黄瓜果实表皮毛性状研究进展

doi:10.16420/j.issn.0513-353x.2024-0939

摘要/Abstract

摘要：

黄瓜是一种在全球范围内广泛种植的重要蔬菜作物，其不同部位均存在表皮毛。果实表面覆盖的非腺体型表皮毛称为果刺，与果瘤一并统称为刺瘤，对黄瓜表皮外观商品性状具有重要影响。刺瘤的大小及稀密程度除影响消费者的选购、栽培育种的偏好之外，还影响果实运输及贮藏、参与生物及非生物胁迫响应等。目前黄瓜中已鉴定了多个果实刺瘤调控基因并初步揭示了其作用机制。本文综述了黄瓜刺瘤发育相关基因的研究进展，总结了黄瓜表皮果刺系统发育的调控网络，并讨论了黄瓜表皮毛在逆境胁迫中的功能，为进一步培育出高商品性、高抗性的黄瓜新品种提供理论基础。

关键词: 黄瓜, 表皮毛, 果刺, 果瘤, 刺瘤

Abstract:

Cucumber（Cucumis sativus L.）is a globally significant vegetable crop characterized by the presence of trichomes on various plant organs. On the fruit surface，these non-glandular trichomes，known as spines，along with protuberances called tubercules，collectively form warts，which significantly influence fruit epidermal appearance and commercial value. The density and size of warts influence consumer acceptance and shape cultivation and breeding decisions，postharvest processes，and responses to both biotic and abiotic stresses. Numerous genes regulating spine and tubercule development have been identified in cucumber，and their underlying molecular mechanisms have been investigated. This review synthesizes current research on the genetic control of cucumber wart development，describes the regulatory networks governing spine ontogeny，and discusses the role of cucumber trichomes in stress responses. This knowledge provides a foundation for developing improved cucumber varieties with enhanced commercial traits and stress resistance.

Key words: cucumber, trichome, spine, tubercule, wart

柴兴文, 姚宏鑫, 郝宁, 王春华, 杜亚琳, 武涛, 曹嘉健. 黄瓜果实表皮毛性状研究进展[J]. 园艺学报, 2025, 52(12): 3386-3400.

CHAI Xingwen, YAO Hongxin, HAO Ning, WANG Chunhua, DU Yalin, WU Tao, CAO Jiajian. Research Progress of Fruit Trichome Traits in Cucumber[J]. Acta Horticulturae Sinica, 2025, 52(12): 3386-3400.

