGAATCA | 基因克隆Gene cloning |
DMP-R | CTAACTAGCACTGCACCCAATCCC | |
qDMP-F | CACTTCACGGACAGTTTCAAGGATC | 定量反转录聚合酶连锁反应Quantificational RT-PCR |
qDMP-R | CAAGCCACTACAAACAACACCAACC | |
GAPDH-F | CTTTACCCTCAGCAAATCCG | 内参Reference gene |
GAPDH-R | AGGTTCTTCCCGCTCTCAAT |
表1 本研究所用引物
Table 1 Primers used in this research
引物名称Primer name | 序列Primer sequence (5′-3′) | 用途Usage |
---|---|---|
DMP-F | ATGGAGGAAGACGACTATGGAATCA | 基因克隆Gene cloning |
DMP-R | CTAACTAGCACTGCACCCAATCCC | |
qDMP-F | CACTTCACGGACAGTTTCAAGGATC | 定量反转录聚合酶连锁反应Quantificational RT-PCR |
qDMP-R | CAAGCCACTACAAACAACACCAACC | |
GAPDH-F | CTTTACCCTCAGCAAATCCG | 内参Reference gene |
GAPDH-R | AGGTTCTTCCCGCTCTCAAT |
基因名Gene name | 基因号 Gene-ID | 染色体位置 Chromosome location | 外显子数 No. of exons | 蛋白 长度 Protein length (aa) | 总平均亲水性Grand average of hydropathicity (GRAVY) | 等电点 Isoelectric point | 分子量 Molecular weight (Da) | 亚细胞定位 Subcellular location | 跨膜结 构域 Transmembrane domain |
---|---|---|---|---|---|---|---|---|---|
LuDMP-1 | L.us.o.g.scaffold34.203 | Chr1:885721-886419(+) | 1 | 232 | 0.107 | 8.08 | 25392.66 | CM,Nucleus | 5 |
LuDMP-2 | L.us.o.g.scaffold0.456 | Chr2:21280161-21280973(+) | 1 | 198 | 0.231 | 8.63 | 21293.48 | CM | 4 |
LuDMP-3 | L.us.o.g.scaffold0.36 | Chr2:21635014-21637816(-) | 1 | 195 | 0.244 | 8.28 | 20946.10 | CM | 4 |
LuDMP-4 | L.us.o.g.scaffold30.59 | Chr4:20534605-20535300(+) | 1 | 231 | 0.202 | 4.89 | 25126.95 | CM | 2 |
LuDMP-5 | L.us.o.g.scaffold30.58 | Chr4:20536021-20536638(+) | 1 | 205 | 0.364 | 6.93 | 22745.70 | CP | 3 |
LuDMP-6 | L.us.o.g.scaffold30.57 | Chr4:20537649-20538317(+) | 1 | 222 | 0.310 | 6.18 | 23847.58 | CM | 4 |
LuDMP-7 | L.us.o.g.scaffold70.212 | Chr7:9831303-9832001(+) | 1 | 232 | 0.057 | 8.08 | 25375.63 | CM,Nucleus | 4 |
LuDMP-8 | L.us.o.g.scaffold136.194 | Chr8:1238120-1238776(+) | 1 | 218 | 0.064 | 9.19 | 23883.09 | CP | 2 |
LuDMP-9 | L.us.o.g.scaffold69.277 | Chr10:17296812-17298353(-) | 2 | 229 | 0.093 | 9.71 | 25372.99 | CM | 5 |
LuDMP-10 | L.us.o.g.scaffold305.10 | Chr12:5379664-5380827(-) | 1 | 217 | 0.313 | 9.06 | 23577.41 | CM | 4 |
LuDMP-11 | L.us.o.g.scaffold48.30 | Chr14:1446411-1447221(+) | 1 | 237 | 0.061 | 9.19 | 23746.92 | CM,CP | 4 |
LuDMP-12 | L.us.o.g.scaffold28.148 | Chr15:5929885-5930780(-) | 1 | 226 | 0.283 | 9.28 | 24020.92 | CM | 4 |
LuDMP-13 | L.us.o.g.scaffold107.51 | Chr15:7942536-7943540(+) | 1 | 233 | 0.184 | 4.76 | 25373.19 | CM | 2 |
LuDMP-14 | L.us.o.g.scaffold107.50 | Chr15:7944336-7945076(+) | 1 | 205 | 0.582 | 6.50 | 22782.72 | CM,Nucleus | 3 |
