[1]唐军,张奥妮,吕丹,等.CRISPR/Cas基因编辑系统在作物遗传改良上的应用研究进展[J].江苏农业科学,2026,54(3):11-21.
　Tang Jun,et al.Research progress on application of CRISPR/Cas gene editing system in crop genetic improvement[J].Jiangsu Agricultural Sciences,2026,54(3):11-21.
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CRISPR/Cas基因编辑系统在作物遗传改良上的应用研究进展()

《江苏农业科学》[ISSN:1002-1302/CN:32-1214/S]

卷:
第54卷
期数:
2026年第3期
页码:
11-21
栏目:
专论与综述
出版日期:
2026-02-05

文章信息/Info

Title:
Research progress on application of CRISPR/Cas gene editing system in crop genetic improvement
作者:
唐军; 张奥妮; 吕丹; 陈佳宁; 吴亚兰; 曾庆平; 邓敏
湖南农业大学农学院/岳麓山实验室/湖南省玉米工程技术研究中心，湖南长沙 410128
Author(s):
Tang Jun; et al
关键词:
基因编辑; CRISPR/Cas; 遗传改良; 靶标基因; 农作物
Keywords:
-
分类号:
S188+.1
DOI:
-
文献标志码:
A
摘要:
CRISPR/Cas系统作为第3代基因编辑技术的核心工具，由成簇规则间隔短回文重复序列(CRISPR)及其关联蛋白(Cas)共同构成。该技术通过向导RNA精确定位靶标基因，利用Cas蛋白的核酸酶活性实现基因组特定位点的精准编辑。相较于传统育种技术，CRISPR/Cas系统在编辑效率、操作周期和靶向准确性方面具有显著优势，已成功应用于玉米、水稻、小麦等主粮作物的性状改良。在作物遗传改良领域，CRISPR/Cas系统的应用主要聚焦于5个关键方向：(1)产量相关性状调控；(2)株型结构优化；(3)生物/非生物抗性增强；(4)品质性状改良；(5)杂种优势利用。重点探讨CRISPR/Cas蛋白在作物产量、株高、抗性、品质以及杂种优势利用上的具体应用，并剖析该技术面临的潜在挑战。
Abstract:
-

参考文献/References:

［1］于熙婷，黄学辉. 现代玉米起源新见解：两类大刍草的混血［J］. 植物学报，2023，58(6)：857-860.
［2］Animasaun D A，Oguntoye E O. Mutagenesis in crop improvement：methods and applications［J］. Journal of Crop Improvement，2024，38(3)：156-178.
［3］Zafra M P，Schatoff E M，Katti A，et al. Optimized base editors enable efficient editing in cells，organoids and mice［J］. Nature Biotechnology，2018，36(9)：888-893.
［4］Usman B，Zhao N，Nawaz G，et al. CRISPR/Cas9 guided mutagenesis of grain size 3 confers increased rice(Oryza sativa L.) grain length by regulating cysteine proteinase inhibitor and ubiquitin-related proteins［J］. International Journal of Molecular Sciences，2021，22(6)：3225.
［5］Liu L，Gallagher J，Arevalo E D，et al. Enhancing grain-yield-related traits by CRISPR-Cas9 promoter editing of maize CLE genes［J］. Nature Plants，2021，7(3)：287-294.
［6］Kwon C T，Tang L L，Wang X G，et al. Dynamic evolution of small signalling peptide compensation in plant stem cell control［J］. Nature Plants，2022，8(4)：346-355.
［7］Bolduc N，Yilmaz A，Mejia-Guerra M K，et al. Unraveling the KNOTTED1 regulatory network in maize meristems［J］. Genes & Development，2012，26(15)：1685-1690.
［8］Fei X H，Wang Y F，Zheng Y X，et al. Identification of two new QTLs of maize(Zea mays L.) underlying kernel row number using the HNAU-NAM1 population［J］. BMC Genomics，2022，23(1)：593.
