植物启动子研究进展

doi:10.16420/j.issn.0513-353x.2024-0408

摘要/Abstract

摘要： 探讨了不同类型启动子的结构特性及其在基因功能研究、代谢工程和基因编辑（CRISPR/ Cas9）中的作用，并总结了启动子研究的内容及方法，以期为深入理解和研究植物启动子的结构特性和调控机制提供帮助。

Abstract: In this paper， the structural characteristics of different types of promoters and their roles in gene function research，metabolic engineering，and gene editing（CRISPR/Cas9）were discusses，and summarized the content and methods for promoter identification，aiming to provide insights for a deeper understanding and investigation of the structural characteristics and regulatory mechanisms of plant promoters.

Key words: promoter, genetic engineering, metabolic engineering, CRISPR/Cas9

中图分类号:

Q 78

张志愿, 王丹, 惠竹梅, 王现行. 植物启动子研究进展[J]. 园艺学报, 2025, 52(4): 1065-1092.

ZHANG Zhiyuan, WANG Dan, XI Zhumei, WANG Xianhang. Advances in Plant Promoter Research[J]. Acta Horticulturae Sinica, 2025, 52(4): 1065-1092.

参考文献

[1]	Ahammed G J，Li C X，Li X，Liu A，Chen S，Zhou J. 2021. Overexpression of tomato RING E3 ubiquitin ligase gene SlRING1 confers cadmium tolerance by attenuating cadmium accumulation and oxidative stress. Physiologia plantarum，173 (1)：449–459.
[2]	Amack S C，Antunes M S. 2020. CaMV35S promoter-A plant biology and biotechnology workhorse in the era of synthetic biology. Current Plant Biology，24：100179.
[3]	Archer L，Mondal H A，Behera S，Twayana M，Patel M，Louis J，Nalam V J，Keereetaweep J，Chowdhury Z，Shah J. 2023. Interplay between MYZUS PERSICAE-INDUCED LIPASE 1 and OPDA signaling in limiting green peach aphid infestation on Arabidopsis thaliana. Journal of Experimental Botany，74 (21)：6860–6873.
[4]	Atkinson R G，Bolitho K M，Wright M A，Iturriagagoitia-Bueno T，Reid S J，Ross G S. 1998. Apple ACC-oxidase and polygalacturonase：ripening-specific gene expression and promoter analysis in transgenic tomato. Plant Molecular Biology，38：449–460.
[5]	Bai J，Wang X，Wu H，Ling F，Zhao Y，Lin Y，Wang R. 2020. Comprehensive construction strategy of bidirectional green tissue-specific synthetic promoters. Plant Biotechnology Journal，18 (3)：668–678.
[6]	Bhullar S，Datta S，Advani S，Chakravarthy S，Gautam T，Pental D，Burma P K. 2007. Functional analysis of cauliflower mosaic virus 35S promoter：re-evaluation of the role of subdomains B5，B4 and B2 in promoter activity. Plant Biotechnology Journal，5 (6)：696–708.
[7]	Boruah R R，Konwar T，Nath P K，Acharjee S，Sarmah B K. 2023. Activity of Arabidopsis Rubisco small subunit promoter in various tissues of chickpea. 3 Biotech，13 (3)：89.
[8]	Brown R L，Kazan K，McGrath K C，Maclean D J，Manners J M. 2003. A role for the GCC-box in jasmonate-mediated activation of the PDF1.2 gene of Arabidopsis. Plant Physiology，132 (2)：1020–1032.
[9]	Butler J E，Kadonaga J T. 2002. The RNA polymerase II core promoter：a key component in the regulation of gene expression. Genes & Development，16 (20)：2583–2592.
[10]	Büttner M，Singh K B. 1997. Arabidopsis thaliana ethylene-responsive element binding protein（AtEBP），an ethylene-inducible，GCC box DNA-binding protein interacts with an ocs element binding protein. Proceedings of the National Academy of Sciences，94 (11)：5961–5966.
[11]	Cai C，Wang W，Ye S，Zhang Z，Ding W，Xiang M，Wu C，Zhu Q. 2020. Overexpression of a novel Arabidopsis gene SUPA leads to various morphological and abiotic stress tolerance alternations in Arabidopsis and Poplar. Frontiers in Plant Science，11：560985.
[12]	Cai M，Wei J，Li X，Xu C，Wang S. 2007. A rice promoter containing both novel positive and negative cis-elements for regulation of green tissue-specific gene expression in transgenic plants. Plant Biotechnology Journal，5 (5)：664–674.
[13]	Chen J，Ren C，Wang J，Xian B，Wang R，Wu Q，Pei J. 2023a. pCtFSG2，a flower-specific promoter with suitable promoter activity in safflower. Journal of Plant Biochemistry and Biotechnology，32 (3)：478–486.
[14]	Chen K，Liu W，Li X，Li H. 2021a. Overexpression of GmGASA32 promoted soybean height by interacting with GmCDC25. Plant Signaling & Behavior，16 (2)：1855017.
[15]	Chen L，Jiang B，Wu C，Sun S，Hou W，Han T. 2015. The characterization of GmTIP，a root-specific gene from soybean，and the expression analysis of its promoter. Plant Cell，Tissue and Organ Culture，121：259–274.
[16]	Chen L，Li Y，Wang Y，Li W，Feng X，Zhao L. 2021b. Use of high resolution spatiotemporal gene expression data to uncover novel tissue-specific promoters in tomato. Agriculture，11 (12)：1195.
[17]	Chen L，Su P，Yang G，H G，Gao C. 2023b. PSG076 promoter shows late pollen-specific activity in wheat（Triticum aestivum L.）. Plant Molecular Biology Reporter，41 (4)：690–698.