LuDMP-15 | L.us.o.g.scaffold107.49 | Chr15:7946004-7946771(+) | 1 | 222 | 0.252 | 6.18 | 23865.56 | CM | 4 |
LuDMP-16 | L.us.o.g.scaffold107.46 | Chr15:7954525-7955820(+) | 2 | 230 | 0.275 | 5.28 | 24596.32 | CM | 4 |
LuDMP-17 | L.us.o.g.scaffold107.45 | Chr15: 7957608-7959009(+) | 2 | 240 | 0.256 | 5.01 | 25829.65 | CM | 2 |
表2 亚麻LuDMP成员基本特征
Table 2 Basic characteristics of flax LuDMP members
基因名Gene name | 基因号 Gene-ID | 染色体位置 Chromosome location | 外显子数 No. of exons | 蛋白 长度 Protein length (aa) | 总平均亲水性Grand average of hydropathicity (GRAVY) | 等电点 Isoelectric point | 分子量 Molecular weight (Da) | 亚细胞定位 Subcellular location | 跨膜结 构域 Transmembrane domain |
---|---|---|---|---|---|---|---|---|---|
LuDMP-1 | L.us.o.g.scaffold34.203 | Chr1:885721-886419(+) | 1 | 232 | 0.107 | 8.08 | 25392.66 | CM,Nucleus | 5 |
LuDMP-2 | L.us.o.g.scaffold0.456 | Chr2:21280161-21280973(+) | 1 | 198 | 0.231 | 8.63 | 21293.48 | CM | 4 |
LuDMP-3 | L.us.o.g.scaffold0.36 | Chr2:21635014-21637816(-) | 1 | 195 | 0.244 | 8.28 | 20946.10 | CM | 4 |
LuDMP-4 | L.us.o.g.scaffold30.59 | Chr4:20534605-20535300(+) | 1 | 231 | 0.202 | 4.89 | 25126.95 | CM | 2 |
LuDMP-5 | L.us.o.g.scaffold30.58 | Chr4:20536021-20536638(+) | 1 | 205 | 0.364 | 6.93 | 22745.70 | CP | 3 |
LuDMP-6 | L.us.o.g.scaffold30.57 | Chr4:20537649-20538317(+) | 1 | 222 | 0.310 | 6.18 | 23847.58 | CM | 4 |
LuDMP-7 | L.us.o.g.scaffold70.212 | Chr7:9831303-9832001(+) | 1 | 232 | 0.057 | 8.08 | 25375.63 | CM,Nucleus | 4 |
LuDMP-8 | L.us.o.g.scaffold136.194 | Chr8:1238120-1238776(+) | 1 | 218 | 0.064 | 9.19 | 23883.09 | CP | 2 |
LuDMP-9 | L.us.o.g.scaffold69.277 | Chr10:17296812-17298353(-) | 2 | 229 | 0.093 | 9.71 | 25372.99 | CM | 5 |
LuDMP-10 | L.us.o.g.scaffold305.10 | Chr12:5379664-5380827(-) | 1 | 217 | 0.313 | 9.06 | 23577.41 | CM | 4 |
LuDMP-11 | L.us.o.g.scaffold48.30 | Chr14:1446411-1447221(+) | 1 | 237 | 0.061 | 9.19 | 23746.92 | CM,CP | 4 |
LuDMP-12 | L.us.o.g.scaffold28.148 | Chr15:5929885-5930780(-) | 1 | 226 | 0.283 | 9.28 | 24020.92 | CM | 4 |
LuDMP-13 | L.us.o.g.scaffold107.51 | Chr15:7942536-7943540(+) | 1 | 233 | 0.184 | 4.76 | 25373.19 | CM | 2 |
LuDMP-14 | L.us.o.g.scaffold107.50 | Chr15:7944336-7945076(+) | 1 | 205 | 0.582 | 6.50 | 22782.72 | CM,Nucleus | 3 |
LuDMP-15 | L.us.o.g.scaffold107.49 | Chr15:7946004-7946771(+) | 1 | 222 | 0.252 | 6.18 | 23865.56 | CM | 4 |
LuDMP-16 | L.us.o.g.scaffold107.46 | Chr15:7954525-7955820(+) | 2 | 230 | 0.275 | 5.28 | 24596.32 | CM | 4 |
LuDMP-17 | L.us.o.g.scaffold107.45 | Chr15: 7957608-7959009(+) | 2 | 240 | 0.256 | 5.01 | 25829.65 | CM | 2 |
基因ID Gene ID | AtDMP8 | AtDMP9 | ZmDMP | |||
