柑橘单性结实研究进展

doi:10.16420/j.issn.0513-353x.2020-0881

摘要/Abstract

摘要：

单性结实是柑橘无核的重要原因。对柑橘属种质资源单性结实能力、激素调控柑橘单性结实果实发育机理、无核基因及基因工程无核育种进展予以综述,同时总结了其他园艺植物单性结实最新研究技术,以期为单性结实研究提供参考。

Abstract:

Parthenocarpy is an important reason for citrus to produce seedless fruit. The parthenocarpy ability of citrus germplasm resources,mechanism of hormones regulating development of parthenocarpy fruit,the progress of seedless genes and genetic engineering seedless breeding are reviewed. Meanwhile,the latest research techniques on parthenocarpy of other horticultural plants are summarized,which may provide reference for parthenocarpy research.

Key words: citrus, parthenocarpy, seedless, hormone, male sterility

中图分类号:

S666

赵科科, 崔璐璐, 李春秀, 党江波, 梁国鲁, 向素琼. 柑橘单性结实研究进展[J]. 园艺学报, 2021, 48(4): 811-824.

ZHAO Keke, CUI Lulu, LI Chunxiu, DANG Jiangbo, LIANG Guolu, XIANG Suqiong. Research Advances on Parthenocarpy in Citrus[J]. Acta Horticulturae Sinica, 2021, 48(4): 811-824.

参考文献 92

[1]	Aarts M, Hodge R, Kalantidis K, Florack D, Wilson Z, Mulligan B, Stiekema W Scott R Pereira A. 1997. The Arabidopsis MALE STERILITY 2 protein shares similarity with reductases in elongation/condensation complexes. The Plant Journal, 12 (3):615-623. doi: 10.1046/j.1365-313X.1997.00615.x URL
[2]	Abouzari A, Nezhad N M. 2016. The investigation of citrus fruit quality. popular characteristic and breeding. Acta Universitatis Agriculturae et Silviculturae Mendelianae Brunensis, 64 (3):725-740. doi: 10.11118/actaun201664030725 URL
[3]	Acciarri N, Restaino F, Vitelli G, Perrone D, Zottini M, Pandolfini T, Spena A, Rotino G L. 2002. Genetically modified parthenocarpic eggplants:improved fruit productivity under both greenhouse and open field cultivation. BMC Biotechnology, 2:4. doi: 10.1186/1472-6750-2-4 URL
[4]	Baumbach J, Pudake R N, Johnson C, Kleinhans K, Ollhoff A, Palmer R G, Bhattacharyya M K, Sandhu D. 2016. Transposon tagging of a male-sterility,female-sterility gene,St8,revealed that the meiotic MER3 DNA helicase activity is essential for fertility in soybean. PLoS ONE, 11 (3):e0150482. doi: 10.1371/journal.pone.0150482 URL
[5]	Chen L, Luo X, Yang X, Jing D, Xia X, Li H, Poudel K, Cao S. 2020. Small RNA and mRNA sequencing reveal the roles of micrornas involved in pomegranate female sterility. International Journal of Molecular Sciences, 21 (2):558. doi: 10.3390/ijms21020558 URL
[6]	Chen X, Zhang M, Tan J, Huang S, Wang C, Zhang H, Tan T. 2017. Comparative transcriptome analysis provides insights into molecular mechanisms for parthenocarpic fruit development in eggplant(Solanum melongena L.). PloS ONE, 12 (6):e0179491. doi: 10.1371/journal.pone.0179491 URL
[7]	Chen Jun-wei, Feng Jian-jun, Qin Qia-ping, Liu Xiao-kun, Wu Jiang, Xie Ming. 2006. Characteristics of sugar metabolism and accumulation in GA ₃ induced parthenocarpic white flesh loquat‘Ninghai Bai’Fruit. Acta Horticulturae Sinica, 33 (3):471-476. (in Chinese)
	陈俊伟, 冯健君, 秦巧平, 刘晓坤, 吴江, 谢鸣. 2006. GA ₃诱导的单性结实‘宁海白’白沙枇杷糖代谢的研究. 园艺学报, 33 (3):471-476.
[8]	Chai P, Dong S, Chai L, Chen S, Flaishman M, Ma H. 2019. Cytokinin-induced parthenocarpy of San Pedro type fig( Ficus carica L.)main crop:explained by phytohormone assay and transcriptomic network comparison. Plant Molecular Biology, 99 (4-5):329-346.
