生物技术通报 ›› 2026, Vol. 42 ›› Issue (3): 283-293.doi: 10.13560/j.cnki.biotech.bull.1985.2025-0704
罗威1,2(
), 宫奥1, 仲阳3, 胡迪1, 周洪源1, 张泓欣1, 艾菊1, 罗有卫4, 高冬丽1()
收稿日期:2025-07-01
出版日期:2026-03-26
发布日期:2026-04-23
通讯作者:
高冬丽，女，博士，研究员，研究方向 ：果实发育调控；E-mail: gdongli@126.com作者简介:罗威，男，博士研究生，研究方向 ：基因功能解析；E-mail: 1136689701@qq.com基金资助:
LUO Wei1,2(), GONG Ao1, ZHONG Yang3, HU Di1, ZHOU Hong-yuan1, ZHANG Hong-xin1, AI Ju1, LUO You-wei4, GAO Dong-li1()
Received:2025-07-01
Published:2026-03-26
Online:2026-04-23
摘要：
目的 SEPALLATA2（SEP2）编码MADS家族转录因子，利用Cssep2突变体揭示其在黄瓜（Cucumis sativus L.）果实和疣状结构发育中的作用，并探究其调控机制，为黄瓜重要性状的深度解析提供理论支持。 方法 通过实时荧光定量分析CsSEP2的表达模式；亚细胞定位分析CsSEP2在细胞内的具体位置；通过稳定遗传转化创制Cssep2敲除突变体，从细胞学和转录组层面探究其表型出现的分子机制；利用酵母双杂交和萤火虫荧光素酶互补实验分析CsSEP2和其他蛋白之间的相互作用。 结果 CsSEP2的mRNA特异性积累于花和果实组织，其编码蛋白定位于细胞核。利用CRISPR/Cas9技术构建功能缺失突变体。突变体果实因细胞分裂与伸长进程受阻而显著缩短，且呈现开裂表型；果实疣状结构发育异常，表现为果刺密度增加、果刺基部缩小及果瘤萎缩。外果皮与胎座的转录组测序分析显示，CsSEP2功能缺失显著改变基因表达谱，尤其是与果实大小和疣状结构发育相关基因表达发生显著变化。CsSEP2与多个已知的调控疣状结构发育的蛋白存在互作关系。 结论 CsSEP2突变对果实和疣状结构发育产生了多效性影响。CsSEP2通过影响一系列基因的表达和蛋白间的互作来调控果实长度和疣状结构发育。
罗威, 宫奥, 仲阳, 胡迪, 周洪源, 张泓欣, 艾菊, 罗有卫, 高冬丽. 黄瓜SEPALLATA2基因敲除对果实及疣状结构发育的多效性影响[J]. 生物技术通报, 2026, 42(3): 283-293.
LUO Wei, GONG Ao, ZHONG Yang, HU Di, ZHOU Hong-yuan, ZHANG Hong-xin, AI Ju, LUO You-wei, GAO Dong-li. Pleiotropic Effects of SEPALLATA2 Knock-out on Fruit and Wart Development in Cucumber[J]. Biotechnology Bulletin, 2026, 42(3): 283-293.