---|---|---|---|---|---|---|
E值 E value | 同源性Per ident (%) | E值 E value | 同源性 Per ident (%) | E值 E value | 同源性 Per ident (%) | |
LuDMP-1 | 1×1086 | 57.55 | 3×1086 | 67.39 | 2×1090 | 67.98 |
LuDMP-2 | 2×1034 | 33.15 | 9×1033 | 32.58 | 9×1028 | 36.05 |
LuDMP-3 | 9×1036 | 33.71 | 2×1033 | 32.58 | 3×1029 | 35.80 |
LuDMP-4 | 2×1047 | 33.76 | 6×1046 | 38.71 | 4×1045 | 44.94 |
LuDMP-5 | 7×1043 | 35.68 | 4×1041 | 35.14 | 6×1041 | 39.33 |
LuDMP-6 | 1×1042 | 34.76 | 1×1042 | 36.22 | 3×1040 | 39.11 |
LuDMP-7 | 1×1088 | 57.96 | 4×1086 | 67.39 | 1×1090 | 67.98 |
LuDMP-8 | 6×1041 | 39.56 | 2×1038 | 38.46 | 1×1036 | 39.33 |
LuDMP-9 | 7×1028 | 33.88 | 8×1026 | 32.07 | 5×1025 | 33.88 |
LuDMP-10 | 7×1046 | 36.94 | 6×1043 | 39.13 | 5×1044 | 43.58 |
LuDMP-11 | 4×1040 | 37.50 | 1×1037 | 36.41 | 1×1035 | 37.64 |
LuDMP-12 | 2×1041 | 36.02 | 8×1040 | 36.56 | 1×1039 | 44.89 |
LuDMP-13 | 3×1047 | 33.47 | 7×1046 | 38.71 | 1×1045 | 45.51 |
LuDMP-14 | 5×1041 | 34.59 | 7×1040 | 33.51 | 6×1040 | 38.76 |
LuDMP-15 | 2×1042 | 35.11 | 9×1044 | 36.76 | 8×1040 | 40.56 |
LuDMP-16 | 1×1040 | 35.48 | 3×1042 | 37.30 | 7×1041 | 40.45 |
LuDMP-17 | 4×1044 | 37.63 | 2×1045 | 38.38 | 9×1043 | 42.46 |
表3 LuDMP基因同源性比对
Table 3 LuDMP gene homology comparison
基因ID Gene ID | AtDMP8 | AtDMP9 | ZmDMP | |||
---|---|---|---|---|---|---|
E值 E value | 同源性Per ident (%) | E值 E value | 同源性 Per ident (%) | E值 E value | 同源性 Per ident (%) | |
LuDMP-1 | 1×1086 | 57.55 | 3×1086 | 67.39 | 2×1090 | 67.98 |
LuDMP-2 | 2×1034 | 33.15 | 9×1033 | 32.58 | 9×1028 | 36.05 |
LuDMP-3 | 9×1036 | 33.71 | 2×1033 | 32.58 | 3×1029 | 35.80 |
LuDMP-4 | 2×1047 | 33.76 | 6×1046 | 38.71 | 4×1045 | 44.94 |
LuDMP-5 | 7×1043 | 35.68 | 4×1041 | 35.14 | 6×1041 | 39.33 |
LuDMP-6 | 1×1042 | 34.76 | 1×1042 | 36.22 | 3×1040 | 39.11 |
LuDMP-7 | 1×1088 | 57.96 | 4×1086 | 67.39 | 1×1090 | 67.98 |
LuDMP-8 | 6×1041 | 39.56 | 2×1038 | 38.46 | 1×1036 | 39.33 |
LuDMP-9 | 7×1028 | 33.88 | 8×1026 | 32.07 | 5×1025 | 33.88 |
LuDMP-10 | 7×1046 | 36.94 | 6×1043 | 39.13 | 5×1044 | 43.58 |
LuDMP-11 | 4×1040 | 37.50 | 1×1037 | 36.41 | 1×1035 | 37.64 |
LuDMP-12 | 2×1041 | 36.02 | 8×1040 | 36.56 | 1×1039 | 44.89 |
LuDMP-13 | 3×1047 | 33.47 | 7×1046 | 38.71 | 1×1045 | 45.51 |
LuDMP-14 | 5×1041 | 34.59 | 7×1040 | 33.51 | 6×1040 | 38.76 |
LuDMP-15 | 2×1042 | 35.11 | 9×1044 | 36.76 | 8×1040 | 40.56 |
LuDMP-16 | 1×1040 | 35.48 | 3×1042 | 37.30 | 7×1041 | 40.45 |
LuDMP-17 | 4×1044 | 37.63 | 2×1045 | 38.38 | 9×1043 | 42.46 |
图2 LuDMP-1基因全长扩增M:DL1000 marker;1:阴性对照;2:DNA扩增;3:cDNA扩增。1: Negative control; 2: DNA amplification; 3: cDNA amplification.