[9]	Caruso M, Merelo P, Distefano G, Malfa S L, Piero A R L, Tadeo F R, Talon M, Gentile A. 2012. Comparative transcriptome analysis of stylar canal cells identifies novel candidate genes implicated in the self-incompatibility response of Citrus clementina. BMC Plant Biology, 12:20. doi: 10.1186/1471-2229-12-20 pmid: 22333138
[10]	Cong L, Yue R, Wang H, Liu J L, Zhai R, Yang J, Wu M, Si M, Zhang H Q, Yang C Q, Xu L F, Wang Z G. 2018. 2,4-D-induced parthenocarpy in pear is mediated by enhancement of GA ₄ biosynthesis. Physiologia Plantarum, 166 (3):812-820. doi: 10.1111/ppl.2019.166.issue-3 URL
[11]	de Jong M, Wolters-Arts M, Feron R, Mariani C, Vriezen W H. 2009. The solanum lycopersicum auxin response factor 7(slarf7)regulates auxin signaling during tomato fruit set and development. The Plant Journal, 57 (1):160-170. doi: 10.1111/tpj.2008.57.issue-1 URL
[12]	Dexter-Boone A, Humphry M, Shi R, Lewis R S. 2019. Genetic control of facultative parthenocarpy in Nicotiana tabacum L. The Journal of Heredity, 110 (5):610-617. doi: 10.1093/jhered/esz025 pmid: 31002335
[13]	Dong Dian-qian, Gong Gui-zhi, Peng Zhu-chun, Hou Yan-hong, Luo Ai, Hong Qi-bin. 2020. Analysis on the relationship between W-murcott tangor parthenocarpy ability and content of 4 endogenous hormone in different parts in cirtus. Molecular Plant Breeding, 18 (4):1326-1337. (in Chinese)
	董倩倩, 龚桂芝, 彭祝春, 侯艳红, 罗艾, 洪棋斌. 2020. W-默科特单性结实能力与果实不同部位四种内源激素含量的关系分析. 分子植物育种, 18 (4):1326-1337.
[14]	Ding J, Chen B, Xia X J, Mao W H, Shi K, Zhou Y H, Yu J Q. 2013. Cytokinin-induced parthenocarpic fruit development in tomato is partly dependent on enhanced gibberellin and auxin biosynthesis. PLoS ONE, 8 (7):e70080. doi: 10.1371/journal.pone.0070080 URL
[15]	Distefano G, Gentile A. 2011. Pollen-pistil interactions and early fruiting in parthenocarpic citrus. Annals of Botany, 108 (3):499-509. doi: 10.1093/aob/mcr187 URL pmid: 21795277
[16]	Ficcadenti N, Sestili S, Pandolfini T, Cirillo C, Rotino G L, Spena A. 1999. Genetic engineering of parthenocarpic fruit development in tomato. Molecular Breeding, 5:463-470. doi: 10.1023/A:1009665409959 URL
[17]	Gambetta G, Gravina A, Fasiolo C, Fornero C, Galiger S, Inzaurralde C, Rey F. 2013. Self-incompatibility,parthenocarpy and reduction of seed presence in‘Afourer’mandarin. Scientia Horticulturae, 164 (17):182-188.
[18]	Garmendia A, Beltrán R, Zornoza C, Breijo F, Reig J, Bayona I, Merle H. 2019. Insect repellent and chemical agronomic treatments to reduce seed number in‘Afourer’mandarin. Effect on yield and fruit diameter. Scientia Horticulturae, 246:437-447. doi: 10.1016/j.scienta.2018.11.025
[19]	García-Hurtado N, Carrera E, Ruiz-Rivero O, López-Gresa M P, Hedden P, Gong F, García-Martínez J L. 2012. The characterization of transgenic tomato overexpressing gibberellin 20-oxidase reveals induction of parthenocarpic fruit growth,higher yield,and alteration of the gibberellin biosynthetic pathway. Journal of Experimental Botany, 63 (16):5803-5813. doi: 10.1093/jxb/ers229 URL pmid: 22945942
[20]	George P, Rédei . 2008. Navel Oranges// Encyclopedia of genetics,genomics, proteomics and informatics. Netherlands Springer.
