园艺学报 ›› 2022, Vol. 49 ›› Issue (4): 816-826.doi: 10.16420/j.issn.0513-353x.2021-0007
刘筱玮1, 夏斌1, 李子葳1, 杨宇佳1, 陈斌1, 周蕴薇2,**(), 何淼1,**(
)
收稿日期:
2021-11-29
修回日期:
2022-03-07
出版日期:
2022-04-25
发布日期:
2022-04-24
通讯作者:
周蕴薇,何淼
E-mail:dlzhyw@126.com;hm2017148@126.com
基金资助:
LIU Xiaowei1, XIA Bin1, LI Ziwei1, YANG Yujia1, CHEN Bin1, ZHOU Yunwei2,**(), HE Miao1,**(
)
Received:
2021-11-29
Revised:
2022-03-07
Online:
2022-04-25
Published:
2022-04-24
Contact:
ZHOU Yunwei,HE Miao
E-mail:dlzhyw@126.com;hm2017148@126.com
摘要:
从野菊(Chrysanthemum indicum)中克隆了cin-miR396a前体序列及其启动子,对前体序列进行比对与进化分析,对启动子顺式作用元件进行分析,并研究了cin-miR396a前体序列对拟南芥的遗传转化及对其耐盐性的影响。结果表明,cin-MIR396a长度为145 bp,含有完整的茎环结构,与拟南芥ath-MIR396a进化关系最近。启动子区具有光信号、茉莉酸甲酯、赤霉素、水杨酸、脱落酸等多种响应元件,推测cin-MIR396a参与多种逆境胁迫下的应答及调控。在拟南芥中过表达cin-miR396a前体可以提高盐胁迫下种子发芽率以及成苗期的耐盐性,cin-miR396a在拟南芥响应盐胁迫中发挥正调控作用。
中图分类号:
刘筱玮, 夏斌, 李子葳, 杨宇佳, 陈斌, 周蕴薇, 何淼. 拟南芥中过表达野菊miR396a基因增强其耐盐性[J]. 园艺学报, 2022, 49(4): 816-826.
LIU Xiaowei, XIA Bin, LI Ziwei, YANG Yujia, CHEN Bin, ZHOU Yunwei, HE Miao. Overexpression of Chrysanthemum indicum miR396a Gene in Arabidopsis Enhances Its Salt Tolerance[J]. Acta Horticulturae Sinica, 2022, 49(4): 816-826.
用途 Usage | 引物名称 Primer name | 引物序列(5′-3′) Sequence |
---|---|---|
前体基因克隆 Precursor gene cloning | MIR396a-F | TCTAGACGCTATTGTTTTACACACAAGTGTG |
MIR396a-R | CCCGGGCTGTGTTGATTCTTTCTCTATCTT | |
启动子克隆 Promoter cloning | P-miR396a-F | GCTCTCCATCTTGTTTCTCGTC |
转化植株PCR验证 PCR assay of the transgenic plants | P-miR396a-R | GCAGTTCAAGAAAGCTGTGGAAG |
35S-F | TCCCACTATCCTTCGCAAGAC | |
miR396a-R | CTGTGTTGATTCTTTCTCTATCTT | |
实时荧光定量PCR | athU6-F | CGATAAAATTGGAACGATACAGA |
Fluorescent quantitative real-time | athU6-R | ATTTGGACCATTTCTCGATTTGT |
PCR(qRT-PCR) | cin-miR396aF | TTCCACAGCTTTCTTGAACTG |
表1 本研究中所用引物及序列
Table 1 Primers and sequences used in this study
用途 Usage | 引物名称 Primer name | 引物序列(5′-3′) Sequence |
---|---|---|
前体基因克隆 Precursor gene cloning | MIR396a-F | TCTAGACGCTATTGTTTTACACACAAGTGTG |
MIR396a-R | CCCGGGCTGTGTTGATTCTTTCTCTATCTT | |
启动子克隆 Promoter cloning | P-miR396a-F | GCTCTCCATCTTGTTTCTCGTC |
转化植株PCR验证 PCR assay of the transgenic plants | P-miR396a-R | GCAGTTCAAGAAAGCTGTGGAAG |
35S-F | TCCCACTATCCTTCGCAAGAC | |
miR396a-R | CTGTGTTGATTCTTTCTCTATCTT | |
实时荧光定量PCR | athU6-F | CGATAAAATTGGAACGATACAGA |
Fluorescent quantitative real-time | athU6-R | ATTTGGACCATTTCTCGATTTGT |
PCR(qRT-PCR) | cin-miR396aF | TTCCACAGCTTTCTTGAACTG |
图3 cin-MIR396a系统进化分析 ath:拟南芥;bdi:二穗短柄草;fve:野草莓;gma:大豆;mtr:蒺藜状苜蓿;nta:烟草;osa:水稻;sly:番茄;stu:马铃薯;sbi:高粱;tcc:可可树;vvi:葡萄;zma:玉米。
Fig. 3 Phylogenetic relationship analysis of cin-MIR396a ath:Arabidopsis thaliana;bdi:Brachypodium distachyon;fve:Fragaria vesca;gma:Glycie max;mtr:Medicago truncatula;nta:Nicotiana tabacum;osa:Oryza sativa;sly:Solanum lycopersicum;stu:Solanum tuberosum;sbi:Sorghum bicolor;tcc:Theobroma cacao;vvi:Vitis vinifera;zma:Zea mays.