[18]	Chen Z，Zhou L，Jiang P，Lu R，Halford N G，Liu C. 2021c. Genome-wide identification of sucrose nonfermenting-1-related protein kinase（SnRK） genes in barley and RNA-seq analyses of their expression in response to abscisic acid treatment. BMC Genomics，22 (1)：300.
[19]	Choi H I，Hong J H，Ha J O，Kang J Y，Kim S Y. 2000. ABFs，a family of ABA-responsive element binding factors. Journal of Biological Chemistry，275 (3)：1723–1730.
[20]	Chow C N，Lee T Y，Hung Y C，Li G Z，Tseng K C，Liu Y H，Kuo P L，Zheng H Q，Chang W C. 2019. PlantPAN3.0：a new and updated resource for reconstructing transcriptional regulatory networks from ChIP-seq experiments in plants. Nucleic Acids Research，47 (D1)：D1155–D1163.
[21]	Christensen A H，Sharrock R A，Quail P H. 1992. Maize polyubiquitin genes：structure，thermal perturbation of expression and transcript splicing，and promoter activity following transfer to protoplasts by electroporation. Plant Molecular Biology，18：675–689.
[22]	Chung M Y，Naing A H，Vrebalov J，Shanmugam A，Lee D J，Park I H，Kim C K，Giovannon J. 2020. The use of SlAdh2 promoter as a novel fruit-specific promoter in transgenic tomato. Journal of Plant Biotechnology，47 (2)：172–178.
[23]	Cobb R E，Sun N，Zhao H. 2013. Directed evolution as a powerful synthetic biology tool. Methods，60 (1)：81–90.
[24]	Colebrook E H，Thomas S G，Phillips A L，Hedden P. 2014. The role of gibberellin signalling in plant responses to abiotic stress. Journal of Experimental Biology，217 (1)：67-75.
[25]	Dasgupta K，Hotton S，Belknap W，Syed Y，Dardick C，Thilmony R，Thomson J G. 2020. Isolation of novel citrus and plum fruit promoters and their functional characterization for fruit biotechnology. BMC Biotechnology，20：1–15.
[26]	de Ruijter N C A，Verhees J，Van Leeuwen W，Van Der Krol A R. 2003. Evaluation and comparison of the GUS，LUC and GFP reporter system for gene expression studies in plants. Plant Biology，5 (2)：103–115.
[27]	de Silva W S I，Perera M M N，Perera K L N S，Wickramasuriya A M，Jayasekera G A U. 2017. In silico analysis of osr40c1 promoter sequence isolated from Indica variety Pokkali. Rice science，24 (4)：228–234.
[28]	Deb D，Dey N. 2019. Synthetic salicylic acid inducible recombinant promoter for translational research. Journal of Biotechnology，297：9–18.
[29]	Dong G，Fan M，Wang H，Leng Y，Sun J，Huang J，Zhang H，Yan J. 2023. Functional characterization of TkSRPP promoter in response to hormones and wounding stress in transgenic tobacco. Plants，12 (2)：252.
[30]	Dong H，Liu L，Fan X，Asghar S，Li Y，Wang Y，Xu X，Wu T，Zhang X，Qiu C，Han Z，Li W. 2019. The artificial promoter rMdAG2I confers flower-specific activity in Malus. International Journal of Molecular Sciences，20 (18)：4551.
[31]	Elliott A R，Campbell J A，Dugdale B，Brettell R I S，Grof C P L. 1999. Green-fluorescent protein facilitates rapid in vivo detection of genetically transformed plant cells. Plant Cell Reports，18：707–714.
[32]	Fan J，Jiang F，Sun H，He T，Liu Y，Jiao G，Ahmad B，Bokhari S A M，Chen Q，Wen Z. 2023. Expression analysis of trihelix transcription factor family in strawberries and functional characterization of FvTrihelix6. Horticulturae，9 (6)：633.
[33]	Fang R X，Nagy F，Sivasubramaniam S，Chua N H. 1989. Multiple cis regulatory elements for maximal expression of the cauliflower mosaic virus 35S promoter in transgenic plants. The Plant Cell，1 (1)：141–150.
[34]	Feng C，Su H，Bai H，Wang R，Liu Y，Guo X，Liu C，Zhang J，Yuan J，Birchler J A，Han F. 2018. High-efficiency genome editing using a dmc1 promoter-controlled CRISPR/Cas9 system in maize. Plant Biotechnology Journal，16 (11)：1848–1857.
[35]	Feng Z，Liu N，Zhang G，Bu Y，Wang B，Gong Y. 2022. Promoter of vegetable soybean GmTIP1;6 responds to diverse abiotic stresses and hormone signals in transgenic Arabidopsis. International Journal of Molecular Sciences，23 (20)：12684.
[36]	Fernandez-Moreno J P，Yaschenko A E，Neubauer M，Marchi A J，Zhao C，Ascencio-Ibanez J T，Alonso J M，Stepanova A N. 2024. A rapid and scalable approach to build synthetic repetitive hormone-responsive promoters. Plant Biotechnology Journal，22 (7)：1942–1956
[37]	Field A，Adelman K. 2020. Evaluating enhancer function and transcription. Annual Review of Biochemistry，89：213–234.
[38]	Fitzgerald H A，Chern M S，Navarre R，Ronald P C. 2004. Overexpression of (At)NPR1 in rice leads to a BTH-and environment-induced lesion-mimic/cell death phenotype. Molecular Plant-Microbe Interactions，17 (2)：140–151.