[21]	Goto S, Yoshioka T, Ohta S, Kita M, Hamada H, Shimizu T. 2018. QTL mapping of male sterility and transmission pattern in progeny of Satsuma mandarin. PLoS ONE, 13 (7):e0200844. doi: 10.1371/journal.pone.0200844 URL
[22]	Goetz M, Hooper L C, Johnson S D, Rodrigues J C M, Vivian-Smith A, Koltunow A M. 2007. Expression of aberrant forms of auxin response factor 8 stimulates parthenocarpy in Arabidopsis and tomato. Plant Physiology, 145 (2):351-366. doi: 10.1104/pp.107.104174 URL
[23]	Guo Rong-yu. 2015. Inheritance analysis and QTL mapping of parthenocarpy in cucumber[M. D. Dissertation]. Qinhuangdao:Hebei Normal University of Science &Technology. (in Chinese)
	郭荣雨. 2015. 旱黄瓜单性结实性遗传分析及主效QTL定位[硕士论文]. 秦皇岛:河北科技师范学院.
[24]	Gustafson F G. 1939. The cause of natural parthenocarpy. American Journal of Botany, 26 (3):135-138. doi: 10.1002/ajb2.1939.26.issue-3 URL
[25]	Gustafson F G. 1942. Parthenocarpy:natural and artificial. The Botanical Review, 8 (9):599-654. doi: 10.1007/BF02881046 URL
[26]	He Meng-xiao. 2019. Fine QTL mapping of parthenocapy in cucumber( Cucumis sativus L.)and validation of candidate gene expression[M. D. Dissertation]. Yangzhou:Yangzhou University. (in Chinese)
	何梦晓. 2019. 黄瓜单性结实QTL精细定位及候选基因表达验证[硕士论文]. 扬州:扬州大学.
[27]	Hoang T T, Nguyen T H, Le H T, Ngo B X, Wakana A. 2014. Self-incompatibility in pummelo[ Citrus grandis (L.) osbeck]with focus on vietnamese cultivars with or without parthenocarpy. Journal Faculty of Agriculture Kyushu University, 59 (1):65-70.
[28]	Huang J H, Wen S X, Zhang Y F, Zhong Q Z, Yang L, Chen L S. 2017. Abnormal megagametogenesis results in seedlessness of a polyembryonic ‘Meiguicheng’orange( Citrus sinensis)mutant created with gamma-rays. Scientia Horticulturae, 217:73-83. doi: 10.1016/j.scienta.2017.01.034 URL
[29]	Iwamasa M, Oba Y. 1980. Seedlessness due to self-incompatibility in Egami-buntan,a Japanese pummelo cultivar. Agricultural Bulletin of Saga University, 49:39-45.
[30]	Kakade V, Dubey A K, Sharma R M, Malik S K. 2017. Gametophytic self-incompatibility causes seedlessness in‘Kagzi Kalan’lemon(citrus limon). The Journal of Horticultural Science and Biotechnology, 92 (3):303-312. doi: 10.1080/14620316.2016.1276415 URL
[31]	Kim J S, Ezura K, Lee J, Ariizumi T, Ezura H. 2019. Genetic engineering of parthenocarpic tomato plants using transient SlIAA9 knockdown by novel tissue-specific promoters. Scientific Reports, 9 (1):18871. doi: 10.1038/s41598-019-55400-7 URL
[32]	Kim J S, Ezura K, Lee J, Kojima M, Takebayashi Y, Sakakibara H, Ariizumi T, Ezura H. 2020. The inhibition of SlIAA9 mimics an increase in endogenous auxin and mediates changes in auxin and gibberellin signalling during parthenocarpic fruit development in tomato. Journal of Plant Physiology,252.