图/表 3

参考文献 79

[1]	Abdeldaym E A, Hassan H A, El-Mogy M M, Mohamed M S, Abuarab M E, Omar H S. 2024. Elevated concentrations of soil carbon dioxide with partial root-zone drying enhance drought tolerance and agro-physiological characteristics by regulating the expression of genes related to aquaporin and stress response in cucumber plants. BMC Plant Biology, 24(1):917.
[2]	Alonso A M, Espinosa J N, Carvajal M, Bárzana G. 2024. The differential expressions of aquaporins underline the diverse strategies of cucumber and tomato against salinity and zinc stress. Physiologia Plantarum, 176(1):e14222.
[3]	An J, Fang C, Yuan Z H, Hu Q, Huang W X, Li H Y, Ma R R, Wang L S, Su T, Li S C, Wang L D, Duan Y, Wang Y Q, Zhang C B, Xu R, Zhang D J, Cao Y M, Hou J J, Kong F J, Sun L J. 2023. A retrotransposon insertion in the Mao1 promoter results in erect pubescence and higher yield in soybean. Proceedings of the National Academy of Sciences of the United States of America, 120(13):e2210791120.
[4]	Chang J, Yu T, Yang Q H, Li C X, Xiong C, Gao S H, Xie Q M, Zheng F Y, Li H X, Tian Z D, Yang C X, Ye Z B. 2018. Hair,encoding a single C2H2 zinc-finger protein,regulates multicellular trichome formation in tomato. The Plant Journal, 96(1):96-102.
[5]	Chen C H, Liu M L, Jiang L, Liu X F, Zhao J Y, Yan S S, Yang S, Ren H Z, Liu R Y, Zhang X L. 2014. Transcriptome profiling reveals roles of meristem regulators and polarity genes during fruit trichome development in cucumber(Cucumis sativus L.). Journal of Experimental Botany, 65(17):4943-4958.
[6]	Chen C H, Yin S, Liu X W, Liu B, Yang S, Xue S D, Cai Y L, Kezia B, Liu H L, Dong M M, Zhang Y Q, Zhao B Y, Ren H Z. 2016. The WD-repeat protein CsTTG 1 regulates fruit wart formation through interaction with the homeodomain-leucine zipper I protein mict. Plant Physiology, 171(2):1156-1168.
[7]	Chen Yang. 2002. Study on physiological and ecological adaptability of halophytes in Xinjiang and the mechanism of Silicon enhancing plant salt tolerance[Ph. D. Dissertation]. Lanzhou: Gansu Agricultural University. (in Chinese)
	陈阳. 2002. 新疆盐生植物生理生态适应性及硅提高植物抗盐作用机制的研究[博士论文]. 兰州: 甘肃农业大学.
[8]	Chen Yucha. 2011. Relationship between nutrients and volatile substances and aphid resistance in cucumber[M. D. Dissertation]. Tai’an: Shandong Agricultural University. (in Chinese)
	陈玉茶. 2011. 营养物质和挥发性物质与黄瓜抗蚜的关系[硕士论文]. 泰安: 山东农业大学.
[9]	Cui J Y, Miao H, Ding L H, Todd C.W, Liu P N, Wang Y, Zhang S P, Gu X F. 2016. A new glabrous gene(csgl3)identified in trichome development in cucumber(Cucumis sativus L.). PLoS ONE, 11(2):e0148422.
[10]	Deng W, Yang Y W, Ren Z X, Corinne A D, Isabelle M, Wang X Y, Song H L, Hu Y H, Mondher B, Li Z G. 2012. The tomato SlIAA15 is involved in trichome formation and axillary shoot development. New Phytologist, 194(2):379-390.
[11]	Du H, Wang G, Pan J, Chen Y, Xiao T T, Zhang L Y, Zhang K Y, Wen H F, Xiong L R, Yu Y, He H L, Pan J S, Cai R. 2020. The HD-ZIP IV transcription factor Tril regulates fruit spine density through dose effects in cucumber. Journal of Experimental Botany, 71(20):6297-6310.
[12]	Ewas M, Gao Y Q, Wang S C, Liu X Q, Zhang H Y, Nishawy E M.E., Ali F, Shahzad R, Ziaf K, Subthain H, Martin C, Luo J. 2016. Manipulation of SlMXl for enhanced carotenoids accumulation and drought resistance in tomato. Science Bulletin, 61(18):1413-1418.
[13]	Feng Z X, Sun L, Dong M M, Fan S S, Shi K X, Qu Y X, Zhu L Y, Shi J F, Wang W J, Liu Y H, Song L Y, Weng Y Q, Liu X W, Ren H Z. 2023. Novel players in organogenesis and flavonoid biosynthesis in cucumber glandular trichomes. Plant Physiology, 192(4):2723-2736.
[14]	Guo C L, Yang X Q, Wang Y L, Nie J T, Yang Y, Sun J X, Du H, Zhu W Y, Pan J, Chen Y, Lv D, He H L, Lian H L, Pan J S, Cai R. 2017. Identification and mapping of ts(tender spines),a gene involved in soft spine development in Cucumis sativus. Theoretical and Applied Genetics, 131(1):1-12.
[15]	Guo C L, Yang X Q, Wang Y L, Nie J T, Yang Y, Sun J X, Du H, Zhu X Y, Pan J, Chen Y, Lv D, He H L, Lian H L, Pan J S, Cai R. 2023. TARGET OF EAT3(TOE3)specifically regulates fruit spine initiation in cucumber(Cucumis sativus L.). The Plant Journal, 115(3):678-689.