[39]	Freitas E O，Melo B P，Lourenço-Tessutti I T，Arraes F B，Amorim R M，Lisei-de-Sá M E，Costa J A，Leite A G B，Faheem M，Ferreira M A，Morgante C V，Fontes E P B，Grossi-de-Sa M F. 2019. Identification and characterization of the GmRD26 soybean promoter in response to abiotic stresses：potential tool for biotechnological application. BMC Biotechnology，19：1–14.
[40]	Gatz C，Lenk I. 1998. Promoters that respond to chemical inducers. Trends in Plant Science，3 (9)：352–358.
[41]	Geem K R，Kim D H，Lee D W，Kwon Y，Lee J，Kim J H，Hwang I. 2019. Jasmonic acid-inducible TSA 1 facilitates ER body formation. The Plant Journal，97 (2)：267–280.
[42]	Gong P，Wei R，Li Y，Wang R，Tang Y，Wang L，Zhu H，Wang Y，Zhang C. 2019. Molecular cloning and functional characterization of a seed-specific VvβVPE gene promoter from Vitis vinifera. Planta，250：657–665.
[43]	Gong Q，Wang Y，Jin Z，Hong Y，Liu Y. 2022. Transcriptional and post-transcriptional regulation of RNAi-related gene expression during plant-virus interactions. Stress Biology，2 (1)：33.
[44]	He Y，Zhang T，Sun H，Zhan H，Zhao Y. 2020. A reporter for noninvasively monitoring gene expression and plant transformation. Horticulture Research，7 (1)：152.
[45]	He Z M，Jiang X L，Qi Y，Luo D Q. 2008. Assessment of the utility of the tomato fruit-specific E8 promoter for driving vaccine antigen expression. Genetica，133：207–214.
[46]	Hua L，Li M，Sun X，Wang J，Li Z，Xu Y，Hu S，Chen H. 2012. A dual color fluorescent reporter system for the real time detection of promoter activity. Biotechnology Letters，34：823–830.
[47]	Huang L Y，Zhang F，Qiao Q I N，Wang W S，Zhang T，Fu B Y. 2015. Identification and validation of root-specific promoters in rice. Journal of Integrative Agriculture，14 (1)：1–10.
[48]	Jakočiūnas T，Pedersen L E，Lis A V，Jensen M K，Keasling J D. 2018. CasPER，a method for directed evolution in genomic contexts using mutagenesis and CRISPR/Cas9. Metabolic Engineering，48：288–296.
[49]	Jameel A，Noman M，Liu W，Ahmad N，Wang F，Li X，Li H. 2020. Tinkering cis motifs jigsaw puzzle led to root-specific drought-inducible novel synthetic promoters. International Journal of Molecular Sciences，21 (4)：1357.
[50]	Jefferson R A，Kavanagh T A，Bevan M W. 1987. GUS fusions：beta-glucuronidase as a sensitive and versatile gene fusion marker in higher plants. The EMBO Journal，6 (13)：3901–3907.
[51]	Jiang Y，Yu D. 2015. WRKY transcription factors：links between phytohormones and plant processes. Science China. Life Sciences，58 (5)：501.
[52]	Jin B，Sheng Z，Muhammad I，Chen J，Yang H. 2019. Cloning and functional analysis of the promoter of a stress-inducible gene（Zmap）in maize. PLoS ONE，14 (2)：e0211941.
[53]	Jores T，Tonnies J，Wrightsman T，Buckler E S，Cuperus J T，Fields S，Queitsch C. 2021. Synthetic promoter designs enabled by a comprehensive analysis of plant core promoters. Nature Plants，7 (6)：842–855.
[54]	Jupin I，Chua N H. 1996. Activation of the CaMV as-1 cis-element by salicylic acid：differential DNA-binding of a factor related to TGA1a. The EMBO Journal，15 (20)：5679–5689.
[55]	Kagaya Y，Hobo T，Murata M，Ban A，Hattori T. 2002. Abscisic acid-induced transcription is mediated by phosphorylation of an abscisic acid response element binding factor，TRAB1. The Plant Cell，14 (12)：3177–3189.
[56]	Kesanakurti D，Kolattukudy P E，Kirti P B. 2012. Fruit-specific overexpression of wound-induced tap1 under E8 promoter in tomato confers resistance to fungal pathogens at ripening stage. Physiologia Plantarum，146 (2)：136–148.
[57]	Khadanga B，Chanwala J，Sandeep I S，Dey N. 2021. Synthetic promoters from strawberry vein banding virus（SVBV）and dahlia mosaic virus （DaMV）. Molecular Biotechnology，63 (9)：792–806.
[58]	Kim A Y，Kim H M，Ma S H，Park S Y，Dat M T，Jang G，Joung Y H. 2019. The promoter of tomato HISTIDINE DECARBOXYLASE A is fruit-specific，and its expression is stably maintained in fruits during ripening. Plant Biotechnology Reports，13：43–50.
[59]	Kim H M，Park S H，Park S Y，Ma S H，Do J H，Kim A Y，Jeon M J，Shim J S，Joung Y H. 2022. Identification of essential element determining fruit-specific transcriptional activity in the tomato HISTIDINE DECARBOXYLASE A gene promoter. Plant Cell Reports，41 (8)：1721–1731.
[60]	Kneissl M L，Deikman J. 1996. The tomato E8 gene influences ethylene biosynthesis in fruit but not in flowers. Plant Physiology，112 (2)：537–547.
[61]	Kobayashi S，Goto-Yamamoto N，Hirochika H. 2004. Retrotransposon-induced mutations in grape skin color. Science，304 (5673)：982.
[62]	Konishi M，Donner T J，Scarpella E，Yanagisawa S. 2015. MONOPTEROS directly activates the auxin-inducible promoter of the Dof5.8 transcription factor gene in Arabidopsis thaliana leaf provascular cells. Journal of Experimental Botany，66 (1)：283–291.