［9］贾婷. 玉米灌浆调控转录因子ZmNAC128和ZmNAC130优异单倍型的鉴定［D］. 雅安：四川农业大学，2023.
［10］ Wang Y H，Tang Q L，Kang Y L，et al. Analysis of the utilization and prospects of CRISPR-Cas technology in the annotation of gene function and creation new germplasm in maize based on patent data［J］. Cells，2022，11(21)：3471-3471.
［11］Chen Q Q，Guo Y M，Zhang J，et al. RNA polymerase common subunit ZmRPABC5b is transcriptionally activated by Opaque2 and essential for endosperm development in maize［J］. Nucleic Acids Research，2023，51(15)：7832-7850.
［12］Baye W，Xie Q，Xie P，et al. Genetic architecture of grain yield-related traits in sorghum and maize［J］. International Journal of Molecular Sciences，2022，23(5)：2405-2405.
［13］Yang N，Liu J，Gao Q，et al. Genome assembly of a tropical maize inbred line provides insights into structural variation and crop improvement［J］. Nature Genetics，2019，51(6)：1052-1059.
［14］梁彦，王永红. 水稻株型功能基因及其在育种上的应用［J］. 生命科学，2016，28(10)：1156-1167.
［15］Lai D L，Fan Y，Xue G X，et al. Genome-wide identification and characterization of the SPL gene family and its expression in the various developmental stages and stress conditions in foxtail millet(Setaria italica)［J］. BMC Genomics，2022，23(1)：389.
［16］Jiao Y Q，Wang Y H，Xue D W，et al. Regulation of OsSPL14 by OsmiR156 defines ideal plant architecture in rice［J］. Nature Genetics，2010，42(6)：541-544.
［17］Chen W K，Chen L，Zhang X，et al. Convergent selection of a WD40 protein that enhances grain yield in maize and rice［J］. Science，2022，375(6587)：eabg7985.
［18］Usman B，Nawaz G，Zhao N，et al. Programmed editing of rice(Oryza sativa L.) OsSPL16 gene using CRISPR/Cas9 improves grain yield by modulating the expression of pyruvate enzymes and cell cycle proteins［J］. International Journal of Molecular Sciences，2020，22(1)：249.
［19］Li Y J，Wu S，Huang Y Y，et al. OsMADS17 simultaneously increases grain number and grain weight in rice［J］. Nature Communications，2023，14：3098.
［20］Yang H，Wang Y L，Tian Y L，et al. Rice FLOURY ENDOSPERM22，encoding a pentatricopeptide repeat protein，is involved in both mitochondrial RNA splicing and editing and is crucial for endosperm development［J］. Journal of Integrative Plant Biology，2023，65(3)：755-771.
［21］Liu Z M，Jiang S，Jiang L L，et al. Transcription factor OsSGL is a regulator of starch synthesis and grain quality in rice［J］. Journal of Experimental Botany，2022，73(11)：3417-3430.
［22］卢寰，黎凌，张景六，等. OsRAMOSA2基因调控水稻的粒形［J］. 植物生理学报，2015，51(3)：307-316.
［23］Ashokkumar S，Jaganathan D，Ramanathan V，et al. Creation of novel alleles of fragrance gene OsBADH2 in rice through CRISPR/Cas9 mediated gene editing［J］. PLoS One，2020，15(8)：e0237018.
［24］Jouanin A，Gilissen L J W J，Schaart J G，et al. CRISPR/Cas9 gene editing of gluten in wheat to reduce gluten content and exposure-reviewing methods to screen for coeliac safety［J］. Frontiers in Nutrition，2020，7：51.
［25］Shi J H，Ni X Y，Huang J X，et al. CRISPR/Cas9-mediated gene editing of BnFAD2 and BnFAE1 modifies fatty acid profiles in Brassica napus［J］. Genes，2022，13(10)：1681.
［26］Wu W X，Zheng X M，Lu G W，et al. Association of functional nucleotide polymorphisms at DTH2 with the northward expansion of rice cultivation in Asia［J］. PNAS，2013，110(8)：2775-2780.