[16]	Hu B L, Wan Y, Li X, Zhang F T, Yan W G, Xie J K. 2013. Phenotypic characterization and genetic analysis of rice with pubescent leaves and glabrous hulls(PLgh). Crop Science, 53(5):1878-1886.
[17]	Hao H, Yao H X, Michio S, Li B H, Wang C H, Cao J J, Toru F, Wu T, Takehiro K. 2024. Novel lignin-based extracellular barrier in glandular trichome. Nature Plants, 10:381-389.
[18]	Hua B, Chang J, Wu M L, Xu Z J, Zhang F Y, Yang M N, Xu H M, Wang L J, Chen X Y, Wu S. 2020. Mediation of JA signalling in glandular trichomes by the woolly/SlMYC1 regulatory module improves pest resistance in tomato. Plant Biotechnology Journal, 19(2):375-393.
[19]	Jing W K, Li Y H, Zhang S, Zhou X F, Gao J P, Ma N. 2022. Aquaporin,beyond a transporter. Horticultural Plant Journal, 9(1):29-34.
[20]	Ji Xiuzhi, Zhang Qingwen, Liu Xiaoxia, Yang Maisheng. 2005. Effects of different cucumber varieties on adult and egg laying of Whitefly tabaci//The 5th International Symposium on High and New Science and Technology for Biodiversity Conservation and Utilization and International Symposium on Insect Conservation,Utilization and Industrialization. Beijing. (in Chinese)
	姬秀枝, 张青文, 刘小侠, 杨麦生. 2005. 不同黄瓜品种对烟粉虱成虫及产卵的影响//第五届生物多样性保护与利用高新科学技术国际研讨会暨昆虫保护、利用与产业化国际研讨会. 北京.
[21]	Kim H J, Han J H, Kwon J K, Park M, Kim B D, Choi D. 2010. Fine mapping of pepper trichome locus 1 controlling trichome formation in Capsicum annuum L. CM334. Theoretical and Applied Genetics, 120(6):1099-1106.
[22]	Kong Xiangxin, Liu Yuanyuan, Yang Weinan, He Meijun, Jin Yongling. 2020. Effects of leaf morphological characteristics of different cucumber varieties on preference of white whitefly in greenhouse. China Plant Protection Guide, 40(1):20-24. (in Chinese)
	孔祥鑫, 刘媛媛, 杨伟男, 何美君, 金永玲. 2020. 不同黄瓜品种叶片形态特征对温室白粉虱偏好的影响. 中国植保导刊, 40(1):20-24.
[23]	Lei Guanhong. 2013. Physiological mechanism of anti-aphid in hairless cucumber[M. D. Dissertation]. Tai’an: Shandong Agricultural University. (in Chinese)
	雷关红. 2013. 无毛黄瓜抗蚜生理机理探究[硕士论文]. 泰安: 山东农业大学.
[24]	Li L, Zhao Y F, McCaig B C, Wingerd B A, Wang J H, Whalon M E, Pichersky E, Howe G A. 2003. The tomato homolog of CORONATINE-INSENSITIVE 1 is required for the maternal control of seed maturation,jasmonate-signaled defense responses,and glandular trichome development. the Plant Cell, 16(1):126-143.
[25]	Li Q, Cao C X, Zhang C J, Zheng S S, Wang Z H, Wang L N, Ren Z H. 2015. The identification of Cucumis sativus Glabrous 1(CsGL1)required for the formation of trichomes uncovers a novel function for the homeodomain-leucine zipper I gene. Journal of Experimental Botany, 66(9):2515-2526.
[26]	Li Tiantian, Jiang Wenzhi, Gu Tingting, Liu Yongyue, Xu Jianpeng, Liu Guijun, Cao Chenxing. 2016. Study on the relationship between nutrients and aphid resistance in cucumber leaves. Shandong Agricultural Sciences, 48(10):44-47,50. (in Chinese)
	李田田, 姜文芝, 谷停停, 刘永月, 许建鹏, 刘桂军, 曹辰兴. 2016. 黄瓜叶片营养物质与抗蚜性的关系研究. 山东农业科学, 48(10):44-47,50.
[27]	Liu J, Cui J J, Dong J C, Zhong J, Zhong C F, Yuan F C, Guan W D, Hu F, Cheng J W, Hu K L. 2023. A 1-bp deletion in the MC04g1399 is highly associated with failure to produce fruit wart in bitter gourd. Horticultural Plant Journal, 10(1):171-180.
[28]	Liu Jinqiu. 2017. The role of SlHZ45 gene in skin and fur development in tomato[M. D. Dissertation]. Harbin: Northeast Agricultural University. (in Chinese)
	刘金秋. 2017. 番茄SlHZ45基因在表皮毛发育中的角色[硕士论文]. 哈尔滨: 东北农业大学.
[29]	Liu Lei, Sun Jian, Wu Dianxing, Shu Xiaoli. 2015. Research progress on photophyllleaf traits and epidermal fur development of rice. Journal of Nuclear Agronomy, 29(11):7. (in Chinese)
	柳蕾, 孙健, 吴殿星, 舒小丽. 2015. 水稻光叶性状及表皮毛发育研究进展. 核农学报, 29(11):7.
[30]	Liu M Y, Zhang C J, Duan L X, Luan Q Q, Li J L, Yang A G, Qi X Q, Ren Z H. 2019. CsMYB60 is a key regulator of flavonols and proanthocyanidans that determine the colour of fruit spines in cucumber. Journal of Experimental Botany, 70(1):69-84.
