Retrospection and Prospect of Chrysanthemum Genetic Breeding for Last Six Decades in China

doi:10.16420/j.issn.0513-353x.2022-0859

Acta Horticulturae Sinica ›› 2022, Vol. 49 ›› Issue (10): 2143-2162.doi: 10.16420/j.issn.0513-353x.2022-0859

• Reviews • Previous Articles Next Articles

Retrospection and Prospect of Chrysanthemum Genetic Breeding for Last Six Decades in China

SU Jiangshuo, JIA Diwen, WANG Siyue, ZHANG Fei, JIANG Jiafu, CHEN Sumei, FANG Weimin, and CHEN Fadi()

State Key Laboratory of Crop Genetics and Germplasm Enhancement,Key Laboratory of Landscaping,Ministry of Agriculture and Rural Affairs,Key Laboratory of Biology of Ornamental Plants in East China,National Forestry and Grassland Administration,College of Horticulture,Nanjing Agricultural University,Nanjing 210095,China

Received:2022-07-27 Revised:2022-08-22 Online:2022-10-25 Published:2022-10-31
Contact: and CHEN Fadi E-mail:chenfd@njau.edu.cn

Abstract

Abstract:

This review briefly summaries the processes for chrysanthemum development,and recalls the progress and main achievements that have been made in the germplasm resources,inheritance and molecular mechanisms underlying important traits formation,traditional breeding as well as modern biotechnology breeding programs during the last six decades in China. We also summarize the application status of modern biological technologies such as transgenic,genome editing,molecular marker-assisted selection,and multi-omics in the area of chrysanthemum genetics and breeding,and propose suggestions for the development of chrysanthemum genetics and breeding accordingly.

Key words: chrysanthemum, germplasm resources, genetic breeding, important trait, biotechnology, molecular mechanism

CLC Number:

S682.11

SU Jiangshuo, JIA Diwen, WANG Siyue, ZHANG Fei, JIANG Jiafu, CHEN Sumei, FANG Weimin, and CHEN Fadi. Retrospection and Prospect of Chrysanthemum Genetic Breeding for Last Six Decades in China[J]. Acta Horticulturae Sinica, 2022, 49(10): 2143-2162.