Fig.2 Full-length amplification of LuDMP-1
图6 LuDMP基因染色体定位横线代表片段复制,方框代表串联复制。The horizontal line represents segmental duplication, and the box represents tandem duplication. Chr: 染色体Chromosome.
Fig.6 Chromosomal locations of LuDMP genes
基因对 Gene pair | 复制事件 Duplication events | 同义替换率 Ks | 非同义替换率 Ka | Ka/Ks | 选择类型 Select type | 分歧时间 Divergence Time |
---|---|---|---|---|---|---|
LuDMP-4/LuDMP-5 | 串联复制 Tandem duplication | 1.6616 | 0.2819 | 0.169656 | 纯化选择 Purifying selection | 136.19670 |
LuDMP-13/LuDMP-14 | 串联复制 Tandem duplication | 2.0752 | 0.2957 | 0.142492 | 纯化选择 Purifying selection | 170.09840 |
LuDMP-16/LuDMP-17 | 串联复制 Tandem duplication | 0.8487 | 0.6472 | 0.762578 | 纯化选择 Purifying selection | 69.56557 |
LuDMP-1/LuDMP-7 | 片段复制 Segmental duplication | 0.2499 | 0.0097 | 0.038816 | 纯化选择 Purifying selection | 20.48361 |
LuDMP-2/LuDMP-3 | 片段复制 Segmental duplication | 0.1296 | 0.0278 | 0.214506 | 纯化选择 Purifying selection | 10.62295 |
LuDMP-4/LuDMP-13 | 片段复制 Segmental duplication | 0.2225 | 0.0025 | 0.011236 | 纯化选择 Purifying selection | 18.23770 |
LuDMP-6/LuDMP-9 | 片段复制 Segmental duplication | n.a. | 0.4789 | n.a. | n.a. | n.a. |
LuDMP-8/LuDMP-9 | 片段复制 Segmental duplication | 1.6678 | 0.2323 | 0.139285 | 纯化选择 Purifying selection | 136.70490 |
LuDMP-8/LuDMP-11 | 片段复制 Segmental duplication | 0.2826 | 0.0484 | 0.171267 | 纯化选择 Purifying selection | 23.16393 |
LuDMP-9/LuDMP-11 | 片段复制 Segmental duplication | 1.6134 | 0.2380 | 0.147515 | 纯化选择 Purifying selection | 132.24590 |
LuDMP-9/LuDMP-15 | 片段复制 Segmental duplication | n.a. | 0.4402 | n.a. | n.a. | n.a. |
表4 LuDMP基因复制分析
Table 4 Duplication analysis of LuDMP genes
基因对 Gene pair | 复制事件 Duplication events | 同义替换率 Ks | 非同义替换率 Ka | Ka/Ks | 选择类型 Select type | 分歧时间 Divergence Time |
---|---|---|---|---|---|---|
LuDMP-4/LuDMP-5 | 串联复制 Tandem duplication | 1.6616 | 0.2819 | 0.169656 | 纯化选择 Purifying selection | 136.19670 |
LuDMP-13/LuDMP-14 | 串联复制 Tandem duplication | 2.0752 | 0.2957 | 0.142492 | 纯化选择 Purifying selection | 170.09840 |
LuDMP-16/LuDMP-17 | 串联复制 Tandem duplication | 0.8487 | 0.6472 | 0.762578 | 纯化选择 Purifying selection | 69.56557 |
LuDMP-1/LuDMP-7 | 片段复制 Segmental duplication | 0.2499 | 0.0097 | 0.038816 | 纯化选择 Purifying selection | 20.48361 |
LuDMP-2/LuDMP-3 | 片段复制 Segmental duplication | 0.1296 | 0.0278 | 0.214506 | 纯化选择 Purifying selection | 10.62295 |
LuDMP-4/LuDMP-13 | 片段复制 Segmental duplication | 0.2225 | 0.0025 | 0.011236 | 纯化选择 Purifying selection | 18.23770 |