［27］Tian J G，Wang C L，Chen F Y，et al. Maize smart-canopy architecture enhances yield at high densities［J］. Nature，2024，632(8025)：576-584.
［28］Tian J G，Wang C L，Xia J L，et al. Teosinte ligule allele narrows plant architecture and enhances high-density maize yields［J］. Science，2019，365(6454)：658-664.
［29］Chen H，Gong X P，Guo Y，et al. ZmbZIP27 regulates nitrogen-mediated leaf angle by modulating lignin deposition in maize［J］. The Crop Journal，2024，12(5)：1404-1413.
［30］Wang X K，Wang X L，Sun S L，et al. Characterization of regulatory modules controlling leaf angle in maize［J］. Plant Physiology，2022，190(1)：500-515.
［31］Liu Y，Chen Z Q，Zhang C，et al. Gene editing of ZmGA20ox3 improves plant architecture and drought tolerance in maize［J］. Plant Cell Reports，2023，43(1)：18.
［32］Sun C R，Liu Y，Li G F，et al. ZmCYP90D1 regulates maize internode development by modulating brassinosteroid-mediated cell division and growth［J］. The Crop Journal，2024，12(1)：58-67.
［33］Liang Y M，Liu Q，Wang X F，et al. ZmMADS69 functions as a flowering activator through the ZmRap2.7-ZCN8 regulatory module and contributes to maize flowering time adaptation［J］. The New Phytologist，2019，221(4)：2335-2347.
［34］Yu J，Song G S，Guo W J，et al. ZmBELL10 interacts with other ZmBELLs and recognizes specific motifs for transcriptional activation to modulate internode patterning in maize［J］. New Phytologist，2023，240(2)：577-596.
［35］杨晨曦，周文期，周香艳，等. 控制玉米株高基因PHR1的基因克隆［J］. 作物学报，2024，50(1)：55-66.
［36］徐崟海，刘佳. IGT基因家族调控作物株型研究进展［J］. 生物技术进展，2022，12(5)：673-682.
［37］王凯婕，安文静，刘亚菲，等. CRISPR/Cas9技术编辑OsRhoGDI2基因导致水稻半矮化［J］. 生物工程学报，2020，36(4)：707-715.
［38］Liu H L，Xu Y Y，Xu Z H，et al. A rice YABBY gene，OsYABBY4，preferentially expresses in developing vascular tissue［J］. Development Genes and Evolution，2007，217(9)：629-637.
［39］Sharan A，Soni P，Nongpiur R C，et al. Mapping the ‘Two-component system’ network in rice［J］. Scientific Reports，2017，7(1)：9287.
［40］Miao J，Guo D S，Zhang J Z，et al. Targeted mutagenesis in rice using CRISPR-Cas system［J］. Cell Research，2013，23(10)：1233-1236.
［41］Das D，Singha D L，Paswan R R，et al. Recent advancements in CRISPR/Cas technology for accelerated crop improvement［J］. Planta，2022，255(5)：109.
［42］Tang H W，Fan J L，Wang R，et al. Changes in the expression pattern of OsWUS negatively regulate plant stature and panicle development in rice［J］. G3，2023，13(7)：jkad100.
［43］Brabham H J，Gómez De La Cruz D，Were V，et al. Barley MLA3 recognizes the host-specificity effector Pwl2 from Magnaporthe Oryzae［J］. The Plant Cell，2024，36(2)：447-470.
［44］李彩华，赵彦坤，李占坤，等. 小麦矮秆基因研究进展［J］. 生物技术进展，2024，14(6)：980-992.
［45］Xu D A，Jia C F，Lyu X R，et al. In silico curation of QTL-rich clusters and candidate gene identification for plant height of bread wheat［J］. The Crop Journal，2023，11(5)：1480-1490.
［46］卢昆丽，尹燕枰，王振林，等. 施氮期对小麦茎秆木质素合成的影响及其抗倒伏生理机制［J］. 作物学报，2014，40(9)：1686-1694.