[33]	Koltunow A M, Brennan P, Bond J E, Barker S J. 1998. Evaluation of genes to reduce seed size in shape Arabidopsis and tobacco and their application to shape citrus. Molecular Breeding, 4 (3):235-251. doi: 10.1023/A:1009610819338 URL
[34]	Kotoda N, Matsuo S, Honda I, Yano K, Shimizu T. 2017. Gibberellin 2-oxidase genes from satsuma mandarin( Citrus unshiu Marc.)caused late flowering and dwarfism in transgenic Arabidopsis. The Horticulture Journal, 86 (2):183-193. doi: 10.2503/hortj.OKD-016 URL
[35]	Lietzow C D, Zhu H, Pandey S, Havey M J, Weng Y. 2016. QTL mapping of parthenocarpic fruit set in North American processing cucumber. TAG. Theoretical and applied genetics. Theoretical and Applied Genetics, 129 (12):2387-2401. doi: 10.1007/s00122-016-2778-z URL
[36]	Liao F, Wang Y, Chen M, Chen W, Zong Y, Li Y, Yang L, Han X, Guo W. 2017. Low transcription of CmsIAA9 in the basal pistil is related to parthenocarpic fruiting of fingered citron(Foshou). Molecular Breeding, 37 (8):1-11. doi: 10.1007/s11032-016-0586-4 URL
[37]	Liang M, Cao Z, Zhu A, Liu Y, Tao M, Yang H, Jr Q X, Wang S, Liu J, Li Y, Chen C, Xie Z, Deng C, Ye J, Guo W, Xu Q, Xia R, Larkin R M, Deng X, Bosch M, Franklin-Tong V E, Chai L. 2020. Evolution of self-compatibility by a mutant Sm-RNase in citrus. Nature Plants, 6 (2):131-142. doi: 10.1038/s41477-020-0597-3 pmid: 32055045
[38]	Liu W, Chen M, Bai L, Zhuang Z, Fan C, Jiang N, Zhao J, Ma S, Xiang X. 2017. Comprehensive transcriptomics and proteomics analyses of pollinated and parthenocarpic litchi( Litchi chinensis Sonn.)fruits during early development. Scientific Reports, 7 (1):5401. doi: 10.1038/s41598-017-05724-z URL
[39]	Lin L, Zhang S, Ding F, He X, Luo C, Huang G, Do M, Wang Q, Yang Z, Su L, Hu G. 2019. Two genes(ClS1 and ClF-box)involved the Self-Incompatibility of“Xiangshui”Lemon[Citrus limon (L.)Burm. f.]. Plant Molecular Biology Reporter, 37 (1-2):50-62. doi: 10.1007/s11105-018-1133-8 URL
[40]	Liu L, Wang Z, Liu J, Liu F, Zhai R, Zhu C, Wang H, Ma F, Xu L. 2018. Histological,hormonal and transcriptomic reveal the changes upon gibberellin-induced parthenocarpy in pear fruit. Horticulture Research, 5:1. doi: 10.1038/s41438-017-0012-z URL
[41]	Liu Gao-ping. 2005. Study on the transformation of male sterility gene in citrus mediated by Agrobacterium tumefaciens[M. D. Dissertation]. Wuhan:Huazhong Agricultural University. (in Chinese)
	刘高平. 2005. 根癌农杆菌介导雄性不育基因转化柑橘的研究[硕士论文]. 武汉:华中农业大学.
[42]	Luo Sai-nan, Deng Zi-niu, Zhong Xiao-hong, Zhang Jia-yin, Yuan Fei-rong, Yang Li. 2008. Transformation of succari sweet orange with DefH9-iaaM gene. Journal of Hunan Agricultural University(Natural Sciences), 34 (2):177-181. (in Chinese)
	罗赛男, 邓子牛, 钟晓红, 张家银, 袁飞荣, 杨莉. 2008. 用单性结实基因 defH9-iaaM转化糖橙的研究. 湖南农业大学学报(自然科学版), 34 (2):177-181.