图1 CsSEP2基因表达分析（A）与亚细胞定位（B）图A中，检测CsSEP2在-3、0、5、10、16 DAA的果实，以及16 DAA果实中部外果皮、中果皮和胎座组织中的基因表达。数据以“均值±标准差”表示，3次生物学重复。DAA表示花后天数，下同
Fig. 1 Expression analyses (A) and subcellular localization (B) of gene CsSEP2In Fig. A, the expressions of CsSEP2 in thefruits of -3, 0, 5, 10, and 16 DAA and epicarp, mesocarp and placenta from the central parts of fruits at 16 DAA were examined. Data are means ± SD, n = 3. DAA refers to days after anthesis, the same below
图2 CsSEP2基因敲除导致果实发育异常A：CsSEP2基因结构及Cssep2突变体的突变位点示意图；外显子和内含子分别以黑色方框和直线表示，红色核苷酸序列为基因编辑靶点，下划线标注的3个碱基为前间隔序列邻近基序（PAM）；B、C：WT和Cssep2突变体植株（B）及果实（C）的代表性表型；D：WT和Cssep2突变体果实长度测定，5次生物学重复；E：Cssep2突变体果实开裂的表型；F：WT和Cssep2突变体果实胎座的对比；G：WT和Cssep2突变体果实胎座面积与横截面积的比值分析，样本量n ≥ 5，数据以“均值 ± 标准差”表示
Fig. 2 Knock-out of gene CsSEP2 resulted in abnormal fruit developmentA: Structure of the CsSEP2 gene and the mutated loci in the Cssep2 mutants. Exons and introns are denoted by black boxes and lines, respectively. The red nucleotide sequences are the editing target site of the gene, and the three underlined bases indicate the protospacer adjacent motif (PAM). B, C: Representative plants (B) and fruits (C) of WT and Cssep2 mutants. D: Measurement of the fruit lengths of WT and Cssep2 mutants. Data are means ± SD, n = 5. E: Macroscopic observation of fruit dehiscence of the Cssep2 mutants. F: Comparative observation of the fruit placentas of WT and Cssep2 mutants. G: The ratio of placenta areas to fruit cross areas of WT and Cssep2 mutants. Values are means ± SD, n ≥ 5
图3 Cssep2突变体果实及疣状结构的细胞学分析A、B：中果皮纵向细胞数目与细胞大小测定，样本量n = 540；C、D：16 DAA中果皮纵向切片观察；E、F：中果皮横向细胞数目与细胞大小测定，样本量n = 540；G、H：16 DAA中果皮横向切片观察：I：-3 DAA WT和Cssep2突变体果刺表型；J：-3 DAA WT和Cssep2突变体疣状结构表型；K：-3 DAA WT和Cssep2突变体果刺基部尺寸测量，TD：横向直径，LD：纵向直径；样本量n = 18；* P < 0.05；**** P < 0.000 1。数据以“均值 ± 标准差”表示。下同
Fig. 3 Cellular dissection of the fruit and wart phenotype of the Cssep2 mutantsA, B: Cell number and size of fruit mesocarp along the longitudinal axis. Values are means ± SD, n = 540. C, D: Longitudinal sections of the fruit mesocarp at 16 DAA. E, F: Cell number and size of fruit mesocarp along the transversal axis. Values are means ± SD, n = 540. G, H: Transversal sections of the fruit mesocarp at 16 DAA. I: Fruit spine of WT and Cssep2 mutants at -3 DAA. J: Macroscopic observation of the tubercule at -3 DAA. K: Measurement of the spine base of WT and Cssep2 mutants at -3 DAA. TD: Traverse diameter, LD: longitudinal diameter; * P < 0.05; **** P < 0.000 1. Values are means ± SD, n = 18. The same below
图4 差异表达基因的GO富集分析与RT-qPCR验证A：5T（5 DAA胎座）、16T（16 DAA胎座）、0W（0 DAA外果皮）和16W（16 DAA外果皮）样本中差异表达基因（DEGs）的GO富集分析结果；与DNA结合和转录调控相关的GO条目以红色高亮标注；B：WT和Cssep2突变体0 DAA外果皮组织中关键基因的表达分析；C：WT和Cssep2突变体5 DAA胎座组织中关键基因的表达分析；**** P < 0.000 1；** P < 0.01；* P < 0.05；ns：无显著差异