[31]	Liu W, Fu R, Li Q, Li J, Wang L N, Ren Z H. 2013. Genome-wide identification and expression profile of homeodomain-leucine zipper Class I gene family in Cucumis sativus. Gene, 531(2):279-287.
[32]	Liu X P, Dong S Y, Miao H, Bo K L, Li C X, Yang Y Y, Gu X F, Zhang S P. 2021. Genome-wide analysis of expansins and their role in fruit spine. Horticultural Plant Journal, 8(6):757-768.
[33]	Liu X P, Yang X Y, Xie Q, Miao H, Bo K L, Dong S Y, Xin T X, Gu X F, Sun J Q, Zhang S P. 2022. NS encodes an auxin transporter that regulates the‘numerous spines’trait in cucumber(Cucumis sativus)fruit. The Plant Journal, 110(2):325-336.
[34]	Liu Yongli, Song Tiefeng, Zhao Juyong. 2013. Evaluation of sexual type and economic characteristics of germplasm resources in cucumber. Horticulture and Seedlings,(9):22-25. (in Chinese)
	刘永丽, 宋铁峰, 赵聚勇. 2013. 黄瓜种质资源的性型及经济学性状评价. 园艺与种苗,(9):22-25.
[35]	Liu Yongyue. 2016. Relationship between the number of hairless(gl)genes and aphid resistance in cucumber[M. D. Dissertation]. Tai’an: Shandong Agricultural University. (in Chinese)
	刘永月. 2016. 黄瓜无毛(gl)基因数量与抗蚜的关系[硕士论文]. 泰安: 山东农业大学.
[36]	Machado A, Wu Y R, Yang Y M, Llewellyn D J, Dennis E S. 2009. The MYB transcription factor GhMYB25 regulates early fibre and trichome development. The Plant Journal, 59(1):52-62.
[37]	Martin C, Bhatt K, Baumann K, Jin H L, Zachgo S, Roberts K, Sommer Z S, Glover B, Rodrigues M P. 2002. The mechanics of cell fate determination in petals. Philosophical Transactions of the Royal Society of London, 357(1422):809-813.
[38]	Nadakuduti S S, Pollard M, Kosma D K, Allen C, Ohlrogge J B, Barry C S. 2012. Pleiotropic phenotypes of the sticky peel mutant provide new insight into the role of CUTIN DEFICIENT 2 in epidermal cell function in tomato. Plant Physiology, 159(3):945-960.
[39]	Pan J, Zhang L Y, Chen G Q, Wen H F, Chen Y, Du H, Zhao J L, He H L, Lian H L, Chen H M, Shi J X, Cai R, Wang G, Pan J S. 2021. Study of micro-trichome(mict)reveals novel connections between transcriptional regulation of multicellular trichome development and specific metabolism in cucumber. Horticulture Research, 8(2):21.
[40]	Pan Y P, Bo K, Cheng Z H, Weng Y Q. 2015. The loss-of-function GLABROUS 3mutation in cucumber is due to LTR-retrotransposon insertion in a class IV HD-ZIP transcription factor gene CsGL3 that is epistatic over CsGL1. Bmc Plant Biology, 15(1):302.
[41]	Pattanaik S, Patra B, Singh S K, Yuan L. 2014. An overview of the gene regulatory network controlling trichorne development in the model plant,Arabidopsis. Front Plant Sci, 5:259.
[42]	Peng Jialin, Nie Jingtao, Qu Meilin, Pan Junsong, He Huanle. 2015. Mapping and candidate gene diversity of nigra thorn in cucumber fruit. Journal of Shanghai Jiaotong University, 33(4):12-19. (in Chinese)
	彭佳林, 聂京涛, 曲美玲, 潘俊松, 何欢乐. 2015. 黄瓜果实黑刺基因的定位及候选基因多样性研究. 上海交通大学学报, 33(4):12-19.
[43]	Pu Li, Suo Jinfeng, Xue Yongbiao. 2003. Molecular genetic control of plant skin and fur development. Journal of Genetics,(11):1078-1084. (in Chinese)
	普莉, 索金凤, 薛勇彪. 2003. 植物表皮毛发育的分子遗传控制. 遗传学报,(11):1078-1084.
[44]	Simmons A T, Gurr G M. 2010. Trichomes of Lycopersicon species and their hybrids:effects on pests and natural enemies. Agricultural and Forest Entomology, 7(4):265-276.
[45]	Sun C, Wei J B, Gu X Y, Wu M L, Li M, Liu Y X, An N K, Wu K M, Wu S S, Wu J Q, Xu M Z, Wu J C, Wang Y L, Chao D Y, Zhang Y J, Wu S. 2024. Different multicellular trichome types coordinate herbivore mechanosensing and defense in tomato. the Plant Cell, 36(12):4952-4969.
[46]	Shi J, Wang J F, Li R, Li D B, Xu F F, Sun Q Q, Zhao B, Mao A J, Guo Y D. 2015. Expression patterns of genes encoding plasma membrane aquaporins during fruit development in cucumber(Cucumis sativus L.). Plant Physiology and Biochemistry, 96(8):329-336.
[47]	Sun Qing, Yang Yan, Luo Donglan, Ba Liangjie. 2018. Research progress for technology of cucumber storage and fresh-keeping after picking. South China Agriculture, 12(34):54-55,61. (in Chinese)
	孙情, 杨炎, 罗冬兰, 巴良杰. 2018. 黄瓜采后贮藏保鲜技术研究进展. 南方农业, 12(34):54-55,61.