[43]	Martí C, Orzáez D, Ellul P, Moreno V, Carbonell J, Granell A. 2007. Silencing of DELLA induces facultative parthenocarpy in tomato fruits. The Plant Journal, 52 (5):865-76. doi: 10.1111/tpj.2007.52.issue-5 URL
[44]	Mesejo C, Yuste R, Martínez-Fuentes A, Reig C, Domingo J, Iglesias, Primo-Millo E, Agustí M. 2013. Self-pollination and parthenocarpic ability in developing ovaries of self-incompatible Clementine mandarins (Citrus clementina). Physiologia Plantarum, 148 (1):87-96. doi: 10.1111/ppl.2013.148.issue-1 URL
[45]	Mesejo C, Yuste R, Reig C, Martínez-Fuentes A, Domingo J, Iglesias, Muñoz-Fambuena N, Bermejo A, Germanà M, Primo-Millo E, Agustí M. 2016. Gibberellin reactivates and maintains ovary-wall cell division causing fruit set in parthenocarpic Citrus species. Plant Science, 247:13-24. doi: 10.1016/j.plantsci.2016.02.018 URL
[46]	Mezzetti B, Landi L, Pandolfini T, Spena A. 2004. The defH9-iaaM auxin-synthesizing gene increases plant fecundity and fruit production in strawberry and raspberry. BMC Biotechnology, 4 (1):4. doi: 10.1186/1472-6750-4-4 URL
[47]	Martínez-Bello L, Thomas M, López-Díaz I. 2015. Silencing C19-GA 2-oxidases induces parthenocarpic development and inhibits lateral branching in tomato plants. Journal of Experimental Botany, 66 (19):5897-910. doi: 10.1093/jxb/erv300 pmid: 26093022
[48]	Molesini B, Pandolfini T, Rotino G L, Dani V, Spena A. 2009. Aucsia gene silencing causes parthenocarpic fruit development in tomato. Plant Physiology, 149 (1):534-548. doi: 10.1104/pp.108.131367 URL
[49]	Mounet F, Moing A, Kowalczyk M, Rohrmann J, Petit J, Garcia V, Maucourt M, Yano K, Deborde C, Aoki K, Berges H, Granell A, Fernie AR, Bellini C, Rothan C, Lemaire-Chamley M. 2012. Down-regulation of a single auxin efflux transport protein in tomato induces precocious fruit development. Journal of Experimental Botany, 63 (13):4901-4917. doi: 10.1093/jxb/ers167 URL
[50]	Olimpieri I, Siligato F, Caccia R, Soressi G P, Mazzucato A, Mariotti L, Ceccarelli N. 2007. Tomato fruit set driven by pollination or by the parthenocarpic fruit allele are mediated by transcriptionally regulated gibberellin biosynthesis. Planta, 226 (4):877-88. URL pmid: 17503074
[51]	Pandolfini T. 2009. Seedless fruit production by hormonal regulation of fruit set. Nutrients, 1 (2):168-177. doi: 10.3390/nu1020168 pmid: 22253976
[52]	Pomares-Viciana T, Río-Celestino M D, Román B, Die J, Pico B, Gómez P. 2019. First RNA-seq approach to study fruit set and parthenocarpy in zucchini( Cucurbita pepo L.). BMC Plant Biology, 19 (1):61. doi: 10.1186/s12870-019-1632-2 pmid: 30727959
[53]	Ponti O M B D, Garretsen F. 1976. Inheritance of parthenocarpy in pickling cucumbers( Cucumis sativus L.)and linkage with other characters. Euphytica, 25 (1):633-642. doi: 10.1007/BF00041600 URL
[54]	Qian C, Ren N, Wang J, Xu Q, Chen X, Qi X. 2018. Effects of exogenous application of CPPU,NAA and GA ₄₊₇ on parthenocarpy and fruit quality in cucumber( Cucumis sativus L.). Food Chemistry, 243:410-413. doi: 10.1016/j.foodchem.2017.09.150 URL
[55]	Qiu W M, Zhu A D, Wang Y, Chai L J, Ge X X, Deng X X, Guo W W. 2012. Comparative transcript profiling of gene expression between seedless Ponkan mandarin and its seedy wild type during floral organ development by suppression subtractive hybridization and cDNA microarray. BMC Genomics, 13 (1):397. doi: 10.1186/1471-2164-13-397 URL
[56]	Qiu Wen-ming. 2013. Mechanism of seedlessness in Qianyang seedless Ponkanmandarin and identification of male sterility related genes[M. D. Dissertation]. Wuhan:Huazhong Agricultural University. (in Chinese)
	邱文明. 2013. “黔阳无核”椪柑无核机理及雄性不育相关基因发掘[硕士论文]. 武汉:华中农业大学.
[57]	Qiu Wen ming, Zhang Zu ming, Xu Yu hai, He Xiu juan, Sun Zhong hai, Tong Tao, Xiao Cui, Wu Li ming, Guo Wen wu. 2019. Cloning and functional analysis of male sterile 2-like protein gene(MSLP)in Citrus. Acta Horticulturae Sinica, 46 (12):2323-2334. (in Chinese)
	邱文明, 张祖铭, 徐育海, 何秀娟, 孙中海, 仝铸, 肖翠, 吴黎明, 郭文武. 2019. 柑橘雄性不育类蛋白基因MSLP的克隆与分析. 园艺学报, 46 (12):2323-2334.