［47］Park J R，Jang Y H，Kim E G，et al. Quantitative trait loci mapping identified candidate genes involved in plant height regulation in rice［J］. International Journal of Molecular Sciences，2023，24(23)：16895.
［48］Pan Y H，Chen L，Zhao Y，et al. Natural variation in OsMKK3 contributes to grain size and chalkiness in rice［J］. Frontiers in Plant Science，2021，12：784037.
［49］Qie Y M，Liu Y，Wang M N，et al. Development，validation，and re-selection of wheat lines with pyramided genes Yr64 and Yr15 linked on the short arm of chromosome 1B for resistance to stripe rust［J］. Plant Disease，2019，103(1)：51-58.
［50］Wang Y P，Cheng X，Shan Q W，et al. Simultaneous editing of three homoeoalleles in hexaploid bread wheat confers heritable resistance to powdery mildew［J］. Nature Biotechnology，2014，32(9)：947-951.
［51］Ali Q，Yu C J，Hussain A，et al. Genome engineering technology for durable disease resistance：recent progress and future outlooks for sustainable agriculture［J］. Frontiers in Plant Science，2022，13：860281.
［52］Yang P，Scheuermann D，Kessel B，et al. Alleles of a wall-associated kinase gene account for three of the major northern corn leaf blight resistance loci in maize［J］. The Plant Journal，2021，106(2)：526-535.
［53］Zhu M，Tong L X，Xu M L，et al. Genetic dissection of maize disease resistance and its applications in molecular breeding［J］. Molecular Breeding，2021，41(5)：32.
［54］Ku L X，Tian L，Su H H，et al. Dual functions of the ZmCCT-associated quantitative trait locus in flowering and stress responses under long-day conditions［J］. BMC Plant Biology，2016，16(1)：239.
［55］Skamnioti P，Gurr S J. Against the grain：safeguarding rice from rice blast disease［J］. Trends in Biotechnology，2009，27(3)：141-150.
［56］林春姿，黄其伟，侯艳，等. 水稻多胺氧化酶基因(OsPAO4)CRISPR/Cas9编辑突变体的创制［J］. 广东农业科学，2023，50(3)：1-10.
［57］Huang Y Y，Liu X X，Xie Y，et al. Identification of FERONIA-like receptor genes involved in rice-Magnaporthe oryzae interaction［J］. Phytopathology Research，2020，2(1)：14.
［58］Gao Y，Zhang C，Han X，et al. Inhibition of OsSWEET11 function in mesophyll cells improves resistance of rice to sheath blight disease［J］. Molecular Plant Pathology，2018，19(9)：2149-2161.
［59］Wang C L，Zhang X P，Fan Y L，et al. XA23 is an executor R protein and confers broad-spectrum disease resistance in rice［J］. Molecular Plant，2014：ssu132.
［60］Sha G，Sun P，Kong X J，et al. Genome editing of a rice CDP-DAG synthase confers multipathogen resistance［J］. Nature，2023，618(7967)：1017-1023.
［61］赵丽丽，郭晨，牟春雨，等. 利用CRISPR/Cas9技术敲除BnERF019降低甘蓝型油菜对黑胫病的抗性［J］. 中国油料作物学报，2025，47(2)：319-327.
［62］Pramanik D，Shelake R M，Park J，et al. CRISPR/Cas9-mediated generation of pathogen-resistant tomato against tomato yellow leaf curl virus and powdery mildew［J］. International Journal of Molecular Sciences，2021，22(4)：1878-1878..
［63］Shawky A，Hatawsh A，Al-Saadi N，et al. Revolutionizing tomato cultivation：CRISPR/Cas9 mediated biotic stress resistance［J］. Plants(Basel)，2024，13(16)：2269.
［64］Bhaskar P B，Raasch J A，Kramer L C，et al. Sgt1，but not Rar1，is essential for the RB-mediated broad-spectrum resistance to potato late blight［J］. BMC Plant Biology，2008，8(1)：8.