LuDMP-6/LuDMP-9 | 片段复制 Segmental duplication | n.a. | 0.4789 | n.a. | n.a. | n.a. |
LuDMP-8/LuDMP-9 | 片段复制 Segmental duplication | 1.6678 | 0.2323 | 0.139285 | 纯化选择 Purifying selection | 136.70490 |
LuDMP-8/LuDMP-11 | 片段复制 Segmental duplication | 0.2826 | 0.0484 | 0.171267 | 纯化选择 Purifying selection | 23.16393 |
LuDMP-9/LuDMP-11 | 片段复制 Segmental duplication | 1.6134 | 0.2380 | 0.147515 | 纯化选择 Purifying selection | 132.24590 |
LuDMP-9/LuDMP-15 | 片段复制 Segmental duplication | n.a. | 0.4402 | n.a. | n.a. | n.a. |
顺式元件 Cis-element | 典型序列 Typical sequence | 特性 Characteristic | 基因Gene |
---|---|---|---|
TATA-box | TATA | 转录开始的-30左右的核心启动子元件Core promoter element around -30 of transcript-ion start | LuDMP-1、LuDMP-2、LuDMP-4、LuDMP-5、LuDMP-6、LuDMP-7、LuDMP-8、LuDMP-9、LuDMP-10、LuDMP-11、LuDMP-12、LuDMP-13、LuDMP-14、 LuDMP-15、LuDMP-16、LuDMP-17 |
ARE | AAACCA | 厌氧诱导所必需的顺式调节元件Cis-acting regulatory element essential for the anaerobic induction | LuDMP-1、LuDMP-2、LuDMP-3、LuDMP-4、LuDMP-5、LuDMP-6、LuDMP-7、LuDMP-8、LuDMP-9、LuDMP-10、LuDMP-11、LuDMP-12、LuDMP-13、LuDMP-14、 LuDMP-15、LuDMP-16、LuDMP-17 |
Sp1 | GGGCGG | 光响应元件Light responsive element | LuDMP-1、LuDMP-3、LuDMP-6、LuDMP-8 |
ABRE | ACGTG | 参与脱落酸反应的顺式作用元件Cis-acting element involved in the abcisic acid responsiveness | LuDMP-1、LuDMP-4、LuDMP-5、LuDMP-6、LuDMP-7、LuDMP-8、LuDMP-9、LuDMP-10、LuDMP-11、LuDMP-13、LuDMP-16 |
TGACG-motif | TGACG | 参与甲基茉莉酸(MeJA)反应的顺式调节元件Cis-acting regulatory element involved in the MeJA-responsiveness | LuDMP-1、LuDMP-4、LuDMP-5、LuDMP-7、LuDMP-8、LuDMP-9、LuDMP-12、LuDMP-13、LuDMP-14、 LuDMP-15、LuDMP-16、LuDMP-17 |
CGTCA-motif | CGTCA | 参与甲基茉莉酸(MeJA)反应的顺式调节元件Cis-acting regulatory element involved in the MeJA-responsiveness | LuDMP-1、LuDMP-3、LuDMP-4、LuDMP-5、LuDMP-7、LuDMP-8、LuDMP-9、LuDMP-12、LuDMP-13、LuDMP-14、 LuDMP-16、LuDMP-17 |
GCN4_motif | TGAGTCA | 参与胚乳表达的顺式调控元件Cis-regulatory elements involved in endosperm expression | LuDMP-2、LuDMP-9、LuDMP-12、LuDMP-15 |
O2-site | GATGATGTGG | 参与玉米醇溶蛋白代谢调控的顺式调控元件Cis-acting regulatory element involved in zein metabolism regulation | LuDMP-1、LuDMP-2、LuDMP-3、LuDMP-4、LuDMP-5、LuDMP-8、LuDMP-10、LuDMP-12、LuDMP-16、LuDMP-17 |
MBS | CAACTG | 与干旱诱导相关的MYB结合位点MYB binding site involved in drought-inducibility | LuDMP-1、LuDMP-3、LuDMP-6、LuDMP-7、LuDMP-8、LuDMP-9、LuDMP-10、LuDMP-14 |
SARE | TTCGACCATCTT | 水杨酸响应元件Cis-acting element involved in salicylic acid responsiveness | LuDMP-1、LuDMP-7 |
TC-rich repeats | GTTTTCTTAC | 防御和应激响应元件Cis-acting element involved in defense and stress responsiveness | LuDMP-1、LuDMP-5、LuDMP-13、LuDMP-15、LuDMP-16 |