Figures/Tables 5

References 140

[1]	An C, Sheng L P, Du X P, Wang Y J, Zhang Y, Song A P, Jiang J F, Guan Z Y, Fang W M, Chen F D, Chen S M. 2019. Overexpression of CmMYB15 provides chrysanthemum resistance to aphids by regulating the biosynthesis of lignin. Horticulture Research, 6:84. doi: 10.1038/s41438-019-0166-y URL
[2]	Bai H R, Lin P, Li X, Liao X Q, Wan L H, Yang X H, Luo Y C, Zhang L, Zhang F, Liu S L, Liu Q L. 2021. DgC3H1,a CCCH zinc finger protein gene,confers cold tolerance in transgenic chrysanthemum. Scientia Horticulturae, 281:109901. doi: 10.1016/j.scienta.2021.109901 URL
[3]	Bi Meng-meng, Zhu Peng-fang, Li Xue-ying, Mao Hong-yu. 2019. Functional verification of CmDREBa-2 in Chrysanthemum. Journal of Shenyang Agricultural University, 50:406-413. (in Chinese)
	毕蒙蒙, 祝朋芳, 李雪莹, 毛洪玉. 2019. 菊花CmDREBa-2基因的功能验证. 沈阳农业大学学报, 50:406-413.
[4]	Cai Hai-yan, Wen Li-zhu, Zheng Cheng-shu, Li Ying-ying, Sun Xia, Sun Xian-zhi. 2013. Mutation types of chyrsanthemum induced with EMS and molecular identification of the important mutation characteristics. Journal of Shandong Agricultural University(Natural Science), 44 (2):171-175. (in Chinese)
	蔡海燕, 温立柱, 郑成淑, 李莹莹, 孙霞, 孙宪芝. 2013. EMS诱发菊花突变类型及重要性状的分子鉴定. 山东农业大学学报(自然科学版), 44 (2):171-175.
[5]	Chang Shu-lin. 1965. Studies on the classification of garden varieties of florists' Chrysanthemum. Acta Horticulturae Sinica, 4 (1):35-46. (in Chinese)
	张树林. 1965. 菊花品种分类的研究. 园艺学报, 4 (1):35-46.
[6]	Chen Fa-di, Chen Pei-du, Fang Wei-min, Li Hong-jian. 1998. Cytogenetics of F₁-hybrids between two small-headed cultivars of Dendranthema × grandiflorum and two wild Dendranthema species. Acta Horticulturae Sinica, 25 (3):308-309. (in Chinese)
	陈发棣, 陈佩度, 房伟民, 李鸿渐. 1998. 栽培小菊与野生菊间杂交一代的细胞遗传学初步研究. 园艺学报, 25 (3):308-309.
[7]	Chen Fa-di, Chen Pei-du, Li Hong-jian. 1996. Genome analysis and their phylogenetic relationships of several wild species of Dendranthema in China. Acta Horticulturae Sinica, 23 (1):67-72. (in Chinese)
	发棣, 陈佩度, 李鸿渐. 1996. 几种中国野生菊的染色体组分析及亲缘关系初步研究. 园艺学报, 23 (1):67-72.
[8]	Chen Fa-di, Chen Su-mei, Fang Wei-min, Zhang Fei, Jiang Jia-fu, Teng Nian-jun, Guan Zhi-yong, Wang Hai-bin, Song Ai-ping, Zhao Shuang. 2016. Discovery of excellent chrysanthemum germplasms and germplasm enhancement. Bulletin of National Natural Science Foundation of China, 30 (2):112-116. (in Chinese)
	陈发棣, 陈素梅, 房伟民, 张飞, 蒋甲福, 滕年军, 管志勇, 王海滨, 宋爱萍, 赵爽. 2016. 菊花优异种质资源挖掘与种质创新研究. 中国科学基金, 30 (2):112-116.
[9]	Chen Feng-huai, Wang Qui-pu. 1979. Chrysanthemum retraces. Chinese Journal of Nature,(10):62-63. (in Chinese)
	陈封怀, 王秋圃. 1979. 菊花探源. 自然杂志,(10):62-63.
[10]	Chen Jun-yu. 2012. The origin of garden chrysanthemum. Hefei: Anhui Science and Technology Press:67-87. (in Chinese)
	陈俊愉. 2012. 菊花起源. 合肥: 安徽科技出版社:67-87.
[11]	Chen L, Chen Y, Jiang J F, Chen S M, Chen F D, Guan Z Y, Fang W M. 2012. The constitutive expression of Chrysanthemum dichrum ICE1 in Chrysanthemum grandiflorum improves the level of low temperature,salinity and drought tolerance. Plant Cell Reports, 31:1747-1758. doi: 10.1007/s00299-012-1288-y pmid: 22645020
[12]	Chen X, Zhou X, Xi L, Li J, Zhao R, Ma N, Zhao L. 2013. Roles of DgBRC1 in regulation of lateral branching in chrysanthemum(Dendranthema × grandiflora cv. Jinba). PLoS ONE, 8:e61717. doi: 10.1371/journal.pone.0061717 URL
[13]	Cheng Li-na, Wei Qian, Muhammad Imtiaz, Gao Jun-ping, Hong Bo. 2013. Advances in application of transgenic breeding technology in the traits improvement of chrysanthemum. Acta Horticulturae Sinica, 40 (9):1813-1825. (in Chinese)
	成丽娜, 魏倩, Muhammad Imtiaz, 高俊平, 洪波. 2013. 转基因育种技术在菊花性状改良中的应用进展. 园艺学报, 40 (9):1813-1825.
[14]	Cheng P L, Liu Y N, Yang Y M, Chen H, Cheng H, Hu Q, Zhang Z X, Gao J J, Zhang J X, Ding L, Fang W M, Chen S M, Chen F D, Jiang J F. 2020. CmBES1 is a regulator of boundary formation in chrysanthemum ray florets. Horticulture Research, 7:129. doi: 10.1038/s41438-020-00351-8 URL