［65］Li J，Coates B S，Kim K S，et al. The genetic structure of Asian corn borer，Ostrinia furnacalis，populations in China：haplotype variance in northern populations and potential impact on management of resistance to transgenic maize［J］. The Journal of Heredity，2014，105(5)：642-655.
［66］Rice R .Researchers at huazhong agricultural university report new data on rice research ［Molecular breeding of rice restorer lines and hybrids for brown planthopper(BPH) resistance using the Bph14 and Bph15 genes］［J］. Agriculture Week，2017，132-.
［67］Wu K，Shirk P D，Taylor C E，et al. CRISPR/Cas9 mediated knockout of the abdominal-a homeotic gene in fall armyworm moth(Spodoptera frugiperda)［J］. PLoS One，2018，13(12)：e0208647.
［68］Wu X，Feng H，Wu D，et al. Using high-throughput multiple optical phenotyping to decipher the genetic architecture of maize drought tolerance［J］. Genome Biology，2021，22(1)：185.
［69］Liu S X，Wang X L，Wang H W，et al. Genome-wide analysis of ZmDREB genes and their association with natural variation in drought tolerance at seedling stage of Zea mays L.［J］. PLoS Genetics，2013，9(9)：e1003790.
［70］Quiroga G，Erice G，Ding L，et al. The arbuscular mycorrhizal symbiosis regulates aquaporins activity and improves root cell water permeability in maize plants subjected to water stress［J］. Plant，Cell & Environment，2019，42(7)：2274-2290.
［71］Kobayashi N I，Yamaji N，Yamamoto H，et al. OsHKT1；5 mediates Na+exclusion in the vasculature to protect leaf blades and reproductive tissues from salt toxicity in rice［J］. The Plant Journal，2017，91(4)：657-670.
［72］Do P T，Pham H Q，Nguyen H M，et al. Assessment of sequence polymorphism and gene expression of OsSOS1 gene in two contrasting rice genotypes［J］. Acta Biologica Cracoviensia s Botanica，2018，60(1)：7-16.
［73］Wang C T，Ru J N，Liu Y W，et al. The maize WRKY transcription factor ZmWRKY40 confers drought resistance in transgenic Arabidopsis［J］. International Journal of Molecular Sciences，2018，19(9)：2580.
［74］AlHusnain L，AlKahtani M D F，Attia K A，et al. Application of CRISPR/Cas9 system to knock out GluB gene for developing low glutelin rice mutant［J］. Botanical Studies，2024，65(1)：27.
［75］蔡梦颖. 利用基因编辑技术靶向SSSⅡb基因改良稻米品质［D］. 南京：南京农业大学，2020.
［76］Yang Y H，Shen Z Y，Xu C D，et al. Genetic improvement of panicle-erectness Japonica rice toward both yield and eating and cooking quality［J］. Molecular Breeding，2020，40(5)：51.
［77］Wang S Y，Yang Y H，Guo M，et al. Targeted mutagenesis of amino acid transporter genes for rice quality improvement using the CRISPR/Cas9 system［J］. The Crop Journal，2020，8(3)：457-464.
［78］Rao Y C，Yang X，Pan C Y，et al. Advance of clustered regularly interspaced short palindromic repeats-Cas9 system and its application in crop improvement［J］. Frontiers in Plant Science，2022，13：839001.
［79］Gao H R，Gadlage M J，Lafitte H R，et al. Superior field performance of waxy corn engineered using CRISPR-Cas9［J］. Nature Biotechnology，2020，38(5)：579-581.
［80］Chen S H，Yang Y，Shi W W，et al. Badh2，encoding betaine aldehyde dehydrogenase，inhibits the biosynthesis of 2-acetyl-1-pyrroline，a major component in rice fragrance［J］. The Plant Cell，2008，20(7)：1850-1861.
［81］Mano J，Masubuchi T，Hatano S，et al. Development and validation of event-specific quantitative PCR method for genetically modified maize LY038［J］. Journal of the Food Hygienic Society of Japan，2013，54(1)：25-30.