[58]	Ruiu F, Picarella M E, Imanishi S, Mazzucato A. 2015. A transcriptomic approach to identify regulatory genes involved in fruit set of wild-type and parthenocarpic tomato genotypes. Plant Molecular Biology, 89 (3):263-278. doi: 10.1007/s11103-015-0367-1 URL
[59]	Royo C, Carbonell-Bejerano P, Torres-Pérez R, Nebish A, Martínez Ó, Rey M, Aroutiounian R, Ibáñez J, Martínez-Zapater J M. 2016. Developmental,transcriptome,and genetic alterations associated with parthenocarpy in the grapevine seedless somatic variant Corinto bianco. Journal of Experimental Botany, 67 (1):259-73. doi: 10.1093/jxb/erv452 URL
[60]	Sardos J, Rouard M, Hueber Y, Cenci A, Hyma K E, Houwe I, Hribova E, Courtois B, Roux N. 2016. A genome-wide association study on the seedless phenotype in banana( Musa spp.)reveals the potential of a selected panel to detect candidate genes in a vegetatively propagated crop. PloS ONE, 11 (5):e0154448. doi: 10.1371/journal.pone.0154448 URL
[61]	Serrani J C, Carrera E, Ruiz-Rivero O, Gallego-Giraldo L, Peres L E P, Garcia-Martinez J L. 2010. Inhibition of auxin transport from the ovary or from the apical shoot induces parthenocarpic fruit-set in tomato mediated by gibberellins. Plant Physiology, 153 (2):851-62. doi: 10.1104/pp.110.155424 pmid: 20388661
[62]	Serrani J C, Ruiz-Rivero O, Fos M, García-Martínez J L. 2008. Auxin-induced fruit set in tomato is mediated in part by gibberellins. The Plant Journal, 56 (6):922-934. doi: 10.1111/j.1365-313X.2008.03654.x pmid: 18702668
[63]	Shimizu T, Tanizawa Y, Mochizuki T, Nagasaki H, Yoshioka T, Toyoda A, Fujiyama A, Kaminuma E, Nakamura Y. 2017. Draft sequencing of the heterozygous diploid genome of satsuma( Citrus unshiu Marc.)Using a Hybrid Assembly Approach. Frontiers in Genetics, 8:180. doi: 10.3389/fgene.2017.00180 URL
[64]	Takisawa R, Maruyama T, Nakazaki T, Kataoka K, Saito H, Koeda S, Nunome T, Fukuoka H, Kitajima A. 2017. Parthenocarpy in the tomato ( Solanum lycopersicum L.)cultivar‘MPK-1’is controlled by a novel parthenocarpic gene. Horticulture Journal, 86 (4):487-492. doi: 10.2503/hortj.OKD-042 URL
[65]	Tang N, Deng W, Hu G, Hu N, Li Z. 2015. Transcriptome profiling reveals the regulatory mechanism underlying pollination dependent and parthenocarpic fruit set mainly mediated by auxin and gibberellin. PLoS ONE, 10 (4):e0125355. doi: 10.1371/journal.pone.0125355 URL
[66]	Talon M, Zacarias L, Primo-Millo E. 1990. Hormonal changes associated with fruit set and development in mandarins differing in their parthenocarpic ability. Physiologia Plantarum, 79 (2):400-406. doi: 10.1111/ppl.1990.79.issue-2 URL
[67]	Talon M, Zacarias L, Primo-Millo E. 1992. Gibberellins and parthenocarpic ability in developing ovaries of seedless mandarins. Plant Physiology, 99 (4):1575-81. doi: 10.1104/pp.99.4.1575 URL
[68]	Talon M, Tadeo F R, Ben-Cheikh W, Gomez-Cadenas A, Mehouachi J, Pérez-Botella J. 1998. Hormonal regulation of fruit set and abscission in citrus: classical concepts and new evidence. Acta Horticulturae,(463):209-217.