[15]	Chong X R, Su J S, Wang F, Wang H B, Song A P, Guan Z Y, Fang W M, Jiang J F, Chen S M, Chen F D, Zhang F. 2019. Identification of favorable SNP alleles and candidate genes responsible for inflorescence-related traits via GWAS in chrysanthemum. Plant Molecular Biology, 99:407-420. doi: 10.1007/s11103-019-00826-w pmid: 30701353
[16]	Chong X R, Zhang F, Wu Y Y, Yang X D, Zhao N, Wang H B, Guan Z Y, Fang W M, Chen F D. 2016. A SNP-enabled assessment of genetic diversity,evolutionary relationships and the identification of candidate genes in chrysanthemum. Genome Biology and Evolution, 8:3661-3671.
[17]	Cobb J N, Biswas P S, Platten J D. 2019. Back to the future:revisiting MAS as a tool for modern plant breeding. Theoretical and Applied Genetics, 132:647-667. doi: 10.1007/s00122-018-3266-4 URL
[18]	Dai Si-lan, Chen Jun-yu. 1996. Artificial interspecific cross among seven species of dendranthema in China. Journal of Beijing Forestry University,(4):16-22. (in Chinese)
	戴思兰, 陈俊愉. 1996. 菊属7个种的人工种间杂交试验. 北京林业大学学报,(4):16-22.
[19]	Dai Si-lan, Huang He, Fu Jian-xin, Hong Yan. 2013. Advances in molecular breeding of ornamental plants. Chinese Bulletin of Botany, 48 (6):589-607. (in Chinese) doi: 10.3724/SP.J.1259.2013.00589 URL
	戴思兰, 黄河, 付建新, 洪艳. 2013. 观赏植物分子育种研究进展. 植物学报, 48 (6):589-607. doi: 10.3724/SP.J.1259.2013.00589
[20]	Dierck R, Leus L, Dhooghe E, van Huylenbroeck J, de Riek J, van Der Straeten D, de Keyser E. 2018. Branching gene expression during chrysanthemum axillary bud outgrowth regulated by strigolactone and auxin transport. Plant Growth Regulation, 86 (1):23-36. doi: 10.1007/s10725-018-0408-2 URL
[21]	Ding L, Song A P, Zhang X, Li S, Su J S, Xia W K, Zhao K K, Zhao W Q, Guan Y X, Fang W M, Chen S M, Jiang J F, Chen F D. 2020. The core regulatory networks and hub genes regulating flower development in Chrysanthemum morifolium. Plant Molecular Biology, 103:669-688. doi: 10.1007/s11103-020-01017-8 URL
[22]	Ding L, Zhao K K, Zhang X, Song A P, Su J S, Hu Y H, Zhao W Q, Jiang J F, Chen F D. 2019. Comprehensive characterization of a floral mutant reveals the mechanism of hooked petal morphogenesis in Chrysanthemum morifolium. Plant Biotechnology Journal, 17:2325-2340. doi: 10.1111/pbi.13143 URL
[23]	Dong L L, Wang Q, Xiong F, Liu N, Zhang S M. 2018. Isolation and functional analysis of CmMAX1 from chrysanthemum. Journal of the American Society for Horticultural Science, 143:430-435. doi: 10.21273/JASHS04412-18 URL
[24]	Fan M, Gao Y K, Wu Z P, Zhang Q X. 2020. Linkage map development by EST-SSR markers and QTL analysis for inflorescence and leaf traits in chrysanthemum(Chrysanthemum morifolium Ramat.). Plants, 9:1342. doi: 10.3390/plants9101342 URL
[25]	Fan Q Q, Song A P, Xin J J, Chen S M, Jiang J F, Wang Y J, Li X R, Chen F D. 2015. CmWRKY15facilitates Alternaria tenuissima infection of chrysanthemum. PLoS ONE, 10:e0143349. doi: https://doi.org/10.1371/journal.pone.0143349. doi: https://doi.org/10.1371/journal.pone.0143349 URL
[26]	Fu Jing, Dai Si-lan. 2016. Analysis of color phenotypic and pigment contents of chrysanthemum based on hyperspectral imaging. Journal of Beijing Forestry University, 38 (8):88-98. (in Chinese)
	伏静, 戴思兰. 2016. 基于高光谱成像技术的菊花花色表型和色素成分分析. 北京林业大学学报, 38 (8):88-98.
[27]	Fu X, Su J S, Yu K L, Cai Y F, Zhang F, Chen S M, Fang W M, Chen F D, Guan Z Y. 2018. Genetic variation and association mapping of aphid(Macrosiphoniella sanbourni)resistance in chrysanthemum(Chrysanthemum morifolium Ramat.). Euphytica, 214:1-9.
[28]	Fu Xiao, Su Jiang-shuo, Li Yuan-yuan, Zhang Fei, Fang Wei-min, Chen Su-mei, Chen Fa-di, Guan Zhi-yong. 2019. Genetic variation and QTL mapping for aphid resistance in an F₁ population of chrysanthemum. Acta Horticulturae Sinica, 46 (7):1351-1358. (in Chinese)
	付晓, 苏江硕, 李媛媛, 张飞, 房伟民, 陈素梅, 陈发棣, 管志勇. 2019. 菊花F₁代抗蚜性遗传变异与QTL定位. 园艺学报, 46 (7):1351-1358.
[29]	Gai Jun-yi. 2003. Genetic research on quantitative traits in plants—separation analysis method of major gene-polygene genetic system. Scientific Chinese, 11:46-48. (in Chinese)
	盖钧镒. 2003. 植物数量性状遗传研究——主基因-多基因遗传体系分离分析方法. 科学中国人, 11:46-48.
[30]	Gleim S, Lubieniechi S, Smyth S J. 2020. CRISPR-Cas 9 application in Canadian public and private plant breeding. The CRISPR Journal, 3:44-51.