［82］王义鹏. 拟南芥胱硫醚-γ-合成酶(D-AtCGS)基因在大肠杆菌中的表达及抗血清制备［D］. 哈尔滨：东北林业大学，2011.
［83］Yang D N，Ni R Z，Yang S H，et al. Functional characterization of the Stipa purpurea；P5CS gene under drought stress conditions［J］. International Journal of Molecular Sciences，2021，22(17)：9599-9599.
［84］舒心媛，严旭，蒲烨弘，等. CRISPR/Cas系统的作用原理及其在作物遗传改良中的应用［J］. 浙江大学学报(农业与生命科学版)，2018，44(3)：259-268，381.
［85］葛鲁安，王豪，赵宁，等. 玉米田反枝苋对烟嘧磺隆的抗性水平及靶标抗性分子机理［J］. 农药学学报，2018，20(5)：595-600.
［86］Ko E，Karayigˇit B. Assessment of biofortification approaches used to improve micronutrient-dense plants that are a sustainable solution to combat hidden hunger［J］. Journal of Soil Science and Plant Nutrition，2022，22(1)：475-500.
［87］Roberts L A，Pierson A J，Panaviene Z，et al. Yellow stripe1.Expanded roles for the maize iron-phytosiderophore transporter［J］. Plant Physiology，2004，135(1)：112-120.
［88］White P J. Selenium accumulation by plants［J］. Annals of Botany，2016，117(2)：217-235.
［89］He Z Q，Shang X，Zhang T H，et al. Effect of calcium and magnesium on starch synthesis in maize kernels and its physiological driving mechanism［J］. Frontiers in Plant Science，2024，14：1332517.
［90］Zheng Y P，Guo T，Xia T，et al. Utility of Arabidopsis KASII promoter in development of an effective CRISPR/Cas9 system for soybean genome editing and its application in engineering of soybean seeds producing super-high oleic and low saturated oils［J］. Journal of Agricultural and Food Chemistry，2024，72(39)：21720-21730.
［91］Fu Y，Yao M，Qiu P，et al. Identification of transcription factor BnHDG4-A08 as a novel candidate associated with the accumulation of oleic，linoleic，linolenic，and erucic acid in Brassica napus［J］. Theoretical and Applied Genetics，2024，137(10)：243.
［92］Kaur N，Alok A，Shivani，et al. CRISPR/Cas9 directed editing of lycopene Epsilon-cyclase modulates metabolic flux for β-carotene biosynthesis in banana fruit［J］. Metabolic Engineering，2020，59：76-86.
［93］Sagor G H M，Berberich T，Tanaka S，et al. A novel strategy to produce sweeter tomato fruits with high sugar contents by fruit-specific expression of a single bZIP transcription factor gene［J］. Plant Biotechnology Journal，2016，14(4)：1116-1126.
［94］Liu T F，Fang H，Liu J，et al. Cytosolic glyceraldehyde-3-phosphate dehydrogenases play crucial roles in controlling cold-induced sweetening and apical dominance of potato(Solanum tuberosum L.) tubers［J］. Plant，Cell & Environment，2017，40(12)：3043-3054.
［95］Xu X，Yuan Y J，Feng B H，et al. CRISPR/Cas9-mediated gene-editing technology in fruit quality improvement［J］. Food Quality and Safety，2020，4(4)：159-166.
［96］Melonek J，Duarte J，Martin J，et al. The genetic basis of cytoplasmic male sterility and fertility restoration in wheat［J］. Nature Communications，2021，12：1036.
［97］Wu L Y，Jing X H，Zhang B L，et al. A natural allele of OsMS1 responds to temperature changes and confers thermosensitive genic male sterility［J］. Nature Communications，2022，13：2055.
［98］Zhang S M，Wu S W，Niu C F，et al. ZmMs25 encoding a plastid-localized fatty acyl reductase is critical for anther and pollen development in maize［J］. Journal of Experimental Botany，2021，72(12)：4298-4318.