CAT-box | GCCACT | 与分生组织表达相关的顺式作用调控元件Cis-acting regulatory element related to meristem expression | LuDMP-3、LuDMP-5、LuDMP-10、LuDMP-12、LuDMP-13、LuDMP-15 |
P-box | CCTTTTG | 赤霉素响应元件Gibberellin-responsive element | LuDMP-6、LuDMP-10、LuDMP-13、LuDMP-14 |
TGA-element | AACGAC | 生长素响应元件Auxin-responsive element | LuDMP-6、LuDMP-12、LuDMP-14 |
TATC-box | TATCCCA | 赤霉素响应元件Cis-acting element involved in gibberellin-responsiveness | LuDMP-8、LuDMP-9、LuDMP-10、LuDMP-15 |
GC-motif | CCCCCG | 参与缺氧特异性诱导的增强剂样元件Enhancer-like element involved in anoxic specific inducibility | LuDMP-9、LuDMP-15、LuDMP-17 |
DRE | TACCGACAT | 脱水、低温及盐胁迫响应元件Cis-acting element involved in dehydration, low-temperature, salt stresses | LuDMP-3 |
RY-element | CATGCATG | 参与种子特异性调控的顺式调控元件Cis-regulatory elements involved in seed-specific regulation | LuDMP-2 |
GARE-motif | TCTGTTG | 赤霉素响应元件Gibberellin-responsive element | LuDMP-7、LuDMP-8、LuDMP-9、LuDMP-10、LuDMP-12 |
TCA-element | CCATTTTTT | 水杨酸响应元件Cis-acting element involved in salicylic acid responsiveness | LuDMP-2、LuDMP-5、LuDMP-10、LuDMP-14、LuDMP-16、LuDMP-17 |
LTR | CCGAAA | 低温响应元件Cis-acting element involved in low-temperature responsiveness | LuDMP-1、LuDMP-2、LuDMP-5、LuDMP-6、LuDMP-7、LuDMP-8、LuDMP-9、LuDMP-10、LuDMP-11、LuDMP-12、LuDMP-13、LuDMP-14 |
表5 亚麻LuDMP启动子顺式元件分析
Table 5 Analysis of cis-elements of flax LuDMP promoters
顺式元件 Cis-element | 典型序列 Typical sequence | 特性 Characteristic | 基因Gene |
---|---|---|---|
TATA-box | TATA | 转录开始的-30左右的核心启动子元件Core promoter element around -30 of transcript-ion start | LuDMP-1、LuDMP-2、LuDMP-4、LuDMP-5、LuDMP-6、LuDMP-7、LuDMP-8、LuDMP-9、LuDMP-10、LuDMP-11、LuDMP-12、LuDMP-13、LuDMP-14、 LuDMP-15、LuDMP-16、LuDMP-17 |
ARE | AAACCA | 厌氧诱导所必需的顺式调节元件Cis-acting regulatory element essential for the anaerobic induction | LuDMP-1、LuDMP-2、LuDMP-3、LuDMP-4、LuDMP-5、LuDMP-6、LuDMP-7、LuDMP-8、LuDMP-9、LuDMP-10、LuDMP-11、LuDMP-12、LuDMP-13、LuDMP-14、 LuDMP-15、LuDMP-16、LuDMP-17 |
Sp1 | GGGCGG | 光响应元件Light responsive element | LuDMP-1、LuDMP-3、LuDMP-6、LuDMP-8 |
ABRE | ACGTG | 参与脱落酸反应的顺式作用元件Cis-acting element involved in the abcisic acid responsiveness | LuDMP-1、LuDMP-4、LuDMP-5、LuDMP-6、LuDMP-7、LuDMP-8、LuDMP-9、LuDMP-10、LuDMP-11、LuDMP-13、LuDMP-16 |
TGACG-motif | TGACG | 参与甲基茉莉酸(MeJA)反应的顺式调节元件Cis-acting regulatory element involved in the MeJA-responsiveness | LuDMP-1、LuDMP-4、LuDMP-5、LuDMP-7、LuDMP-8、LuDMP-9、LuDMP-12、LuDMP-13、LuDMP-14、 LuDMP-15、LuDMP-16、LuDMP-17 |