[48]	Suo J F, Liang X E, Pu L, Zhang Y S, Xue Y B. 2003. Identification of GhMYB109 encoding a R2R3 MYB transcription factor that expressed specifically in fiber initials and elongating fibers of cotton(Gossypium hirsutum L.). Biochimica et Biophysica Acta(BBA)- Gene Structure and Expression, 1630(1):25-34.
[49]	Tiwari P. 2016. Recent advances and challenges in trichome research and essential oil biosynthesis in Mentha arvensis L. Industrial Crops and Products, 82:141-148.
[50]	Wang S, Wang J W, Yu N, Li C H, Luo B, Gou J Y, Wang L J, Chen X Y. 2004. Control of plant trichome development by a cotton fiber MYB gene. The Plant Cell, 16(9):2323-2334.
[51]	Wang Y L, Nie J T, Chen H M, Guo C L, Pan J, He H L, Pan J S, Cai R. 2015. Identification and mapping of Tril,a homeodomain-leucine zipper gene involved in multicellular trichome initiation in Cucumis sativus. Theoretical and Applied Genetics, 129(2):305-316.
[52]	Wang Yujue, Fu Qiushi, Zheng He, Wen Changlong, Cheng Lin, Zhao Bing, Guo Yangdong. 2010. Effects of drought stress on growth,photosynthetic physiology and stomatal characteristics of cucumber seedlings. Journal of China Agricultural University, 15(5):12-18. (in Chinese)
	王玉珏, 付秋实, 郑禾, 温常龙, 程琳, 赵冰, 郭仰东. 2010. 干旱胁迫对黄瓜幼苗生长、光合生理及气孔特征的影响. 中国农业大学学报, 15(5):12-18.
[53]	Wang Z Y, Wang L M, Han L J, Cheng Z H, Liu X F, Wang S Y, Liu L, Chen J C, Song W Y, Zhao J Y, Zhou Z Y, Zhang X L. 2021. HECATE 2 acts with GLABROUS3 and Tu to boost cytokinin biosynthesis and regulate cucumber fruit wart formation. Plant Physiology, 187(3):1619-1635.
[54]	Xie Q M, Gao Y N, Li J, Yang Q H, Qu X L, Li H X, Zhang J H, Wang T T, Ye Z B, Yang C X. 2020. The HD-Zip IV transcription factor SlHDZIV8 controls multicellular trichome morphology by regulating the expression of Hairless-2. Journal of Experimental Botany, 71(22):7132-7145.
[55]	Xie Q M, Xiong C, Yang Q H, Zheng F Y, Larkin R M, Zhang J H, Wang T T, Zhang Y Y, Bo O Y, Lu Y E, Ye J, Ye Z B, Yang C X. 2022. A novel regulatory complex mediated by Lanata(Ln)controls multicellular trichome formation in tomato. New Phytologist, 236(6):2294-2310.
[56]	Xu Jianpeng. 2016. Screening of cucumber anti-whitefly materials and research on resistance mechanism to whitefly[M. D. Dissertation]. Tai’an: Shandong Agricultural University. (in Chinese)
	许建鹏. 2016. 黄瓜抗粉虱材料筛选及其对烟粉虱抗性机理的研究[硕士论文]. 泰安: 山东农业大学.
[57]	Xu J S, Herwijnen Z O, Dräger D B, Sui C, Haring M A, Schuurink R C. 2018. SlMYC1 regulates type VI glandular trichome formation and terpene biosynthesis in tomato glandular cells. the Plant Cell, 30(12):2988-3005.
[58]	Xu S, Wang Y R, Yang S L, Fan S S, Shi K X, Wang F, An M H, Qi Y, Wang M Q, Feng M, Li Z F, Liu X W, Ren H Z. 2024. The WUSCHEL-related homeobox transcription factor CsWOX3 negatively regulates fruit spine morphogenesis in cucumber(Cucumis sativus L.). Horticulture Research, 11(8):uhae163.
[59]	Xue S D, Dong M M, Liu X W, Xu S, Pang J A, Zhang W Z, Weng Y Q, Ren H Z. 2018. Classification of fruit trichomes in cucumber and effects of plant hormones on type Ⅱ fruit trichome development. Planta, 249:407-416.
[60]	Yan An. 2010. Effects of enhanced UV-B radiation and salt stress on Arabidopsis epidermis mutants[M. D. Dissertation]. Lanzhou: Lanzhou University. (in Chinese)
	颜安. 2010. 增强UV-B辐射及盐胁迫对拟南芥表皮毛突变体的影响[硕士论文]. 兰州: 兰州大学.
[61]	Yan S S, Ning K, Wang Z Y, Liu X F, Zhong Y T, Ding L, Zi H L, Cheng Z H, Li X X, Shan H Y, Lv Q Y, Luo L X, Liu R Y, Yan L Y, Zhou Z Y, Lucas W J, Zhang X L. 2020. CsIVP functions in vasculature development and downy mildew resistance in cucumber. PLos Biology, 18(3):e3000671.
[62]	Yang C X, Li H X, Zhang J H, Luo Z D, Gong P J, Zhang C J, Li J H, Wang T T, Zhang Y Y, Lu Y E, Ye Z B. 2011. A regulatory gene induces trichome formation and embryo lethality in tomato. Proceedings of the National Academy of Sciences of the United States of America, 108(29):11836-11841.
[63]	Yang S, Cai Y L, Liu X W, Dong M M, Zhang Y Q, Chen S Y, Zhang W B, Li Y J, Tang M, Zhai X L, Weng Y Q, Ren H Z. 2018. A CsMYB6-CsTRY module regulates fruit trichome initiation in cucumber. Journal of Experimental Botany, 69(8):1887-1902.