[63]	Koo J，Kim Y，Kim J，Yeom M，Lee I C，Nam H G. 2007. A GUS/luciferase fusion reporter for plant gene trapping and for assay of promoter activity with luciferin-dependent control of the reporter protein stability. Plant and Cell Physiology，48 (8)：1121–1131.
[64]	Köster J，Thurow C，Kruse K，Meier A，Iven T，Feussner I，Gatz C. 2012. Xenobiotic-and jasmonic acid-inducible signal transduction pathways have become interdependent at the Arabidopsis CYP81D11 promoter. Plant Physiology，159 (1)：391–402.
[65]	Krogan N T，Marcos D，Weiner A I，Berleth T. 2016. The auxin response factor MONOPTEROS controls meristem function and organogenesis in both the shoot and root through the direct regulation of PIN genes. New Phytologist，212 (1)：42–50.
[66]	Kummari D，Palakolanu S R，Kishor P K，Bhatnagar-Mathur P，Singam P，Vadez V，Sharma K K. 2020. An update and perspectives on the use of promoters in plant genetic engineering. Journal of Biosciences，45：1–24.
[67]	Lee D K，Jung H，Jang G，Jeong J S，Kim Y S，Ha S H，Choi Y D，Kim J K. 2016. Overexpression of the OsERF71 transcription factor alters rice root structure and drought resistance. Plant Physiology，172 (1)：575–588.
[68]	Lee M，Dominguez-Ferreras A，Kaliyadasa E，Huang W J，Antony E，Stevenson T，Lehmann S，Schäfer P，Knight M R，Ntoukakis V，Knight H. 2021. Mediator subunits MED16，MED14，and MED2 are required for activation of ABRE-dependent transcription in Arabidopsis. Frontiers in Plant Science，12：649720.
[69]	Lescot M，Déhais P，Thijs G，Marchal K，Moreau Y，Van de Peer Y，Rouzé P，Rombauts S. 2002. PlantCARE，a database of plant cis-acting regulatory elements and a portal to tools for in silico analysis of promoter sequences. Nucleic Acids Research，30 (1)：325–327.
[70]	Li C，Li W，Zhou Z，Chen H，Xie C，Lin Y. 2020a. A new rice breeding method：CRISPR/Cas9 system editing of the Xa13 promoter to cultivate transgene-free bacterial blight-resistant rice. Plant Biotechnology Journal，18 (2)：313.
[71]	Li R，Zhu F，Duan D. 2020b. Function analysis and stress-mediated cis-element identification in the promoter region of VqMYB15. Plant Signaling & Behavior，15 (7)：1773664.
[72]	Li X，Niu G，Fan Y，Liu W，Wu Q，Yu C，Wang J，Xiao Y，Hou L，Jin D，Chen S，Hu R，Yang Y，Pei Y. 2023. Synthetic dual hormone-responsive promoters enable engineering of plants with broad-spectrum resistance. Plant Communications，4 (4)：100596.
[73]	Li Y，Yu L，Wang Q，Zhao X，Li X，Qi B. 2021. Analysis of the promoter of Emb5 from Zea mays identifies a region of 523 bp responsible for its embryo-specific activity. Plant Molecular Biology Reporter，39：288–300.
[74]	Liao D，Chen X，Chen A，Wang H，Liu J，Liu J，Gu M，Sun S，Xu G. 2015. The characterization of six auxin-induced tomato GH3 genes uncovers a member，SlGH3.4，strongly responsive to arbuscular mycorrhizal symbiosis. Plant and Cell Physiology，56 (4)：674–687.
[75]	Lim C J，Lee H Y，Kim W B，Lee B S，Kim J，Ahmad R，Kim H A，Yi S Y，Hur C G，Kwon S Y. 2012. Screening of tissue-specific genes and promoters in tomato by comparing genome wide expression profiles of Arabidopsis orthologues. Molecules and Cells，34：53–59.
[76]	Lim H，Hwang H，Kim T，Kim S，Chung H，Lee D，Kim S，Park S，Cho W，Ji H，Lee G. 2021. Transcriptomic analysis of rice plants overexpressing PsGAPDH in response to salinity stress. Genes，12 (5)：641.
[77]	Lin M，Yan J，Ali M M，Wang S，Tian S，Chen F，Lin Z. 2022. Isolation and functional characterization of a green-tissue promoter in japonica rice （Oryza sativa subsp. Japonica）. Biology，11 (8)：1092.
[78]	Liu B，Zhang B，Yang Z，Liu Y，Yang S，Shi Y，Jiang C，Qin F. 2021a. Manipulating ZmEXPA4 expression ameliorates the drought-induced prolonged anthesis and silking interval in maize. The Plant Cell，33 (6)：2058–2071.
[79]	Liu L，Gallagher J，Arevalo E D，Chen R，Skopelitis T，Wu Q，Bartlett M，Jackson D. 2021b. Enhancing grain-yield-related traits by CRISPR-Cas9 promoter editing of maize CLE genes. Nature Plants，7 (3)：287–294.
[80]	Liu S，Liu C，Wang X，Chen H. 2021c. Seed-specific activity of the Arabidopsis β-glucosidase 19 promoter in transgenic Arabidopsis and tobacco. Plant Cell Reports，40：213–221.
[81]	Liu X，Ma X，Wang H，Li S，Yang W，Nugroho R D，Luo L，Zhou X，Tang C，Fan Y，Zhao Q，Zhang J，Chen R. 2021d. Metabolic engineering of astaxanthin-rich maize and its use in the production of biofortified eggs. Plant Biotechnology Journal，19 (9)：1812–1823.