[31]	Guan Y X, Ding L, Jiang J F, Shentu Y Y, Zhao W Q, Zhao K K, Zhang X, Song A P, Chen S M, Chen F D. 2021. Overexpression of the CmJAZ1-like gene delays flowering in Chrysanthemum morifolium. Horticulture Research, 8:87. doi: 10.1038/s41438-021-00525-y URL
[32]	Han X Y, Luo Y T, Lin J Y, Wu H Y, Sun H, Zhou L J, Chen S M, Guan Z Y, Fang W M, Zhang F, Chen F D, Jiang J F. 2021. Generation of purple-violet chrysanthemums via anthocyanin B-ring hydroxylation and glucosylation introduced from Osteospermum hybrid F3'5'H and Clitoria ternatea A3'5'GT. Ornamental Plant Research, 1:1-9.
[33]	He Jun-ping, Chen Fa-di, Chen Su-mei, Fang Wei-min. 2010. Aphid-resistance of chrysanthemum cultivars. Chinese Journal of Eclolgy, 29:1382-1386. (in Chinese)
	何俊平, 陈发棣, 陈素梅, 房伟民. 2010. 不同菊花品种抗蚜虫性鉴定. 生态学杂志, 29:1382-1386.
[34]	He L, Wu Y H, Zhao Q, Wang B, Liu Q L, Zhang L. 2018. Chrysanthemum DgWRKY2 gene enhances tolerance to salt stress in transgenic chrysanthemum. International Journal of Molecular Sciences, 19:2062. doi: 10.3390/ijms19072062 URL
[35]	He Ling. 2019. Joint sequencing analysis of transcriptome in chrysanthemum leaves under low temperature stress and cloning and functional identification of DgMYB1 gene[M. D. Dissertation]. Ya'an: Sichuan Agricultural University. (in Chinese)
	何玲. 2019. 低温胁迫下菊花叶转录组联合测序分析及DgMYB1基因的克隆与功能鉴定[硕士论文]. 雅安: 四川农业大学.
[36]	He Zhen. 2015. Studies on evaluation of standard cut flower of chrysanthemum branching characteristics and heredity analysis[M. D. Dissertation]. Nanjing: China Agricultural University. (in Chinese)
	何臻. 2015. 标准切花菊分枝特性评价及其遗传分析[硕士论文]. 南京: 南京农业大学.
[37]	Hirakawa H, Sumitomo K, Hisamatsu T, Nagano S, Shirasawa K, Higuchi Y, Kusaba M, Koshioka M, Nakano Y, Yagi M. 2019. De novo whole-genome assembly in Chrysanthemum seticuspe,a model species of Chrysanthemums,and its application to genetic and gene discovery analysis. DNA Research, 26:195-203. doi: 10.1093/dnares/dsy048 pmid: 30689773
[38]	Hong Bo, Shi Chun-feng, Zhang Xiao-jiao, Gao Jun-ping. 2009. Advances in research of ornamental and agricultural traits in chrysanthemum by gene engineering. Scientia Agricultura Sinica, 42:1348-1358. (in Chinese)
	洪波, 史春凤, 张晓娇, 高俊平. 2009. 菊花观赏性状和农艺性状基因工程改良研究进展. 中国农业科学, 42:1348-1358.
[39]	Huang H, Hu K, Han K T, Xiang Q Y, Dai S L. 2013. Flower colour modification of chrysanthemum by suppression of F3'H and overexpression of the exogenous Senecio cruentus F3'5'H gene. PLoS ONE, 8:e74395. doi: 10.1371/journal.pone.0074395 URL
[40]	Huang H F, Gao X K, Gao X, Zhang S Q, Zheng Y, Zhang N, Hong B, Zhao X, Gu Z Y. 2022a. Flower color mutation,pink to orange,through CmGATA4-CCD4a-5 module regulates carotenoids degradation in chrysanthemum. Plant Science, 322:111290. doi: 10.1016/j.plantsci.2022.111290 URL
[41]	Huang Q X, Liao X Q, Yang X H, Luo Y C, Lin P, Zeng Q H, Bai H R, Jiang B B, Pan Y Z, Zhang F, Zhang L, Jia Y, Li Q L. 2021. Lysine crotonylation of DgTIL1 at K72 modulates cold tolerance by enhancing DgnsLTP stability in chrysanthemum. Plant Biotechnology Journal, 19:1125-1140. doi: 10.1111/pbi.13533 pmid: 33368971
[42]	Huang Y Y, Xing X J, Tang Y, Jin J Y, Ding L, Song A P, Chen S M, Chen F D, Jiang J F, Fang W M. 2022b. An ethylene-responsive transcription factor and a flowering locus KH domain homologue jointly modulate photoperiodic flowering in chrysanthemum. Plant Cell Environment, 45:1442-1456. doi: 10.1111/pce.14261 URL
[43]	Ji Qing-ping. 1987. Experiments and discussion on the origin of Chinese chrysanthemum[Ph. D. Dissertation]. Beijing: Beijing Forestry University. (in Chinese)
	吉庆萍. 1987. 有关中国菊花起源的实验与探讨[博士论文]. 北京: 北京林业大学.
[44]	Kishi-Kaboshi M, Aida R, Sasaki K. 2017. Generation of gene-edited Chrysanthemum morifolium using multicopy transgenes as targets and markers. Plant and Cell Physiology, 58:216-226. doi: 10.1093/pcp/pcw222 pmid: 28049122
[45]	Lan Feng. 2018. The study on mutagenic effects of ⁶⁰Co γ-rays on two kinds of ground-grow chrysanthemum[M. D. Dissertation]. Harbin: Northeast Forestry University. (in Chinese)