[64]	Yang S, Wang Y L, Zhu H Y, Zhang M J, Wang D K, Xie K X, Fan P F, Dou J L, Liu D M, Liu B, Chen C H, Yan Y, Zhao L J, Yang L M. 2022. A novel HD-Zip I/C2H2-ZFP/WD-repeat complex regulates the size of spine base in cucumber. New Phytologist, 233(6):2643-2658.
[65]	Yang S L, Xue S D, Shan L, Fan S S, Sun L, Dong Y M, Li S, Gao M Y, Qi Y, Yang L, An M H, Wang F, Pang J A, Zhang W Z, Weng Y Q, Liu X W, Ren H Z. 2024. The CsTM alters multicellular trichome morphology and enhances resistance against aphid by interacting with CsTIP1;1 in cucumber. Journal of Advanced Research, 69:17-30.
[66]	Yang X Q, Zhang W W, He H L, Nie J T, Bie B B, Zhao J L, Ren G L, Li Y, Zhang D B, Pan J S, Cai R. 2014. Tuberculate fruit gene Tu encodes a C2H 2 zinc finger protein that is required for the warty fruit phenotype in cucumber(Cucumis sativus L.). The Plant Journal, 78(6):1034-1046.
[67]	Yang Z G, Song M F, Cheng F, Zhang M R, Davoudi M, Chen J F, Lou Q F. 2021. A SNP mutation in homeodomain-DDT(HD-DDT)transcription factor results in Multiple Trichomes(mt)in cucumber(Cucumis sativus L.). Genes, 12(10):1478.
[68]	Ye X L, Hu F Y, Ren J, Huang S N, Liu W J, Feng H, Liu Z Y. 2016. Fine mapping and candidate gene analysis of Brtri1,a gene controlling trichome development in Chinese cabbage(Brassica rapa L.ssp pekinensis). Genetics and Molecular Research, 15(4):doi:10.4238/gmr15048924.
[69]	Yin S, Zhao L L, Liu J Q, Sun Y J, Li B H, Wang L N, Ren Z H, Chen C H. 2023. Pan-genome analysis of WOX gene family and function exploration of CsWOX9 in cucumber. International Journal of Molecular Sciences, 24(24):17568.
[70]	Yu X H, Chen G P, Tang B Y, Zhang J L, Zhou S G, Hu Z L. 2018. The jasmonate ZIM-domain protein gene SlJAZ2 regulates plant morphology and accelerates flower initiation in Solanum lycopersicum plants. Plant Science, 267:65-73.
[71]	Zhang C Y, Win K T, Kim Y C, Lee S. 2019. Two types of mutations in the HEUKCHEEM gene functioning in cucumber spine color development can be used as signatures for cucumber domestication. Planta, 250:1491-1504.
[72]	Zhang H Y, Wang L N, Zheng S S, Liu Z Z, Wu X Q, Gao Z H, Cao C X, Li Q, Ren Z H. 2016. A fragment substitution in the promoter of CsHDZIV11/CsGL3 is responsible for fruit spine density in cucumber(Cucumis sativus L.). Theoretical and Applied Genetics, 129:1289-1301.
[73]	Zhang Haiyang. 2016. Localization,cloning and functional analysis of thin thorn gene in cucumber[Ph. D. Dissertation]. Tai’an: Shandong Agricultural University. (in Chinese)
	张海洋. 2016. 黄瓜稀刺基因的定位、克隆及功能分析[博士论文]. 泰安: 山东农业大学.
[74]	Zhang W W, Chen Y, Zhou P, Bao W M, Yang X Q, Xu T B, She W W, Xu L Q, Yu P G, Pan J S. 2018. Identification and fine mapping of molecular markers closely linked to fruit spines size ss gene in cucumber(Cucumis sativus L.). Euphytica, 214:213.
[75]	Zhang Y Q, Shen J J, Bartholomew E S, Dong M M, Chen S Y, Yin S, Zhai X L, Feng Z X, Ren H Z, Liu X W. 2021. TINY BRANCHED HAIR functions in multicellular trichome development through an ethylene pathway in Cucumis sativus L. The Plant Journal, 106(3):753-765.
[76]	Zhao L J, Fan P F, Wang Y L, Xu N N, Zhang M J, Chen M Y, Zhang M Y, Dou J L, Liu D M, Niu H H, Zhu H Y, Hu J B, Sun S R, Yang L M, Yang S. 2024. ELONGATED HYPOTCOTYL5 and SPINE BASE SIZE 1 together mediate light-regulated spine expansion in cucumber. Plant Physiology, 195(1):552-565.
[77]	Zhao J L, Pan J S, Guan Y, Zhang W W, Bie B B, Wang Y L, He H L, Lian H L, Cai R. 2015. Micro-trichome as a class I homeodomain-leucine zipper gene regulates multicellular trichome development in Cucumis sativus. Journal of Integrative Plant Biology, 57(11):925-935.
[78]	Zhao L J, Zhu H Y, Zhang K G, Wang Y L, Wu L, Chen C H, Liu X W, Yang S, Ren H Z, Yang L M. 2020. The MIXTA-LIKE transcription factor CsMYB6 regulates fruit spine and tubercule formation in cucumber. Plant Science, 300:110636.
[79]	Zhou Xiujie. 2007. Study on the mechanism of silicon promoting drought resistance in cucumber seedlings[M. D. Dissertation]. Tianjin: Tianjin Normal University. (in Chinese)
	周秀杰. 2007. 硅促进黄瓜幼苗抗旱作用机制研究[硕士论文]. 天津: 天津师范大学.