[82]	Liu Y G，Chen Y. 2007. High-efficiency thermal asymmetric interlaced PCR for amplification of unknown flanking sequences. Biotechniques，43 (5)：649–656.
[83]	Liu Z B，Ulmasov T，Shi X，Hagen G，Guilfoyle T J. 1994. Soybean GH3 promoter contains multiple auxin-inducible elements. The Plant Cell，6 (5)：645–657.
[84]	Ly L K，Bui T P，Le A V T，Nguyen P V，Ong P X，Pham N B，Zhang Z J，Do P T，Chu H H. 2022. Enhancing plant growth and biomass production by overexpression of GA20ox gene under control of a root preferential promoter. Transgenic Research，31 (1)：73–85.
[85]	Maiti S，Patro S，Pal A，Dey N. 2015. Identification of a novel salicylic acid inducible endogenous plant promoter regulating expression of CYR1，a CC-NB-LRR type candidate disease resistance gene in Vigna mungo. Plant Cell，Tissue and Organ Culture（PCTOC），120：489–505.
[86]	McElroy D，Blowers A D，Jenes B，Wu R. 1991. Construction of expression vectors based on the rice actin 1（Act1）5’region for use in monocot transformation. Molecular and General Genetics MGG，231：150–160.
[87]	Mohan C，Jayanarayanan A N，Narayanan S. 2017. Construction of a novel synthetic root-specific promoter and its characterization in transgenic tobacco plants. 3 Biotech，7：1–9.
[88]	Molla K A，Karmakar S，Chanda P K，Ghosh S，Sarkar S N，Datta S K，Datta K. 2013. Rice oxalate oxidase gene driven by green tissue-specific promoter increases tolerance to sheath blight pathogen（Rhizoctonia solani）in transgenic rice. Molecular Plant Pathology，14 (9)：910–922.
[89]	Montgomery J，Goldman S，Deikman J，Margossian L，Fischer R L. 1993a. Identification of an ethylene-responsive region in the promoter of a fruit ripening gene. Proceedings of the National Academy of Sciences，90 (13)：5939–5943.
[90]	Montgomery J，Pollard V，Deikman J，Fischer R L. 1993b. Positive and negative regulatory regions control the spatial distribution of polygalacturonase transcription in tomato fruit pericarp. The Plant Cell，5 (9)：1049–1062.
[91]	Negi N，Khurana P. 2021. A salicylic acid inducible mulberry WRKY transcription factor，MiWRKY53 is involved in plant defence response. Plant Cell Reports，40：2151–2171.
[92]	Nielsen J，Keasling J D. 2016. Engineering cellular metabolism. Cell，164 (6)：1185–1197.
[93]	Nitz I，Berkefeld H，Puzio P S，Grundler F M. 2001. Pyk10，a seedling and root specific gene and promoter from Arabidopsis thaliana. Plant Science，161 (2)：337–346.
[94]	Odell J T，Nagy F，Chua N H. 1985. Identification of DNA sequences required for activity of the cauliflower mosaic virus 35S promoter. Nature，313 (6005)：810–812.
[95]	Ohme-Takagi M，Shinshi H. 1995. Ethylene-inducible DNA binding proteins that interact with an ethylene-responsive element. The Plant Cell，7 (2)：173–182.
[96]	Oliva R，Ji C，Atienza-Grande G，Huguet-Tapia J C，Perez-Quintero A，Li T，Eom J，Li C，Nguyen H，Liu B，Auguy F，Sciallano C，Luu V T，Dossa G S，Cunnac S，Schmidt S M，Slamet-Loedin I H，Cruz C V，Szurek B，Frommer W B，White F F，Yang B. 2019. Broad-spectrum resistance to bacterial blight in rice using genome editing. Nature Biotechnology，37 (11)：1344–1350.
[97]	Pellizzaro A，Neveu M，Lalanne D，Ly Vu B，Kanno Y，Seo M，Leprince O，Buitink J. 2020. A role for auxin signaling in the acquisition of longevity during seed maturation. New Phytologist，225 (1)：284–296.
[98]	Qian D，Zhang Z，He J，Zhang P，Ou X，Li T，Niu L，Nan Q，Niu Y，He W，An L，Jiang K，Xiang Y. 2019. Arabidopsis ADF5 promotes stomatal closure by regulating actin cytoskeleton remodeling in response to ABA and drought stress. Journal of Experimental Botany，70 (2)：435–446.
[99]	Quaedvlieg N E，Schlaman H R，Admiraal P C，Wijting S E，Stougaard J，Spaink H P. 1998. Fusions between green fluorescent protein and β-glucuronidase as sensitive and vital bifunctional reporters in plants. Plant Molecular Biology，38：861–873.
[100]	Rahman F U，Khan I A，Aslam A，Liu R，Sun L，Wu Y，Aslam M M，Khan A U，Li P，Jiang J，Fan X，Liu C，Zhang Y. 2022a. Transcriptome analysis reveals pathogenesis-related gene 1 pathway against salicylic acid treatment in grapevine（Vitis vinifera L）. Frontiers in Genetics，13：1033288.
[101]	Rahman F U，Zhang Y，Khan I A，Liu R，Sun L，Wu Y，Jiang J，Fan X，Liu C. 2022b. The promoter analysis of VvPR1 gene：A candidate gene identified through transcriptional profiling of methyl jasmonate treated grapevine（Vitis vinifera L.）. Plants，11 (12)：1540.
[102]	Reidt W，Wohlfarth T，Ellerström M，Czihal A，Tewes A，Ezcurra I，Rask L，Bäumlein H. 2000. Gene regulation during late embryogenesis：the RY motif of maturation-specific gene promoters is a direct target of the FUS3 gene product. The Plant Journal，21 (5)：401–408.