	兰凤. 2018. ⁶⁰Co-γ射线对两种露地菊品种辐射诱变效应的研究[硕士论文]. 哈尔滨: 东北林业大学.
[46]	Li Cui, Hao Fu-shun, Sun Li-rong. 2018. DgGA20ox was effectively silenced by CRISPR/Cas 9 technology in Chrysanthemum. Journal of Henan Institute of Science and Technology(Natural Science Edition), 46:22-26. (in Chinese)
	李翠, 郝福顺, 孙立荣. 2018. 利用 CRISPR/Cas9 技术有效沉默菊花DgGA20ox基因. 河南科技学院学报(自然科学版), 46:22-26.
[47]	Li F, Zhang H R, Zhao H S, Gao T W, Song A P, Jiang J F, Chen F D, Chen S M. 2018. Chrysanthemum CmHSFA4 gene positively regulates salt stress tolerance in transgenic chrysanthemum. Plant Biotechnology Journal, 16:1311-1321. doi: 10.1111/pbi.12871 URL
[48]	Li Hong-jian, Shao Jian-wen. 1990. Investigation,collection and classification of chrysanthemum cultivars in China. Journal of Nanjing Agricultural University, 13 (1):30-36. (in Chinese)
	李鸿渐, 邵建文. 1990. 中国菊花品种资源的调查收集与分类. 南京农业大学学报, 13 (1):30-36.
[49]	Li P, Zhang F, Chen S, Jiang J, Wang H, Su J, Fang W, Guan Z, Chen F. 2016. Genetic diversity,population structure and association analysis in cut chrysanthemum(Chrysanthemum morifolium Ramat.). Molecular Genetics and Genomics, 291:1117-1125. doi: 10.1007/s00438-016-1166-3 URL
[50]	Li P L, Song A P, Gao C Y, Jiang J F, Chen S M, Fang W M, Zhang F, Chen F D. 2015. The over-expression of a chrysanthemum WRKY transcription factor enhances aphid resistance. Plant Physiology and Biochemistry, 95:26-34. doi: 10.1016/j.plaphy.2015.07.002 pmid: 26184088
[51]	Li P R, Su J S, Guan Z Y, Fang W M, Chen F D, Zhang F. 2018. Association analysis of drought tolerance in cut chrysanthemum(Chrysanthemum morifolium Ramat.)at seedling stage. 3 Biotech, 8:1-9.
[52]	Li X, Yang Q, Liao X Q, Tian Y C, Zhang F, Zhang L, Liu Q L. 2022. A natural antisense RNA improves chrysanthemum cold tolerance by regulating the transcription factor DgTCP1. Plant Physiology,doi: https://doi.org/10.1093/plphys/kiac267. doi: https://doi.org/10.1093/plphys/kiac267 URL
[53]	Li Xin-lei, Chen Fa-di. 2004a. Advances of genetic improvement and germplasm resources for chrysanthemum. Chinese Bulletin of Botany, 21:392-401. (in Chinese)
	李辛雷, 陈发棣. 2004a. 菊花种质资源与遗传改良研究进展. 植物学通报, 21:392-401.
[54]	Li Xin-lei, Chen Fa-di. 2004b. RAPD analysis of wild species cultivars and interspecific hybrids in Dendranthema. Journal of Nanjing Agricultural University,(3):29-33. (in Chinese)
	李辛雷, 陈发棣. 2004b. 菊属野生种、栽培菊花及种间杂种的RAPD分析. 南京农业大学学报,(3):29-33.
[55]	Liang J L, Zhao L J, Challis R, Leyser O. 2010. Strigolactone regulation of shoot branching in chrysanthemum(Dendranthema grandiflorum). Journal of Experimental Botany, 61:3069-3078. doi: 10.1093/jxb/erq133 URL
[56]	Liu Rui. 2010. Phylogenetic relationship studies on some wild species and cultivated varieties in chrysanthemum[M. D. Dissertation]. Baoding: Agricultural University of Hebei Province. (in Chinese)
	刘蕤. 2010. 部分菊属野生种与栽培菊花亲缘关系研究[硕士论文]. 保定: 河北农业大学.
[57]	Liu Wei-xin. 2019. Molecular mechanism of sucrose regulates axillary bud growth in chrysanthemum[Ph. D. Dissertation]. Nanjing: Nanjing Agricultural University. (in Chinese)
	刘伟鑫. 2019. 蔗糖调控菊花侧芽生长的分子机理研究[博士论文]. 南京: 南京农业大学.
[58]	Liu X F, Xiang L L, Yin X R, Grierson D, Li F, Chen K S. 2015. The identification of a MYB transcription factor controlling anthocyanin biosynthesis regulation in Chrysanthemum flowers. Scientia Horticulturae, 194:278-285. doi: 10.1016/j.scienta.2015.08.018 URL
[59]	Liu Y, Wang L, Liu C X, Yin H, Liu H, Luo H, He M, Zhou Y W. 2022. CgbZIP1:a bZIP transcription factor from Chrysanthemum grandiflora confers plant tolerance to salinity and drought stress. Agronomy, 12:556. doi: 10.3390/agronomy12030556 URL
[60]	Liu Y, Xin J, Liu L, Song A, Guan Z, Fang W, Chen F. 2020. A temporal gene expression map of Chrysanthemum leaves infected with Alternaria alternata reveals different stages of defense mechanisms. Horticulture Research, 7:23. doi: 10.1038/s41438-020-0245-0 pmid: 32140232
[61]	Liu Yi-qi. 2016. Isolation and function analysis of floral symmetry gene CmDIV in Chrysanthemum morifolium'Maoxiangyu'[M. D. Dissertation]. Beijing: Beijing Foresty University. (in Chinese)