[69]	Teng C, Du D, Xiao L, Yu Q, Shang G, Zhao Z. 2017. Mapping and identifying a candidate gene(Bnmfs)for female-male sterility through whole-genome resequencing and RNA-Seq in rapeseed( Brassica napus L.). Frontiers in Plant Science, 8:2086. doi: 10.3389/fpls.2017.02086 URL
[70]	Ueta R, Abe C, Watanabe T, Sugano S S, Ishihara R, Ezura H, Osakabe Y, Osakabe K. 2017. Rapid breeding of parthenocarpic tomato plants using CRISPR/Cas9. Scientific Reports, 7 (1):507. doi: 10.1038/s41598-017-00501-4 URL
[71]	Vriezen W H, Feron R, Maretto F, Keijman J, Mariani C. 2008. Changes in tomato ovary transcriptome demonstrate complex hormonal regulation of fruit set. New Phytologist, 177 (1):60-76. pmid: 18028300
[72]	Vardi A, Neumann H, Frydman-Shani A, Yaniv Y, Spiegel-Roy P. 2000. Tentative model on the inheritance of juvenility,self-incompatibility and parthenocarpy. Acta Horticulturae, 535:199-203.
[73]	Wang H, Jones B, Li Z, Frasse P, Delalande C, Regad F, Chaabouni S, Latché A, Pech J C, Bouzayen M. 2005. The tomato Aux/IAA transcription factor IAA 9 is involved in fruit development and leaf morphogenesis. The Plant Cell, 17 (10):2676-2692. doi: 10.1105/tpc.105.033415 URL
[74]	Wen B, Song W, Sun M, Chen M, Mu Q, Zhang X, Wu Q, Chen X, Gao D, Wu H. 2019. Identification and characterization of cherry(Cerasus pseudocerasus G. Don)genes responding to parthenocarpy induced by GA ₃through transcriptome analysis BMC Genetics, 20 (1):65. doi: 10.1186/s12863-019-0746-8 URL
[75]	Wu Z, Zhang T, Li L, Xu J, Qin X, Zhang T L, Cui L, Luo Q, Li J, Chen J. 2016. Identification of a stable major-effect QTL(Parth 2.1)controlling parthenocarpy in cucumber and associated candidate gene analysis via whole genome re-sequencing. BMC Plant Biology, 16 (1):182. doi: 10.1186/s12870-016-0873-6 URL
[76]	Wu G A, Terol J, Ibanez V, López-García A, Pérez-Román E, Borredá C, Domingo C, Tadeo F R, Carbonell-Caballero J, Alonso R, Curk F, Du D, Ollitrault P, Roose M L, Dopazo J, Gmitter F G, Rokhsar D S, Talon M. 2018. Genomics of the origin and evolution of Citrus. Nature, 554 (7692):311-316. doi: 10.1038/nature25447 URL
[77]	Wu Zhe. 2015. QTL mapping and cadidate gene screening of parthenocarpy in cucumber[Ph. D. Dissertation]. Nanjing:Nanjing Agricultural University. (in Chinese)
	武喆. 2015. 黄瓜单性结实性状的QTL定位及候选基因筛选[博士论文]. 南京:南京农业大学.
[78]	Xiao J, Zhang L, Fan F, Liu X. 2018. Comparative transcript profiling reveals the mechanism of female sterility associated with seedless Ponkan mandarin( Citrus reticulata Blanco). Genome, 61 (8):595-604. doi: 10.1139/gen-2017-0215 URL
[79]	Xiao Jia-xin, Peng Shu-ang, He Hua-ping. 2005. Changes in Concentrations of IAA,ZR and GA ₃ during fruit development of parthenocarpic and self-pollinated citrus cultivars. Plant Science Journal, 23 (3):272-275. (in Chinese)
	肖家欣, 彭抒昂, 何华平. 2005. 单性结实和自花结实柑橘果实发育过程中IAA、ZR和GA ₃含量的变化. 武汉植物学研究, 23 (3):272-275.
[80]	Yan Li-ying, Lou Li-na, Li Xiao-li, Lou Qun-feng, Feng Zhi-hong, Chen Jin-feng. 2010. Inheritance of parthenocarpy in monoecious cucumber. Scientia Agricultura Sinica, 43 (6):1295-1301. (in Chinese)
	闫立英, 娄丽娜, 李晓丽, 娄群峰, 冯志红, 陈劲枫. 2010. 雌雄同株黄瓜单性结实性遗传分析. 中国农业科学, 43 (6):1295-1301.