[103]	Ren C，Liu Y，Guo Y，Duan W，Fan P，Li S，Liang Z. 2021. Optimizing the CRISPR/Cas9 system for genome editing in grape by using grape promoters. Horticulture Research，8 (1)：52.
[104]	Rocher A，Dumas C，Cock J M. 2005. A W-box is required for full expression of the SA-responsive gene SFR2. Gene，344：181–192.
[105]	Rogers J C，Rogers S W. 1992. Definition and functional implications of gibberellin and abscisic acid cis-acting hormone response complexes. The Plant Cell，4 (11)：1443–1451.
[106]	Roque E，Gómez-Mena C，Hamza R，Beltrán J P，Cañas L A. 2019. Engineered male sterility by early anther ablation using the pea anther-specific promoter PsEND1. Frontiers in Plant Science，10：461361.
[107]	Ruan X M，Xiong X，Li J F. 2024. Identification and application of an exocarp-preferential promoter for genetic engineering of tomato fruit. Horticulture Research，11 (3)：uhae035.
[108]	Rushton P J. 2016. What have we learned about synthetic promoter construction? Plant Synthetic Promoters: Methods and Protocols，1482：1–13.
[109]	Sarkar S，Das A，Khandagale P，Maiti I B，Chattopadhyay S，Dey N. 2018. Interaction of Arabidopsis TGA3 and WRKY53 transcription factors on Cestrum yellow leaf curling virus（CmYLCV） promoter mediates salicylic acid-dependent gene expression in planta. Planta，247：181–199.
[110]	Singh S，Yadav S，Singh A，Mahima M，Singh A，Gautam V，Sarkar A K. 2020. Auxin signaling modulates LATERAL ROOT PRIMORDIUM1（LRP1）expression during lateral root development in Arabidopsis. The Plant Journal，101 (1)：87–100.
[111]	Skriver K，Olsen F L，Rogers J C，Mundy J. 1991. cis-Acting DNA elements responsive to gibberellin and its antagonist abscisic acid. Proceedings of the National Academy of Sciences，88 (16)：7266–7270.
[112]	Smale S T. 2001. Core promoters：active contributors to combinatorial gene regulation. Genes & Development，15 (19)：2503–2508.
[113]	Song S，Wang G，Wu H，Fan X，Liang L，Zhao H，Li S，Hu Y，Liu H，Ayaad M，Xing Y. 2020. OsMFT2 is involved in the regulation of ABA signaling-mediated seed germination through interacting with OsbZIP23/66/72 in rice. The Plant Journal，103 (2)：532–546.
[114]	Sun C，Palmqvist S，Olsson H，Borén M，Ahlandsberg S，Jansson C. 2003. A novel WRKY transcription factor，SUSIBA2，participates in sugar signaling in barley by binding to the sugar-responsive elements of the iso1 promoter. The Plant Cell，15 (9)：2076–2092.
[115]	Sun S R，Wu X B，Chen J S，Huang M T，Fu H Y，Wang Q N，Rott P，Gao S J. 2024. Identification of a sugarcane bacilliform virus promoter that is activated by drought stress in plants. Communications Biology，7 (1)：368.
[116]	Sun X M，Xiong H Y，Jiang C H，Zhang D M，Yang Z L，Huang Y P，Zhu W B，Ma S S，Duan J Z，Wang X，Liu W，Guo H F，Li G L，Qi J W，Liang C B，Zhang Z Y，Li J J，Zhang H L，Han L J，Zhou Y H，Peng Y L，Li Z C. 2022. Natural variation of DROT1 confers drought adaptation in upland rice. Nature Communications，13 (1)：4265.
[117]	Sutoh K，Yamauchi D. 2003. Two cis-acting elements necessary and sufficient for gibberellin-upregulated proteinase expression in rice seeds. The Plant Journal，34 (5)：635–645.
[118]	Szabała B M. 2023. A bifunctional selectable marker for wheat transformation contributes to the characterization of male-sterile phenotype induced by a synthetic Ms2 gene. Plant Cell Reports，42 (5)：895–907.
[119]	Tang H，Liu H，Zhou Y，Liu H，Du L，Wang K，Ye X. 2021. Fertility recovery of wheat male sterility controlled by Ms2 using CRISPR/Cas9. Plant Biotechnology Journal，19 (2)：224.
[120]	Tang W，Luo X，Samuels V. 2004. Regulated gene expression with promoters responding to inducers. Plant Science，166 (4)：827–834.
[121]	Thurow C，Schiermeyer A，Krawczyk S，Butterbrodt T，Nickolov K，Gatz C. 2005. Tobacco bZIP transcription factor TGA2. 2 and related factor TGA2. 1 have distinct roles in plant defense responses and plant development. The Plant Journal，44 (1)：100–113.
[122]	Tian L，Wu K，Hannam C，Latoszek-Green M，Sibbald S，Hu M，Brown D C W，Miki B. 2005. Analysis and use of the tobacco eIF4A-10 promoter elements for transgene expression. Journal of Plant Physiology，162 (12)：1355–1366.
[123]	Tounosu N，Sesoko K，Hori K，Shimojima M，Ohta H. 2023. cis-Regulatory elements and transcription factors related to auxin signaling in the streptophyte algae Klebsormidium nitens. Scientific Reports，13 (1)：9635.
[124]	Vallarino J G，Kubiszewski-Jakubiak S，Ruf S，Rößner M，Timm S，Bauwe H，Carrari F，Rentsch D，Bock R，Sweetlove L J，FernieA R. 2020. Multi-gene metabolic engineering of tomato plants results in increased fruit yield up to 23%. Scientific Reports，10 (1)：17219.