	刘轶奇. 2016. 菊花'毛香玉'花对称性调控CmDIV基因克隆及分析[硕士论文]. 北京: 北京林业大学.
[62]	Liu Z L, Wang J, Tian Y, Dai S L. 2019. Deep learning for image-based large-flowered chrysanthemum cultivar recognition. Plant Methods, 15:1-11. doi: 10.1186/s13007-018-0385-5 URL
[63]	Ma Chao-feng. 2019. Expression analysis of PHYA and PHYB homologous gene in Chrysanthemum lavandulifolium and C. vestitum and ClPHYB functional verification[M. D. Dissertation]. Beijing: Beijing Foresty University. (in Chinese)
	马朝峰. 2019. 甘菊和毛华菊PHYA和PHYB同源基因表达分析及ClPHYB功能验证[硕士论文]. 北京: 北京林业大学.
[64]	Mao Y, Sun J, Cao P, Zhang R, Fu Q, Chen S, Chen F, Jiang J. 2016. Functional analysis of alternative splicing of the FLOWERING LOCUS T orthologous gene in Chrysanthemum morifolium. Horticulture Research, 3:16058. doi: 10.1038/hortres.2016.58 URL
[65]	Nakano M, Hirakawa H, Fukai E, Toyoda A, Kajitani R, Minakuchi Y, Itoh T, Higuchi Y, Kozuka T, Bono H. 2021. A chromosome-level genome sequence of Chrysanthemum seticuspe,a model species for hexaploid cultivated chrysanthemum. Communications Biology, 4:1-11. doi: 10.1038/s42003-020-01566-0 URL
[66]	Nie J, Wen C, Xi L, Lv S H, Zhao Q C, Kou Y P, Ma N, Zhao L J, Zhou X F. 2018. The AP2/ERF transcription factor CmERF053 of chrysanthemum positively regulates shoot branching,lateral root,and drought tolerance. Plant Cell Reports, 37:1049-1060. doi: 10.1007/s00299-018-2290-9 URL
[67]	Peng H, Zhang F, Jiang J F, Chen S M, Fang W M, Guan Z Y, Chen F D. 2015. Identification of quantitative trait loci for branching traits of spray cut chrysanthemum. Euphytica, 202:385-392. doi: 10.1007/s10681-014-1259-1 URL
[68]	Ping Q, Cheng P L, Huang F, Ren L P, Cheng H, Guan Z Y, Fang W M, Chen S M, Chen F D, Jiang J F. 2019. The heterologous expression in Arabidopsis thaliana of a chrysanthemum gene encoding the BBX family transcription factor CmBBX13 delays flowering. Plant Physiology and Biochemistry, 144:480-487. doi: S0981-9428(19)30420-6 pmid: 31655346
[69]	Qi C, Gao J, Chen K, Shu L, Pearson S. 2022. Tea Chrysanthemum detection by leveraging generative adversarial networks and edge computing. Frontiers in Plant Science, 13:850606. doi: 10.3389/fpls.2022.850606 URL
[70]	Qi Meng-wen, Wang Hua-guo. 1997. Progress and analysis of flower radiation breeding in China. Journal of Nuclear Agricultural Sciences,(6):39-41. (in Chinese)
	齐孟文, 王化国. 1997. 我国花卉辐射育种的进展与剖析. 核农学通报,(6): 39-41.
[71]	Song C, Liu Y F, Song A P, Dong G Q, Zhao H B, Sun W, Ramakrishnan S, Wang Y, Wang S B, Li T Z, Niu Y, Jiang J F, Dong B, Xia Y, Chen S M, Hu Z G, Chen F D, Chen S L. 2018. The Chrysanthemum nankingense genome provides insights into the evolution and diversification of chrysanthemum flowers and medicinal traits. Molecular Plant, 11:1482-1491. doi: S1674-2052(18)30308-3 pmid: 30342096
[72]	Song Feng-yan. 2016. Photoperiod pathway related gene AtCO in genetic transformation of Chrysanthemum morifolium'Huoyan'[M. D. Dissertation]. Harbin: Northeast Forestry University. (in Chinese)
	宋凤艳. 2016. 光周期途径相关基因AtCO在露地菊'火焰'中的遗传转化[硕士论文]. 哈尔滨: 东北林业大学.
[73]	Song X B, Xu Y H, Gao K, Fan G X, Zhang F, Deng C Y, Dai S L, Huang H, Xin H G, Li Y Y. 2020. High-density genetic map construction and identification of loci controlling flower-type traits in Chrysanthemum(Chrysanthemum× morifolium Ramat.). Horticulture Research, 7:108. doi: 10.1038/s41438-020-0333-1 URL
[74]	Song X B, Zhao X G, Fan G X, Gao K, Dai S L, Zhang M, Ma C F, Wu X Y. 2018. Genetic analysis of the corolla tube merged degree and the relative number of ray florets in chrysanthemum(Chrysanthemum × morifolium Ramat.). Scientia Horticulturae, 242:214-224. doi: 10.1016/j.scienta.2018.07.010 URL
[75]	Su J S, Jiang J F, Zhang F, Liu Y, Ding L, Chen S M, Chen F D. 2019a. Current achievements and future prospects in the genetic breeding of chrysanthemum:a review. Horticulture Research, 6:109.
[76]	Su J S, Yang X C, Zhang F, Wu S F, Xiong S Y, Shi L M, Guan Z Y, Fang W M, Chen F D. 2018. Dynamic and epistatic QTL mapping reveals the complex genetic architecture of waterlogging tolerance in chrysanthemum. Planta, 247:899-924. doi: 10.1007/s00425-017-2833-2 pmid: 29273861