CGTCA-motif | CGTCA | 参与甲基茉莉酸(MeJA)反应的顺式调节元件Cis-acting regulatory element involved in the MeJA-responsiveness | LuDMP-1、LuDMP-3、LuDMP-4、LuDMP-5、LuDMP-7、LuDMP-8、LuDMP-9、LuDMP-12、LuDMP-13、LuDMP-14、 LuDMP-16、LuDMP-17 |
GCN4_motif | TGAGTCA | 参与胚乳表达的顺式调控元件Cis-regulatory elements involved in endosperm expression | LuDMP-2、LuDMP-9、LuDMP-12、LuDMP-15 |
O2-site | GATGATGTGG | 参与玉米醇溶蛋白代谢调控的顺式调控元件Cis-acting regulatory element involved in zein metabolism regulation | LuDMP-1、LuDMP-2、LuDMP-3、LuDMP-4、LuDMP-5、LuDMP-8、LuDMP-10、LuDMP-12、LuDMP-16、LuDMP-17 |
MBS | CAACTG | 与干旱诱导相关的MYB结合位点MYB binding site involved in drought-inducibility | LuDMP-1、LuDMP-3、LuDMP-6、LuDMP-7、LuDMP-8、LuDMP-9、LuDMP-10、LuDMP-14 |
SARE | TTCGACCATCTT | 水杨酸响应元件Cis-acting element involved in salicylic acid responsiveness | LuDMP-1、LuDMP-7 |
TC-rich repeats | GTTTTCTTAC | 防御和应激响应元件Cis-acting element involved in defense and stress responsiveness | LuDMP-1、LuDMP-5、LuDMP-13、LuDMP-15、LuDMP-16 |
CAT-box | GCCACT | 与分生组织表达相关的顺式作用调控元件Cis-acting regulatory element related to meristem expression | LuDMP-3、LuDMP-5、LuDMP-10、LuDMP-12、LuDMP-13、LuDMP-15 |
P-box | CCTTTTG | 赤霉素响应元件Gibberellin-responsive element | LuDMP-6、LuDMP-10、LuDMP-13、LuDMP-14 |
TGA-element | AACGAC | 生长素响应元件Auxin-responsive element | LuDMP-6、LuDMP-12、LuDMP-14 |
TATC-box | TATCCCA | 赤霉素响应元件Cis-acting element involved in gibberellin-responsiveness | LuDMP-8、LuDMP-9、LuDMP-10、LuDMP-15 |
GC-motif | CCCCCG | 参与缺氧特异性诱导的增强剂样元件Enhancer-like element involved in anoxic specific inducibility | LuDMP-9、LuDMP-15、LuDMP-17 |
DRE | TACCGACAT | 脱水、低温及盐胁迫响应元件Cis-acting element involved in dehydration, low-temperature, salt stresses | LuDMP-3 |
RY-element | CATGCATG | 参与种子特异性调控的顺式调控元件Cis-regulatory elements involved in seed-specific regulation | LuDMP-2 |
GARE-motif | TCTGTTG | 赤霉素响应元件Gibberellin-responsive element | LuDMP-7、LuDMP-8、LuDMP-9、LuDMP-10、LuDMP-12 |
TCA-element | CCATTTTTT | 水杨酸响应元件Cis-acting element involved in salicylic acid responsiveness | LuDMP-2、LuDMP-5、LuDMP-10、LuDMP-14、LuDMP-16、LuDMP-17 |
LTR | CCGAAA | 低温响应元件Cis-acting element involved in low-temperature responsiveness | LuDMP-1、LuDMP-2、LuDMP-5、LuDMP-6、LuDMP-7、LuDMP-8、LuDMP-9、LuDMP-10、LuDMP-11、LuDMP-12、LuDMP-13、LuDMP-14 |
图8 LuDMP-1/7基因在种子及花药中的表达模式不同小写字母表示在0.05水平具有显著性差异。下同。Different small letters mean significant differences at the 0.05 level. The same below.
Fig.8 Expression patterns of LuDMP-1/7 in seeds and anther
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