顺式作 用元件 cis-acting elements | 功能 Function | 野菊 Chrysanthemum indicum | 拟南芥 Arabidopsis thaliana | 水稻 Oryza sativa |
---|---|---|---|---|
A-box | 顺式作用调节元件cis-Acting regulatory element | + | - | - |
ABRE | 脱落酸应答元件cis-Acting element involved in ABA responsiveness | + | + | + |
ACE | 光应答元件cis-Acting element involved in light responsiveness | - | - | + |
AE-box | 部分光应答元件Part of a light responsive element | - | - | + |
ARE | 厌氧诱导应答元件cis-Acting element essential for the anaerobic induction | + | + | - |
Box 4 | 部分光应答元件Part of a light responsive element | + | + | - |
BoxⅡ | 部分光应答元件Part of a light responsive element | + | - | - |
CAAT-box | 增强子元件cis-Acting element playing an enhancing role | + | + | + |
CAT-box | 分生组织表达有关的应答元件cis-Acting element related to meristem expression | - | - | + |
CCAAT-box | MYBHv1结合位点MYBHv1 binding site | + | - | - |
CGTCA-motif | 茉莉酸甲酯应答元件cis-Acting element involved in the MeJA-responsiveness | + | + | + |
circadian | 昼夜节律应答元件cis-Acting element involved in circadian control | - | + | - |
ERE | 乙烯应答元件Ethylene responsive element | + | + | - |
GA-motif | 部分光应答元件Part of a light responsive element | + | - | - |
GARE-motif | 赤霉素应答元件Gibberellin-responsive element | - | - | + |
GATA-motif | 部分光应答元件Part of a light responsive element | + | - | + |
G-Box | 光应答元件cis-Acting element involved in light responsiveness | + | + | + |
GCN4_motif | 胚乳表达的应答元件cis-Acting element involved in endosperm expression | + | - | - |
I-box | 部分光应答元件Part of a light responsive element | + | - | - |
MBS | 干旱诱导中MYB 结合位点MYB binding site involved in drought-inducibility | + | - | - |
MYB | MYB结合位点MYB binding site | + | + | + |
MYC | MYC结合位点MYC binding site | + | + | + |
O2-site | 玉米醇溶蛋白代谢的应答元件cis-Acting element involved in zein metabolism regulation | + | - | + |
P-box | 赤霉素应答元件cis-Acting element involved in gibberellin-responsive | + | + | - |
TATA-box | 核心启动子元件Core promoter elements | + | + | + |
TCA-element | 水杨酸应答元件cis-Acting element involved in salicylic acid responsiveness | + | - | - |
TGACG-motif | 茉莉酸甲酯应答元件cis-Acting element involved in the MeJA-responsiveness | - | + | + |
表2 cin-MIR396a启动子区顺时作用元件比较
Table 2 Comparison of cis-acting elements in promoters of cin-MIR396a
顺式作 用元件 cis-acting elements | 功能 Function | 野菊 Chrysanthemum indicum | 拟南芥 Arabidopsis thaliana | 水稻 Oryza sativa |
---|---|---|---|---|
A-box | 顺式作用调节元件cis-Acting regulatory element | + | - | - |
ABRE | 脱落酸应答元件cis-Acting element involved in ABA responsiveness | + | + | + |
ACE | 光应答元件cis-Acting element involved in light responsiveness | - | - | + |
AE-box | 部分光应答元件Part of a light responsive element | - | - | + |
ARE | 厌氧诱导应答元件cis-Acting element essential for the anaerobic induction | + | + | - |
Box 4 | 部分光应答元件Part of a light responsive element | + | + | - |