[125]	van Haaren M J J，Houck C M. 1991. Strong negative and positive regulatory elements contribute to the high-level fruit-specific expression of the tomato 2A11 gene. Plant Molecular Biology，17：615–630.
[126]	van Haaren M J J，Houck C M. 1993. A functional map of the fruit-specific promoter of the tomato 2A11 gene. Plant Molecular Biology，21：625–640.
[127]	Vaucheret H，Béclin C，Elmayan T，Feuerbach F，Godon C，Morel J B，Mourrain P，Palauqui J C，Vernhettes S. 1998. Transgene-induced gene silencing in plants. The Plant Journal，16 (6)：651–659.
[128]	Walcher C L，Nemhauser J L. 2012. Bipartite promoter element required for auxin response. Plant Physiology，158 (1)：273–282.
[129]	Wan L，Zhang J，Zhang H，Zhang Z，Quan R，Zhou S，Huang R. 2011. Transcriptional activation of OsDERF1 in OsERF3 and OsAP2-39 negatively modulates ethylene synthesis and drought tolerance in rice. PLoS ONE，6 (9)：e25216.
[130]	Wang C，Li W，Chen F，Cheng Y，Huang X，Zou B，Wang Y，Xu W，Qu S. 2022. Genome-wide identification and characterization of members of the ACS gene family in Cucurbita maxima and their transcriptional responses to the specific treatments. International Journal of Molecular Sciences，23 (15)：8476.
[131]	Wang J X，Ming X，Tao Y B，Xu Z F. 2020a. Jatropha curcas ortholog of tomato MADS-box gene 6（JcTM6）promoter exhibits floral-specific activity in Arabidopsis thaliana. PeerJ，8：e9827.
[132]	Wang M，Yan W，Peng X，Chen Z，Xu C，Wu J，Deng X Y，Tang X. 2020b. Identification of late-stage pollen-specific promoters for construction of pollen-inactivation system in rice. Journal of Integrative Plant Biology，62 (8)：1246–1263.
[133]	Wang X，Replogle A M Y，Davis E L，Mitchum M G. 2007a. The tobacco Cel7 gene promoter is auxin-responsive and locally induced in nematode feeding sites of heterologous plants. Molecular Plant Pathology，8 (4)：423–436.
[134]	Wang Y Q，Zhu S Y，Wang Y，Zhang M Y. 2007b. Tissue and inducible expression of a rice glutathione transporter gene promoter in transgenic Arabidopsis. Bot Stud，48：35–41.
[135]	Welsch R，Wust F，Bar C，Al-Babili S，Beyer P. 2008. A third phytoene synthase is devoted to abiotic stress-induced abscisic acid formation in rice and defines functional diversification of phytoene synthase genes. Plant Physiology，147 (1)：367–380.
[136]	Wichmann J，Behrendt G，Boecker S，Klamt S. 2023. Characterizing and utilizing oxygen-dependent promoters for efficient dynamic metabolic engineering. Metabolic Engineering，77：199–207.
[137]	Wiley S R，KRAus R J，Mertz J E. 1992. Functional binding of the“TATA”box binding component of transcription factor TFIID to the-30 region of TATA-less promoters. Proceedings of the National Academy of Sciences，89 (13)：5814–5818.
[138]	Wu A M，Lv S Y，Liu J Y. 2007. Functional analysis of a cotton glucuronosyltransferase promoter in transgenic tobaccos. Cell Research，17 (2)：174–183.
[139]	Wu C Y，Washida H，Onodera Y，Harada K，Takaiwa F. 2000. Quantitative nature of the Prolamin-box，ACGT and AACA motifs in a rice glutelin gene promoter：minimal cis-element requirements for endosperm-specific gene expression. The Plant Journal，23 (3)：415–421.
[140]	Wu J G，Yang Z R，Wang Y，Zheng L J，Ye R Q，Ji Y H，Zhao S S，Ji S Y，Liu R F，Xu L，Zheng H，Zhou Y J，Zhang X，Cao X F，Xie L H，Wu Z J. 2015. Viral-inducible argonaute18 confers broad-spectrum virus resistance in rice by sequestering a host microRNA. Elife，4：e05733.
[141]	Wu P，Xu X，Li J，Zhang J，Chang S，Yang X，Guo X. 2021. Seed-specific overexpression of cotton GhDGAT1 gene leads to increased oil accumulation in cottonseed. The Crop Journal，9 (2)：487–490.
[142]	Xia C，Zhang L C，Zou C，Gu Y Q，Duan J L，Zhao G Y，Wu J J，Liu Y，Fang X H，Gao L F，Jiao Y N，Sun J Q，Pan Y H，Liu X，Jia J Z，Kong X Y. 2017. A TRIM insertion in the promoter of Ms2 causes male sterility in wheat. Nature Communications，8 (1)：15407.
[143]	Xu H，Wang H，Zhang Y，Yang X，Lv S，Hou D，Mo C，Wassie M，Yu B，Hu T. 2023. A synthetic light-inducible photorespiratory bypass enhances photosynthesis to improve rice growth and grain yield. Plant Communications，4 (6)：100641.
[144]	Xu R，Goldman S，Coupe S，Deikman J. 1996. Ethylene control of E4 transcription during tomato fruit ripening involves two cooperative cis- elements. Plant Molecular Biology，31：1117–1127.
[145]	Xu Z S，Yang Q Q，Feng K，Yu X，Xiong A S. 2020. DcMYB113，a root-specific R2R3-MYB，conditions anthocyanin biosynthesis and modification in carrot. Plant Biotechnology Journal，18 (7)：1585–1597.