[81]	Ye W, Qin Y, Ye Z, Silva J A T, Zhang L, Wu X, Lin S, Hu G. 2009. Seedless mechanism of a new mandarin cultivar‘Wuzishatangju’(Citrus reticulata Blanco). Plant Science, 177 (1):19-27. doi: 10.1016/j.plantsci.2009.03.005 URL
[82]	Yin Z, Malinowski R, Ziółkowska A, Sommer H, Plcader W, Malepszy S. 2006. The DefH9-iaaM-containing construct efficiently induces parthenocarpy in cucumber. Cellular & Molecular Biology Letters, 11 (2):279-290.
[83]	Yu Jie. 2009. Agrobacterium tumefaciens-mediated transformation of Barnase::iaaM genes into citrus[M.D. Dissertation]. Huazhong Agricultural University. (in Chinese)
	于洁. 2009. 根癌农杆菌介导的Barnase::iaaM基因转化柑橘的研究[硕士论文]. 华中农业大学.
[84]	Zhang S, Shi Q, Albrecht U, Shatters R G, Stange R, McCollum G, Zhang S, Fan C, Stover E. 2017. Comparative transcriptome analysis during early fruit development between three seedy citrus genotypes and their seedless mutants. Horticulture Research, 4:17041. doi: 10.1038/hortres.2017.41 URL
[85]	Zhang S W, Huang G X, Ding F, He X H, Pan J C. 2012. Mechanism of seedlessness in a new lemon cultivar‘Xiangshui’[Citrus limon(L.)Burm. F.]. Sexual Plant Reproduction, 25 (4):337-45. doi: 10.1007/s00497-012-0201-8 URL
[86]	Zhang Ying, Chen Yu-hui, Lian Yong, Liu Fu-zhong. 2018. Research progress on molecules of parthenocarpy in horticultural plants. Acta Horticulturae Sinica, 45 (7):1402-1414. (in Chinese)
	张映, 陈钰辉, 连勇, 刘富中. 2018. 园艺植物单性结实的分子研究进展. 园艺学报, 45 (7):1402-1414.
[87]	Zhang Zu-ming. 2015. Priliminary functional dissection of male-sterility gene MSLP in citrus[M. D. Dissertation]. Wuhan:Huazhong Agricultural University. (in Chinese)
	张祖铭. 2015. 柑橘雄性不育相关基因MSLP功能初步解析[硕士论文]. 武汉:华中农业大学.
[88]	Zhang Shang-long, Chen Kun-song, Ye Qing-fu, Chen Da-ming, Liu Chun-rong. 1994. Changes of IAA,ABA and ZT contents in ovary of unisexual fruit after pollination. Acta Horticalutruae Sinica, 21 (2):117-123. (in Chinese)
	张上隆, 陈昆松, 叶庆富, 陈大明, 刘春荣. 1994. 柑桔授粉处理和单性结实子房(幼果)内源IAA、ABA和ZT含量的变化. 园艺学报, 21 (2):117-123.
[89]	Zheng Qi-fa, Yang Shu-xi, Yang Jin-hua. 1995. A preliminary study on plant growth regulator induced monosexual fruiting of shatian pomelo. Guangdong Agricultural Sciences,(3):25-26. (in Chinese)
	郑启发, 杨淑兮, 杨锦华. 1995. 植物生长调节剂诱导沙田柚单性结实初探. 广东农业科学,(3):25-26.
[90]	Zhou X, Wakana A, Kim J H, Sakai K, Kajiwara K, Mizunoe Y. 2018a. Parthenocarpy in Citrus accessions with special focus on relatives of Kunenbo ( C. nobilis Lour. var. kunep Tanaka). Scientia Horticulturae, 232:29-39. doi: 10.1016/j.scienta.2017.12.045 URL
[91]	Zhou X, Wakana A, Sakai K, Mizunoe Y, Dewi P S, Kim J H. 2018b. Early diagnosis of parthenocarpic seedlings within one year after pollination with grapefruit( Citrus paradisi Macf.). Scientia Horticulturae, 239:282-288. doi: 10.1016/j.scienta.2018.04.051 URL
[92]	Zhu C, Zheng X, Huang Y, Ye J, Chen P, Zhang C, Zhao F, Xie Z, Zhang S, Wang N, Li H, Wang L, Tang X, Chai L, Xu Q, Deng X. 2019. Genome sequencing and CRISPR/Cas9 gene editing of an early flowering mini-citrus(Fortunella hindsii). Plant Biotechnology Journal, 17 (11):2199-2210. doi: 10.1111/pbi.v17.11 URL