BoxⅡ | 部分光应答元件Part of a light responsive element | + | - | - |
CAAT-box | 增强子元件cis-Acting element playing an enhancing role | + | + | + |
CAT-box | 分生组织表达有关的应答元件cis-Acting element related to meristem expression | - | - | + |
CCAAT-box | MYBHv1结合位点MYBHv1 binding site | + | - | - |
CGTCA-motif | 茉莉酸甲酯应答元件cis-Acting element involved in the MeJA-responsiveness | + | + | + |
circadian | 昼夜节律应答元件cis-Acting element involved in circadian control | - | + | - |
ERE | 乙烯应答元件Ethylene responsive element | + | + | - |
GA-motif | 部分光应答元件Part of a light responsive element | + | - | - |
GARE-motif | 赤霉素应答元件Gibberellin-responsive element | - | - | + |
GATA-motif | 部分光应答元件Part of a light responsive element | + | - | + |
G-Box | 光应答元件cis-Acting element involved in light responsiveness | + | + | + |
GCN4_motif | 胚乳表达的应答元件cis-Acting element involved in endosperm expression | + | - | - |
I-box | 部分光应答元件Part of a light responsive element | + | - | - |
MBS | 干旱诱导中MYB 结合位点MYB binding site involved in drought-inducibility | + | - | - |
MYB | MYB结合位点MYB binding site | + | + | + |
MYC | MYC结合位点MYC binding site | + | + | + |
O2-site | 玉米醇溶蛋白代谢的应答元件cis-Acting element involved in zein metabolism regulation | + | - | + |
P-box | 赤霉素应答元件cis-Acting element involved in gibberellin-responsive | + | + | - |
TATA-box | 核心启动子元件Core promoter elements | + | + | + |
TCA-element | 水杨酸应答元件cis-Acting element involved in salicylic acid responsiveness | + | - | - |
TGACG-motif | 茉莉酸甲酯应答元件cis-Acting element involved in the MeJA-responsiveness | - | + | + |
图4 过表达cin-MIR396a拟南芥抗性筛选(A)与PCR验证(B) M:Marker 2000;1:重组质粒;2:野生型拟南芥;3:阴性对照;4 ~ 10:转化植株。
Fig. 4 Resistance screening(A)and PCR verification(B)of Arabidopsis thaliana overexpressing cin-MIR396a M:Marker 2000;1:Recombinant plasmid;2:Wild-type Arabidopsis;3:Negative control;4-10:Transformed plant.
图5 过表达拟南芥株系(OE1 ~ OE7)中miR396a的表达分析 * 表明在α = 0. 05 水平差异显著;** 表明在α = 0.01 水平差异显著。WT:野生型;OE1 ~ OE7:过表达株系。下同。
Fig. 5 Expression analysis of miR396a in transgenic Arabidopsis(OE1-OE7) * indicates that the difference is significant at the α = 0.05 level;** indicates that the difference is significant at the α = 0.01 level. WT:Wild type;OE1-OE7:Overexpression strains. The same below.
图6 过表达cin-MIR396a拟南芥(OE4、OE6、OE7)在不同盐浓度处理下的发芽情况
Fig. 6 Germination of Arabidopsis(OE4,OE6,OE7)overexpressing cin-MIR396a under different salt concentrations
图7 过表达cin-MIR396a拟南芥(OE4、OE6、OE7)在300 mol · L-1 NaCl处理下的形态特征
Fig. 7 Morphological characteristics of overexpressing cin-MIR396a Arabidopsis(OE4,OE6,OE7)under 300 mol · L-1 NaCl treatment
图8 300 mol · L-1 NaCl胁迫8 d对转基因拟南芥(OE4、OE6、OE7)MDA含量(A)和相对电导率(B)的影响
Fig. 8 Effects of 300 mol · L-1 NaCl stress treatment for 8 d on MDA content(A)and relative conductivity(B)of transgenic Arabidopsis(OE4,OE6,OE7)
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