[146]	Xun H，Zhang X，Yu J，Pang J，Wang S，Liu B，Dong H，Jiang L，Guo D. 2021. Analysis of expression characteristics of soybean leaf and root tissue-specific promoters in Arabidopsis and soybean. Transgenic Research，30：799–810.
[147]	Yadav V K，Yadav V K，Pant P，Singh S P，Maurya R，Sable A，Sawant S V. 2017. GhMYB1 regulates SCW stage-specific expression of the GhGDSL promoter in the fibres of Gossypium hirsutum L. Plant Biotechnology Journal，15 (9)：1163–1174.
[148]	Yan L，Wei S，Wu Y，Hu R，Li H，Yang W，Xie Q. 2015. High-efficiency genome editing in Arabidopsis using YAO promoter-driven CRISPR/Cas9 system. Molecular Plant，8 (12)：1820–1823.
[149]	Yang Jiangtao，Pang Weimin，Wang Xujing，Lü Shaofu，Tang Qiaoling，Wang Zhixing. 2016. Cloning and deletion analysis of GhRACK1 promoter from Gossypium hirsutum. Acta Agronomica Sinica，42 (3)：368–375. (in Chinese)
	杨江涛，庞伟民，王旭静，吕少溥，唐巧玲，王志兴. 2016. 陆地棉GhRACK1启动子的克隆与缺失分析. 作物学报，42 (3)：368–375.
[150]	Yang Y Z，Tan B C. 2014. A distal ABA responsive element in AtNCED3 promoter is required for positive feedback regulation of ABA biosynthesis in Arabidopsis. PLoS ONE，9 (1)：e87283.
[151]	Yuan C，Li C，Yan C，Zhao X，Wang J，Sun Q，Shan S. 2019. Isolation and characterization of a novel seed-specific promoter from peanut（Arachis hypogaea L.）. Molecular Biology Reports，46：3183–3191.
[152]	Zerpa-Catanho D，Clough S J，Ming R. 2022. Characterization and analysis of the promoter region of monodehydroascorbate reductase 4（CpMDAR4）in papaya. Plant Reproduction，35 (4)：233–264.
[153]	Zhai X，Ji L，Gao J，Zhou Y J. 2021. Characterizing methanol metabolism-related promoters for metabolic engineering of Ogataea polymorpha. Applied Microbiology and Biotechnology，105：8761–8769.
[154]	Zhang C，Pan S，Chen H，Cai T，Zhuang C，Deng Y，Zhuang Y，Zeng Y，Chen S，Zhuang W. 2016. Characterization of NtREL1，a novel root-specific gene from tobacco，and upstream promoter activity analysis in homologous and heterologous hosts. Plant Cell Reports，35：757–769.
[155]	Zhang S，Li X，Fan S，Zhou L，Wang Y. 2020a. Overexpression of HcSCL13，a Halostachys caspica GRAS transcription factor，enhances plant growth and salt stress tolerance in transgenic Arabidopsis. Plant Physiology and Biochemistry，151：243–254.
[156]	Zhang S，Wu S，Hu C，Yang Q，Dong T，Sheng O，Deng G，He W，Dou T，Li C，Sun C，Yi G，Bi F. 2022. Increased mutation efficiency of CRISPR/Cas9 genome editing in banana by optimized construct. PeerJ，10：e12664.
[157]	Zhang W，Wang N，Yang J，Guo H，Liu Z，Zheng X，Li S，Xiang F. 2020b. The salt-induced transcription factor GmMYB84 confers salinity tolerance in soybean. Plant Science，291：110326.
[158]	Zhao Y，Sun Y，Huang S，Liu Z，Feng H. 2022. Identification of an anther-specific promoter from a male sterile AB line in Chinese cabbage （Brassica rapa L. ssp. pekinensis）. 3 Biotech，12 (4)：104.
[159]	Zheng C，Zhang L，Tang Z，Zhao C，Yuan S. 2009. TAIL-PCR and its cloning in plant gene. Genomics and Applied Biology，28 (3)：544–548.
[160]	Zheng H，Lei Y，Lin S，Zhang Q，Zhang Z. 2011. Bidirectionalization of a methyl jasmonate-inducible plant promoter. Biotechnology Letters，33：387–393.
[161]	Zheng N，Li T，Dittman J D，Su J，Li R，Gassmann W，Peng D，Whitham S A，Liu S，Yang B. 2020. CRISPR/Cas9-based gene editing using egg cell-specific promoters in Arabidopsis and soybean. Frontiers in Plant Science，11：551133.
[162]	Zhou J，Liu G，Zhao Y，Zhang R，Tang X，Li L，Jia X，Guo Y，Wu Y，Han Y，Bao Y，He Y，Han Q，Yang H，Zheng X，Qi Y，Zhang T，Zhang Y. 2023. An efficient CRISPR–Cas12a promoter editing system for crop improvement. Nature Plants，9 (4)：588–604.
[163]	Zhou Y，Ma B，Tao J J，Yin C C，Hu Y，Huang Y H，Wei W，Xin P Y，Chu J F，Zhang W K，Chen S Y，Zhang J S. 2022. Rice EIL1 interacts with OsIAAs to regulate auxin biosynthesis mediated by the tryptophan aminotransferase MHZ10/OsTAR2 during root ethylene responses. The Plant Cell，34 (11)：4366–4387.
[164]	Zou X，Song E，Peng A，He Y，Xu L，Lei T，Yao L，Chen S. 2014. Activation of three pathogen-inducible promoters in transgenic citrus（Citrus sinensis Osbeck）after Xanthomonas axonopodis pv. citri infection and wounding. Plant Cell，Tissue and Organ Culture，117：85–98.