[77]	Su J S, Zhang F, Chong X R, Song A P, Guan Z Y, Fang W M, Chen F D. 2019b. Genome-wide association study identifies favorable SNP alleles and candidate genes for waterlogging tolerance in chrysanthemums. Horticulture Research, 6:21. doi: 10.1038/s41438-018-0101-7 URL
[78]	Su J S, Zhang F, Li P R, Guan Z Y, Fang W M, Chen F D. 2016. Genetic variation and association mapping of waterlogging tolerance in chrysanthemum. Planta, 244:1241-1252. pmid: 27522648
[79]	Su J S, Zhang F, Yang X C Feng Y X, Yang X D, Wu Y Y, Guan Z Y, Fang W M, Chen F D. 2017. Combining ability,heterosis,genetic distance and their intercorrelations for waterlogging tolerance traits in chrysanthemum. Euphytica, 213 (2):1-15. doi: 10.1007/s10681-016-1788-x URL
[80]	Sun J, Wang H, Ren L P, Chen S M, Jiang J F. 2017. CmFTL2 is involved in the photoperiod- and sucrose-mediated control of flowering time in chrysanthemum. Horticulture Research, 4:17001. doi: 10.1038/hortres.2017.1 URL
[81]	Sun W, Yang X C, Su J S, Guan Z Y, Jiang J F, Chen F D, Fang W M, Zhang F. 2019. The genetics of planting density-dependent branching in chrysanthemum. Scientia Horticulturae, 256:108598. doi: 10.1016/j.scienta.2019.108598 URL
[82]	Tang M W, Xue W J, Li X Q, Wang L S, Wang M, Wang W P, Yin X, Chen B W, Qu X T, Li J Y, Gao X Y, Wei X F, Bu F Q, Zhang L Y, Sui Z R, Ding B, Wang Y, Zhang Q Z, Li Y H, Zhang Y. 2022. Mitotically heritable epigenetic modifications of CmMYB6 control anthocyanin biosynthesis in Chrysanthemum. New Phytologist,doi: https://doi.org/10.1111/nph.18389. doi: https://doi.org/10.1111/nph.18389 URL
[83]	Teixeira da Silva J A, Shinoyama H, Aida R, Matsushita Y, Raj S K, Chen F D. 2013. Chrysanthemum biotechnology:Quo vadis? Critical Reviews in Plant Sciences, 32:21-52. doi: 10.1080/07352689.2012.696461 URL
[84]	van Lieshout N, van Kaauwen M, Kodde L, Arens P, Smulders M J, Visser R G, Finkers R. 2022. De novo whole-genome assembly of Chrysanthemum makinoi, a key wild chrysanthemum. G3, 12:jkab358. doi: https://doi.org/10.1093/g3journal/jkab358. doi: https://doi.org/10.1093/g3journal/jkab358 URL
[85]	Wang Bo-nan. 2007. Mutagenic effect of γ-Rays on Dendrite ma. moratorium Tzvel. Henan Science, 25 (5):758-760. (in Chinese)
	王柏楠. 2007. γ射线对菊花的诱变效应. 河南科学, 25 (5):758-760.
[86]	Wang C C, Zhang F, Guan Z Y, Chen S M, Jiang J F, Fang W M, Chen F D. 2014. Inheritance and molecular markers for aphid(Macrosiphoniella sanbourni)resistance in chrysanthemum(Chrysanthemum morifolium Ramat.). Scientia Horticulturae, 180:220-226. doi: 10.1016/j.scienta.2014.10.038 URL
[87]	Wang Hai, Li Tong, Zhou Yuan-yuan, Wang Wang, Zhao Hui-en. 2021. Cloning and expression analysis of Lignin-regulated transcription factor ClSND1-like from Chrysanthemum lavandulifolium. Acta Botanica Boreali-Occidentalia Sinica, 41:359-367. (in Chinese)
	王海, 李童, 周圆圆, 汪王, 赵惠恩. 2021. 甘菊木质素调控转录因子ClSND1-like的克隆与表达分析. 西北植物学报, 41:359-367.
[88]	Wang Hong, Chen Fa-di, Zhao Hong-bo, Fang Wei-min. 2007. Mutagenic effect of Pingyangmycin on Dendranthema × grandiflorum'Italy Red' with small inflorescence. Journal of Nanjing Agricultural University,(4):39-43. (in Chinese)
	王红, 陈发棣, 赵宏波, 房伟民. 2007. 平阳霉素对盆栽小菊意大利红的诱变效应. 南京农业大学学报,(4):39-43.
[89]	Wang J J, Guan Y X, Ding L, Li P R, Zhao W Q, Jiang J F, Chen S M, Chen F D. 2019. The CmTCP20 gene regulates petal elongation growth in Chrysanthemum morifolium. Plant Science, 280:248-257. doi: 10.1016/j.plantsci.2018.12.008 URL
[90]	Wang L J, Gao J J, Zhang Z X, Liu W M, Cheng P L, Mu W T, Su T, Chen S M, Chen F D, Jiang J F. 2020a. Overexpression of CmSOS1 confers waterlogging tolerance in chrysanthemum. Journal of Integrative Plant Biology, 62:1059-1064. doi: 10.1111/jipb.12889 URL
[91]	Wang L J, Sun J, Ren L P, Zhou M, Han X Y, Ding L, Zhang F, Guan Z Y, Fang W M, Chen S M, Chen F D, Jiang J F. 2020b. CmBBX8 accelerates flowering by targeting CmFTL1directly in summer chrysanthemum. Plant Biotechnology Journal, 18:1562-1572. doi: 10.1111/pbi.13322 URL
