园艺学报 ›› 2023, Vol. 50 ›› Issue (5): 947-958.doi: 10.16420/j.issn.0513-353x.2022-0205
郭静, 廖满余, 金燕, 马小川, 张芬, 卢晓鹏, 邓子牛, 盛玲()
收稿日期:
2022-06-09
修回日期:
2023-01-19
出版日期:
2023-05-25
发布日期:
2023-05-31
通讯作者:
*(E-mail:shengling0629@163.com)基金资助:
GUO Jing, LIAO Manyu, JIN Yan, MA Xiaochuan, ZHANG Fen, LU Xiaopeng, DENG Ziniu, SHENG Ling()
Received:
2022-06-09
Revised:
2023-01-19
Published:
2023-05-25
Online:
2023-05-31
摘要:
转录因子CsbHLH3调控柑橘果实柠檬酸代谢过程中的具体机制尚不清楚。以冰糖橙果实cDNA为材料,克隆了CsbHLH3。瞬时转化烟草亚细胞定位显示CsbHLH3蛋白定位在细胞核。异位表达CsbHLH3的阳性番茄果实中CsbHLH3的表达显著上升,同时柠檬酸含量显著降低。定量分析表明,CsbHLH3在番茄中异位表达引起了柠檬酸降解相关基因SlACO3b、SlGDH及液泡膜质子泵基因SlPH8的上调。相反,在冰糖橙叶片中瞬时抑制CsbHLH3会引起柠檬酸降解相关基因CsGABP、CsACO2、CsACLα1、CsACLα2及液泡膜质子泵基因CsPH8、CsVHP2、CsVHP3的下调表达,其柠檬酸含量显著高于空载对照。通过酵母单杂交和EMSA试验证实CsbHLH3均能够靶定PH8和GABP启动子调控其表达。上述结果表明转录因子CsbHLH3是通过调控柠檬酸代谢途径多个相关基因包括PH8和降解途径基因GABP的表达综合负调控柠檬酸的水平。
中图分类号:
郭静, 廖满余, 金燕, 马小川, 张芬, 卢晓鹏, 邓子牛, 盛玲. 柑橘转录因子CsbHLH3调控柠檬酸代谢的功能解析[J]. 园艺学报, 2023, 50(5): 947-958.
GUO Jing, LIAO Manyu, JIN Yan, MA Xiaochuan, ZHANG Fen, LU Xiaopeng, DENG Ziniu, SHENG Ling. Functional Analysis of Transcription Factor CsbHLH3 in Regulating Citric Acid Metabolism of Citrus Fruit[J]. Acta Horticulturae Sinica, 2023, 50(5): 947-958.
引物名称 | 正向引物(5′-3′) | 反向引物(5′-3′) |
---|---|---|
Primer name | Forward | Reverse |
p1300-CsbHLH3 | GAGAACACGGGGGACGAGCTCATGGCTGAGAAATTTTGGACAAAGG | GCCCTTGCTCACCATGGTACCTTTAGAAAGGGCAGCAAGTAGT |
RNAi-CsbHLH3 | AAAAAGCAGGCTAGCTGTCCTCTGGACGAC | GAAAGCTGGGTTGATTCGTATAGATTCATCATA |
Adapter-attB | GGGACAAGTTTGTACAAAAAAGCAGGCT; | GGGACCACTTTGTACAAGAAAGCTGGGT |
P35s-F | CCCACCCACGAGGAGCAT | |
CsbHLH3(qRT) | TTACTGGCATGCTTCAAACTTGA | TGGCCATCTCCCCAAATTAG |
Actin(qRT) | CCAAGCAGCATGAAGATCAA | ATCTGCTGGAAGGTGCTGAG |
CsCS1(qRT) | GGTGCCCCCAATATTAACAA | AGAGCTCGGTCCCATATCAA |
CsCS2(qRT) | ACTGGTGTATGGATGCGACA | TCTTCGTCTTGTGGCATTTG |
CsPEPC1(qRT) | GTGCGATCCCGTCTATCTGT | AAGGCTCAAGGCCACTTTTT |
CsPEPC2(qRT) | GGCATGCAAAACACTGGTTA | CATGTTCATTACGGCTTGGA |
CsPEPC3(qRT) | GAACAATGACGGACACAACG | TGGACTCGCTTCCAACTTCT |
CsACO1(qRT) | GGCAAGTCATTCACATGCGTT | TGAAGAAGTAGACCCCGGTTGA |
CsACO2(qRT) | GGCAATGATGAAGTGATGGCT | GTTGGAACATGGACCGTCTTT |
CsACO3(qRT) | TGCAGCAATGAGGTACAAGGC | TCACACCCAGAAGCATTGGAC |
CsACLα1(qRT) | GATACTGTTGGAGACTTGGG | GCTCTCTTACGACCATCAGG |
CsACLα2(qRT) | TACAGTGGAGCACCCAACGA | CCTTCAGGGCTTGGATTATG |
CsACLβ(qRT) | GAGGAGATAACAGAGACAAA | AACAAAGAGCCCATTCAGAT |
CsGABP(qRT) | GTTGAGTGTTCCATCTCATA | GAACATGGCCTGAATCAACA |
CsPH8(qRT) | CCGTGAAGGAATTGATTTGG | CCATGACAATGGATTCCACA |
CsVHP2(qRT) | TGAGCCACAGAATCAGAGAGAGAA | GCACCAACAATCAAACCAATAAAC |