［99］Ye M W，Peng Z，Tang D，et al. Generation of self-compatible diploid potato by knockout of S-RNase［J］. Nature Plants，2018，4(9)：651-654.
［100］Lee S，Enciso-Rodriguez F E，Behling W，et al. HT-B and S-RNase CRISPR-Cas9 double knockouts show enhanced self-fertility in diploid Solanum tuberosum［J］. Frontiers in Plant Science，2023，14：1151347.
［101］王春雷，赵宪坤，高季平，等. 芸薹属植物自交不亲和性研究进展［J］. 江苏农业学报，2015，31(6)：1442-1447.
［102］Qin X Q，Chetelat R T. Ornithine decarboxylase genes contribute to S-RNase-independent pollen rejection［J］. Plant Physiology，2021，186(1)：452-468.
［103］Wang C，Liu Q，Shen Y，et al. Clonal seeds from hybrid rice by simultaneous genome engineering of meiosis and fertilization genes［J］. Nature Biotechnology，2019，37(3)：283-286.
［104］Schenkelaars P，de Vriend H，Kalaitzandonakes N. Drivers of consolidation in the seed industry and its consequences for innovation［D］. Amsterdam：Commission on Genetic Modification，2011.
［105］Barman H N，Sheng Z H，Fiaz S，et al. Generation of a new thermo-sensitive genic male sterile rice line by targeted mutagenesis of TMS5 gene through CRISPR/Cas9 system［J］. BMC Plant Biology，2019，19(1)：109.
［106］Hu J C，Li S Y，Li Z L，et al. A barley stripe mosaic virus-based guide RNA delivery system for targeted mutagenesis in wheat and maize［J］. Molecular Plant Pathology，2019，20(10)：1463-1474.
［107］Jiang Y L，An X L，Li Z W，et al. CRISPR/Cas9-based discovery of maize transcription factors regulating male sterility and their functional conservation in plants［J］. Plant Biotechnology Journal，2021，19(9)：1769-1784.
［108］Gautam R D，Kumar A，Singh S，et al. A comprehensive overview of breeding strategy to improve phenotypic and quality traits in Valeriana jatamansi Jones［J］. Heliyon，2023，9(8)：e18294.
［109］Koonin E V，Gootenberg J S，Abudayyeh O O. Discovery of diverse CRISPR-cas systems and expansion of the genome engineering toolbox［J］. Biochemistry，2023，62(24)：3465-3487.
［110］钟静丽，林健香，周建奎，等. 碱基编辑系统的研究进展［J］. 生物工程学报，2024，40(5)：1271-1292.
［111］Xu J Y，Lian W，Jia Y N，et al. Optimized guide RNA structure for genome editing via Cas9［J］. Oncotarget，2017，8(55)：94166-94171.
［112］Anzalone A V，Gao X D，Podracky C J，et al. Programmable deletion，replacement，integration and inversion of large DNA sequences with twin prime editing［J］. Nature Biotechnology，2022，40(5)：731-740.
［113］Li B，Fu C Y，Zhou J W，et al. Highly efficient genome editing using geminivirus-based CRISPR/Cas9 system in cotton plant［J］. Cells，2022，11(18)：2902.

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　Li Li,et al.New progress and prospect of gene editing technology[J].Jiangsu Agricultural Sciences,2018,46(3):5.
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备注/Memo

备注/Memo:
收稿日期：2025-03-29
基金项目：国家自然科学基金(编号：32101700)；中国博士后科学基金面上项目(编号：2022M711122)。
作者简介：唐军(2000—)，男，湖南桃源人，硕士研究生，主要研究方向为玉米遗传育种与品质功能分析。E-mail：1121663859@qq.com。
通信作者：邓敏，博士，副教授，主要研究方向为玉米遗传育种和功能基因组学研究。E-mail：hdengmin@163.com。
更新日期/Last Update: 2026-02-05