基因名 Gene name	基因号 Gene number	编码蛋白 Coding protein	突变体表型 Mutant phenotype	参考文献 Reference
CsGL3	Csa6G514870	HD-ZIP IV transcription factor	光滑无毛Glabrous	Pan et al.,2015 Cui et al.,2016
TRIL	Csa6G514870	HD-ZIP IV transcription factor	无毛Trichome-less	Wang et al.,2015
FS1	Csa6G514870	HD-ZIP IV transcription factor	稀刺Few spines	Zhang et al.,2016
CsGL1	Csa3G748220	HD-ZIP I transcription factor	下胚轴有毛状体，地上部存在未发育成熟表皮毛 The hypocotyl has trichomes，and the aerial parts have undeveloped trichomes	Li et al.,2015
MICT	Csa3G748220	HD-ZIP I transcription factor	微毛Micro-trichome	Zhao et al.,2015
CsTBH	Csa3G748220	HD-ZIP I transcription factor	小分支毛Tiny branched hair	Chen et al.,2014 Zhang et al.,2021
CsTOE3	CsaV3_6G043390	AP2/ERF transcription factor	无刺瘤Spineless	Guo ea al.,2023
NS	Csa2G264590	Auxin transport protein	多毛Numerous spines	Liu et al.,2022
MT	CsaV3_6G050410	HD-DDT protein	多毛Multiple trichomes	Yang et al.,2021
CsMYB6	Csa3G824850	R2R3-MYB transcription factor	过表达使果刺减少 Overexpression reduces fruit spines	Yang et al.,2018
CsGA20ox1	Csa5g172270	Gibberellin oxidase	过表达使果刺变短 Overexpression causes the fruit spine to shorten	Liu et al.,2013
CsTRY	Csa5G139610	R2R3-MYB transcription factor	过表达使果刺减少 Overexpression reduces fruit spines	Yang et al.,2018
CsTTG1	Csa4Gm097650	WD-repeat transcription factor	过表达使果刺增加 Overexpression increases fruit spines	Chen et al.,2016
CsHEC2	Csa2G285890	bHLH transcription factor	刺瘤密度降低Reduced wart density	Wang et al.,2021
CsSBS1	Csa5G577350	C2H2 zinc finger transcription factor	小刺基Small spine base	Yang et al.,2022
CsHY5	CsaV3_2G031960	bZIP transcription factor	小刺基Small spine base	Zhao et al.,2024
CsWOX3	Csa6G301060	HOMEODOMAIN domain and WUS domain	大刺基Big spine base	Xu et al.,2024
CsTM	Csa1G056960	C-lectin receptor-like kinase	小刺基Small spine base	Yang et al.,2024
B2	Csa4G003095	R2R3-MYB transcription factor	黑刺Black spine	彭佳林等,2015
TS	Csa1G056960	C-lectin receptor-like kinase	嫩刺Tender spines	Guo et al.,2017
Tu	Csa016861	C2H2 zinc finger transcription factor	无果瘤Non-warty	Yang et al.,2014