CsVHP3(qRT) | CCCTGCACATACAACACAG | TGCTGACTCCTTTCCTTGCT |
Sl Actin(qRT) | CACCATTGGGTCTGAGCGAT | GGGCGACAACCTTGATCTTC |
Sl CS(qRT) | TTGGGGAACATCACAGTTG | TGATGGCACCTTTCCTGTT |
Sl PEPC1(qRT) | TGTGAACCTGAACCCGACT | GTCCCCTATTCGGGACTTC |
Sl PEPC2(qRT) | TATCACTACTTTAAATGTCTGC | TCAAGGATACATGATTCTTAAT |
Sl ACO3a(qRT) | GCCGCTTGCTTCAACTTCTAC | GACTCCACCTCGGCACAGA |
Sl ACO3b(qRT) | TGGTGCTTATTGCTCTAGTGGGTA | CAACACCGTATCTCCACCTCA |
Sl GAD2(qRT) | CTTTGATCTTCTCCGTCGTTG | ATATCGAGACGCGAAAGTCG |
Sl GAD3(qRT) | CAGGACGTTTCAATATAATC | CCTACGGAGGGTCTCAGAG |
Sl GDH(qRT) | AGCACGACAATGCACGAGGG | ATATTGGCGACCGCTGTCTTCC |
Sl PEPCK(qRT) | AGACGAAACCACTGAGGACGA | CATTCACAAACACCTTCTCCAA |
Sl GABA-T(qRT) | CAGCACAAGCTTGACGATGG | TGGTGATTACTGGTTAAGGG |
Sl SSADH(qRT) | TCTCCGCTGAGGAGGGTAAACG | ACAAGCAAGAGCAGGGCCAACC |
Sl PH8(qRT) | TCGGAAAGGCTGCACATCTT | CGCTTTTGAATCGGCCACAT |
表1 基因克隆和实时荧光定量分析所用引物序列
Table 1 Primers used for cloning and qRT-PCR analysis in this study
引物名称 | 正向引物(5′-3′) | 反向引物(5′-3′) |
---|---|---|
Primer name | Forward | Reverse |
p1300-CsbHLH3 | GAGAACACGGGGGACGAGCTCATGGCTGAGAAATTTTGGACAAAGG | GCCCTTGCTCACCATGGTACCTTTAGAAAGGGCAGCAAGTAGT |
RNAi-CsbHLH3 | AAAAAGCAGGCTAGCTGTCCTCTGGACGAC | GAAAGCTGGGTTGATTCGTATAGATTCATCATA |
Adapter-attB | GGGACAAGTTTGTACAAAAAAGCAGGCT; | GGGACCACTTTGTACAAGAAAGCTGGGT |
P35s-F | CCCACCCACGAGGAGCAT | |
CsbHLH3(qRT) | TTACTGGCATGCTTCAAACTTGA | TGGCCATCTCCCCAAATTAG |
Actin(qRT) | CCAAGCAGCATGAAGATCAA | ATCTGCTGGAAGGTGCTGAG |
CsCS1(qRT) | GGTGCCCCCAATATTAACAA | AGAGCTCGGTCCCATATCAA |
CsCS2(qRT) | ACTGGTGTATGGATGCGACA | TCTTCGTCTTGTGGCATTTG |
CsPEPC1(qRT) | GTGCGATCCCGTCTATCTGT | AAGGCTCAAGGCCACTTTTT |
CsPEPC2(qRT) | GGCATGCAAAACACTGGTTA | CATGTTCATTACGGCTTGGA |
CsPEPC3(qRT) | GAACAATGACGGACACAACG | TGGACTCGCTTCCAACTTCT |
CsACO1(qRT) | GGCAAGTCATTCACATGCGTT | TGAAGAAGTAGACCCCGGTTGA |
CsACO2(qRT) | GGCAATGATGAAGTGATGGCT | GTTGGAACATGGACCGTCTTT |
CsACO3(qRT) | TGCAGCAATGAGGTACAAGGC | TCACACCCAGAAGCATTGGAC |
CsACLα1(qRT) | GATACTGTTGGAGACTTGGG | GCTCTCTTACGACCATCAGG |
CsACLα2(qRT) | TACAGTGGAGCACCCAACGA | CCTTCAGGGCTTGGATTATG |
CsACLβ(qRT) | GAGGAGATAACAGAGACAAA | AACAAAGAGCCCATTCAGAT |
CsGABP(qRT) | GTTGAGTGTTCCATCTCATA | GAACATGGCCTGAATCAACA |
CsPH8(qRT) | CCGTGAAGGAATTGATTTGG | CCATGACAATGGATTCCACA |
CsVHP2(qRT) | TGAGCCACAGAATCAGAGAGAGAA | GCACCAACAATCAAACCAATAAAC |
CsVHP3(qRT) | CCCTGCACATACAACACAG | TGCTGACTCCTTTCCTTGCT |
Sl Actin(qRT) | CACCATTGGGTCTGAGCGAT | GGGCGACAACCTTGATCTTC |
Sl CS(qRT) | TTGGGGAACATCACAGTTG | TGATGGCACCTTTCCTGTT |
Sl PEPC1(qRT) | TGTGAACCTGAACCCGACT | GTCCCCTATTCGGGACTTC |