Fig. 4 GO enrichment and RT-qPCR analysis of differentially expressed genesA: GO enrichment analysis of DEGs at 5T (placenta at 5 DAA), 16T (placenta at 16 DAA), 0W (epicarp at 0 DAA) and 16W (epicarp at 16 DAA). GO terms related with DNA binding and regulation of transcription are highlighted with red. B: Expression analysis of genes in the epicarp tissues of WT and Cssep2 mutants. C: Expression analysis of genes in the placenta tissues of WT and Cssep2 mutants; **** P < 0.000 1; ** P < 0.01; * P < 0.05; ns: not significant difference
图5 与CsSEP2互作的蛋白鉴定A：酵母双杂交实验结果；QDO：四缺培养基（SD/-Leu-Trp-His-Ade），A600：终浓度为600 ng/mL的Aureobasidin A抗性筛选；B：萤火虫荧光素酶互补成像结果
Fig. 5 Identification of proteins interacting with CsSEP2A: Yeast two-hybrid assay results. QDO: Quadruple dropout, SD/-Leu-Trp-His-Ade. A600: Addition of Aureobasidin A with a final concentration of 600 ng/mL. B: Firefly luciferase complementation imaging analysis
图6 CsSEP2调控果实相关性状的分子网络A、B：CsSEP2参与调控果实发育（A）及果疣性状（B）的分子网络示意图。图例位于右上角。CsSEP2通过抑制CRABS CLAW的表达负调控黄瓜果实长度［17］；CsFUL通过抑制CsPIN1、CsPIN7和CsSUP的表达调控果实长度，且可与CsSEP2形成蛋白复合体发挥功能［21］。值得注意的是，Cssep2突变体中CsFUL和CsSUP的表达水平均发生显著变化。与WT相比，Cssep2突变体中CsGL3［22］、CsTS［23］、CsGL1［24］、CsTTG1［25］、CsTu［26-27］和ns［28］的表达水平下调，这些基因已被证实参与果刺与疣状结构的形成和发育调控。其中，ns基因编码生长素转运蛋白，负调控果刺密度［28］；CsGL1不仅抑制CsGA20OX1的表达，影响赤霉素合成并改变果刺密度和果刺基部大小［24］，还能与CsTTG1互作形成复合体调控果刺密度［25］；CsTu通过促进CsCTK［26］和CsTS1［27］的表达，调节细胞分裂素和生长素合成，进而促进果刺起始和疣状结构形成。在Cssep2突变体中，CsGA20OX1、CsCTK和CsTS1的表达水平也发生不同程度的改变。此外，本研究证实CsSEP2可与CsGL3、CsGL1、CsTTG1和CsTu发生蛋白互作
Fig. 6 Molecular network of fruit-related traits regulated by CsSEP2A, B: Schematic diagram of the CsSEP2-invovled molecular network of fruit development (A) and warty traits (B). The captions of the figure are in the upper right corner. CsSEP2 inhibits the lengths of cucumber fruits by suppressing the expression of CRABS CLAW[17]. CsFUL regulates fruit length by inhibiting the expressions of CsPIN1, CsPIN7 and CsSUP, and can also form protein complex with CsSEP2 to exert its functions[21]. Notably, the expressions of both CsFUL and CsSUP changed significantly in Cssep2. Compared to WT, the expressions of CsGL3[22], CsTS[23], CsGL1[24], CsTTG1[25], CsTu[26-27], and ns[28] were lower in Cssep2 mutants. These genes have been confirmed to be involved in the regulation of the formation and development of fruit spines and warts. Among them, ns encodes an auxin transporter and inhibits the density of fruit spines[28]. CsGL1 can not only inhibit the expression of CsGA20OX1, affecting the synthesis of GA and altering the density of fruit spines and the size of the fruit spine base[24], but also interacts with CsTTG1 to form a complex to regulate the density of fruit spines[25]. CsTu promotes the expressions of CsCTK[26] and CsTS1[27], altering the synthesis of cytokinin and auxin and facilitating the initiation of fruit spines and the formation of fruit warts. In Cssep2 mutants, the expressions of CsGA20OX1, CsCTK, and CsTS1 also changed to varying degrees. In addition, this study further revealed that CsSEP2 interacts with CsGL3, CsGL1, CsTTG,1 and CsTu
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