基因名 Gene name	基因号 Gene number	编码蛋白 Coding protein	突变体表型 Mutant phenotype	参考文献 Reference
CsGL3	Csa6G514870	HD-ZIP IV transcription factor	光滑无毛Glabrous	Pan et al.,2015 Cui et al.,2016
TRIL	Csa6G514870	HD-ZIP IV transcription factor	无毛Trichome-less	Wang et al.,2015
FS1	Csa6G514870	HD-ZIP IV transcription factor	稀刺Few spines	Zhang et al.,2016
CsGL1	Csa3G748220	HD-ZIP I transcription factor	下胚轴有毛状体，地上部存在未发育成熟表皮毛 The hypocotyl has trichomes，and the aerial parts have undeveloped trichomes	Li et al.,2015
MICT	Csa3G748220	HD-ZIP I transcription factor	微毛Micro-trichome	Zhao et al.,2015
CsTBH	Csa3G748220	HD-ZIP I transcription factor	小分支毛Tiny branched hair	Chen et al.,2014 Zhang et al.,2021
CsTOE3	CsaV3_6G043390	AP2/ERF transcription factor	无刺瘤Spineless	Guo ea al.,2023
NS	Csa2G264590	Auxin transport protein	多毛Numerous spines	Liu et al.,2022
MT	CsaV3_6G050410	HD-DDT protein	多毛Multiple trichomes	Yang et al.,2021
CsMYB6	Csa3G824850	R2R3-MYB transcription factor	过表达使果刺减少 Overexpression reduces fruit spines	Yang et al.,2018
CsGA20ox1	Csa5g172270	Gibberellin oxidase	过表达使果刺变短 Overexpression causes the fruit spine to shorten	Liu et al.,2013
CsTRY	Csa5G139610	R2R3-MYB transcription factor	过表达使果刺减少 Overexpression reduces fruit spines	Yang et al.,2018
CsTTG1	Csa4Gm097650	WD-repeat transcription factor	过表达使果刺增加 Overexpression increases fruit spines	Chen et al.,2016
CsHEC2	Csa2G285890	bHLH transcription factor	刺瘤密度降低Reduced wart density	Wang et al.,2021
CsSBS1	Csa5G577350	C2H2 zinc finger transcription factor	小刺基Small spine base	Yang et al.,2022
CsHY5	CsaV3_2G031960	bZIP transcription factor	小刺基Small spine base	Zhao et al.,2024
CsWOX3	Csa6G301060	HOMEODOMAIN domain and WUS domain	大刺基Big spine base	Xu et al.,2024
CsTM	Csa1G056960	C-lectin receptor-like kinase	小刺基Small spine base	Yang et al.,2024
B2	Csa4G003095	R2R3-MYB transcription factor	黑刺Black spine	彭佳林等,2015
TS	Csa1G056960	C-lectin receptor-like kinase	嫩刺Tender spines	Guo et al.,2017
Tu	Csa016861	C2H2 zinc finger transcription factor	无果瘤Non-warty	Yang et al.,2014