Sl PEPC2(qRT) | TATCACTACTTTAAATGTCTGC | TCAAGGATACATGATTCTTAAT |
Sl ACO3a(qRT) | GCCGCTTGCTTCAACTTCTAC | GACTCCACCTCGGCACAGA |
Sl ACO3b(qRT) | TGGTGCTTATTGCTCTAGTGGGTA | CAACACCGTATCTCCACCTCA |
Sl GAD2(qRT) | CTTTGATCTTCTCCGTCGTTG | ATATCGAGACGCGAAAGTCG |
Sl GAD3(qRT) | CAGGACGTTTCAATATAATC | CCTACGGAGGGTCTCAGAG |
Sl GDH(qRT) | AGCACGACAATGCACGAGGG | ATATTGGCGACCGCTGTCTTCC |
Sl PEPCK(qRT) | AGACGAAACCACTGAGGACGA | CATTCACAAACACCTTCTCCAA |
Sl GABA-T(qRT) | CAGCACAAGCTTGACGATGG | TGGTGATTACTGGTTAAGGG |
Sl SSADH(qRT) | TCTCCGCTGAGGAGGGTAAACG | ACAAGCAAGAGCAGGGCCAACC |
Sl PH8(qRT) | TCGGAAAGGCTGCACATCTT | CGCTTTTGAATCGGCCACAT |
图1 CsbHLH3的亚细胞定位分析载体骨架示意图(A)和CsbHLH3的亚细胞定位(B) 绿色荧光为CsbHLH3-eYFP融合蛋白:红色荧光为mCherry蛋白(Hy5)核定位标记。
Fig. 1 Schematic diagram of the vector backbone for subcellular localization analysis of CsbHLH3(A)and Subcellular localization of CsbHLH3(B) Green fluorescence is CsbHLH3-eYFP fusion protein;red fluorescence is the nuclear localization marker of mCherry protein(Hy5).
图2 转CsbHLH3基因的番茄(OE-20、OE-24)果实表型(A)及该基因表达量(B)和柠檬酸含量(C) DAF:花后天数;WT:野生型。不同字母表示不同材料在P < 0.05水平差异显著。
Fig. 2 Fruit phenotype(A),relative expression(B)and citrate content(C)of tomato(OE-20,OE-24)transgenic with CsbHLH3 gene DAF:Days After Flowering;WT:Wild Type. Different letters indicate that different materials have significant differences at P < 0.05 level.
图3 转CsbHLH3基因的番茄(OE-20、OE-24)果实中柠檬酸代谢相关基因的表达量 WT:野生型;合成基因:SlCS1、SlPEPC1、SlPEPC2;降解基因:SlACO3a、SlACO3b、SlGAD2、SlGAD3、SlGDH、SlPEPCK、SlGABA-T、SlSSADH;转运基因:SlPH8。不同字母表示不同材料在P < 0.05水平差异显著。
Fig. 3 Expression of citrate metabolism related genes in tomato(OE-20,OE-24)transgenic with CsbHLH3 gene WT:Wild type;Synthetic gene:SlCS1,SlPEPC1,SlPEPC2;Degradable gene:SlACO3a,SlACO3b,SlGAD2,SlGAD3,SlGDH,SlPEPCK,SlGABA-T,SlSSADH;Transport gene:SlPH8. Different letters indicate that different materials have significant differences at P < 0.05 level.
图4 抑制CsbHLH3表达的柑橘叶片中该基因表达量(A)及柠檬酸含量(B)
Fig. 4 The relative expression(A)and citrate content(B)of CsbHLH3 in citrus leaves were suppressed t-test,* α = 0.05.
图5 抑制CsbHLH3表达的柑橘叶片中柠檬酸代谢相关基因的表达定量
Fig. 5 Quantitative expression of citrate metabolism related genes in citrus leaves suppressing CsbHLH3 expression t-test,* α = 0.05.
图6 转录因子CsbHLH3与CsPH8和CsGABP的互作分析 A、B:EMSA分析;C、D:酵母单杂交分析。Positive:p53-pAbAi,阳性对照;Negative:pGADT7,阴性对照。
Fig. 6 Interaction analysis of transcription factor CsbHLH3 with CsPH8 and CsGABP A,B:EMSA analysis;C,D:Yeast one-hybrid analysis. Positive:p53-pAbAi as positive control;Negative:pGADT7 as negative control.
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