[92]	Wang T L, Wei Q, Wang Z L, Liu W W, Zhao X, Ma C, Gao J P, Xu Y J, Hong B. 2022a. CmNF‐YB 8 affects drought resistance in chrysanthemum by altering stomatal status and leaf cuticle thickness. Journal of Integrative Plant Biology, 64:741-755. doi: 10.1111/jipb.13201 URL
[93]	Wang Wei-jiao. 2019. Cloning of terpene synthase genes CmTPS1,2, 3 and functional identification of CmTPS2,3 in chrysanthemum[M. D. Dissertation]. Nanjing: Nanjing Agricultural University. (in Chinese)
	王威姣. 2019. 菊花萜烯合成酶基因CmTPS1,2, 3的克隆及CmTPS2,3的功能鉴定[硕士论文]. 南京: 南京农业大学.
[94]	Wang Yi. 2013. The expression pattern and functional characterization of flowering related genes in Chrysanthemum lavandulifolium(Fisch. Ex Trautv.)Makino[Ph. D. Dissertation]. Beijing: Beijing Foresty University. (in Chinese)
	王翊. 2013. 甘菊成花相关基因表达模式和功能的研究[博士论文]. 北京: 北京林业大学.
[95]	Wang Y G, Zhou L J, Wang Y X, Geng Z Q, Ding B Q, Jiang J F, Chen S M, Chen F D. 2022b. An R2R3-MYB transcription factor CmMYB 21 represses anthocyanin biosynthesis in color fading petals of chrysanthemum. Scientia Horticulturae, 293:110674. doi: 10.1016/j.scienta.2021.110674 URL
[96]	Wang Y G, Zhou L J, Wang Y X, Geng Z Q, Liu S H, Chen C W, Chen S M, Jiang J F, Chen F D. 2022c. CmMYB9a activates floral coloration by positively regulating anthocyanin biosynthesis in chrysanthemum. Plant Molecular Biology, 108:51-63. doi: 10.1007/s11103-021-01206-z URL
[97]	Wang Y J, Sheng L P, Zhang H R, Du X P, An C, Xia X L, Chen F D, Jiang J F, Chen S M. 2017. CmMYB19 over-expression improves aphid tolerance in chrysanthemum by promoting lignin synthesis. International Journal of Molecular Sciences, 18:619. doi: 10.3390/ijms18030619 URL
[98]	Wei Q, Ma C, Xu Y J, Wang T L, Chen Y Y, Lv J, Zhang L L, Jiang C Z, Hong B, Gao J P. 2017. Control of chrysanthemum flowering through integration with an aging pathway. Nature Communications, 8:829. doi: 10.1038/s41467-017-00812-0 pmid: 29018260
[99]	Wei Z Q, Lv M J, Wan W, Yu F, Cao X Y, Meng L S. 2019. Transformation of eIF5B1 gene into chrysanthemum to gain calluses of high temperature tolerance. Biologia, 74:1271-1277. doi: 10.2478/s11756-019-00312-0 URL
[100]	Wen L, Sun X, Ren H, Fan H, Yu Y, Guo Y, Zheng C. 2017. Cloning and functional verification of chrysanthemum CmXTHs genes related to petal elongation. Acta Horticulturae Sinica, 44:2150-2162.
[101]	Wen X H, Li Z J, Wang L L, Lu C F, Gao Q, Xu P, Pu Y, Zhang Q L, Hong Y, Hong L, Huang H, Xin H, Wu X Y, Kang D R, Gao K, Li Y J, Ma C F, Li X M, Zheng H K, Wang Z C, Jiao Y N, Zhang L S, Dai S L. 2022. The Chrysanthemum lavandulifolium genome and the molecular mechanism underlying diverse capitulum types. Horticulture Research, 9:uhab022. doi: https://doi.org/10.1093/hr/uhab022. doi: https://doi.org/10.1093/hr/uhab022 URL
[102]	Wen X H, Qi S, Yang L W, Hong Y, Dai S L. 2019. Expression pattern of candidate genes in early capitulum morphogenesis of Chrysanthemum lavandulifolium. Scientia Horticulturae, 252:332-341. doi: 10.1016/j.scienta.2019.03.064 URL
[103]	Wu Jian, Yang Yan-fang, Wang Hui, Zhu Kai, Liu Li-qing, Chen Fa-di, Yu De-yue. 2016. Study of DmDREBa/b gene enhanced stress tolerance in transgenic tobacco. Molecular Plant Breeding, 14:3063-3072. (in Chinese)
	武剑, 杨艳芳, 王慧, 朱凯, 刘黎卿, 陈发棣, 喻德跃. 2016. 菊花DmDREBa/b基因提高转基因烟草耐逆性研究. 分子植物育种, 14:3063-3072.
[104]	Xi L, Wen C, Fang S, Chen X L, Nie J, Chu J F, Yuan C Q, Yan C Y, Ma N, Zhao L J. 2015. Impacts of strigolactone on shoot branching under phosphate starvation in chrysanthemum(Dendranthema grandiflorum cv.Jinba). Frontiers in Plant Science, 6:694.
[105]	Xia Wei-kang. 2020. Functional analysis of transcription factors CmCUC2and CmCUC3 in chrysanthemum[M. D. Dissertation]. Nanjing: Nanjing Agricultural University. (in Chinese)
	夏伟康. 2020. 菊花转录因子CmCUC2和CmCUC3的功能分析[硕士论文]. 南京: 南京农业大学.
[106]	Xiang L L, Liu X F, Li H, Yin X R, Grierson D, Li F, Chen K S. 2019. CmMYB#7,an R3 MYB transcription factor,acts as a negative regulator of anthocyanin biosynthesis in chrysanthemum. Journal of Experimental Botany, 70:3111-3123. doi: 10.1093/jxb/erz121 pmid: 30994176
[107]	Xin Jing-jing. 2020. Molecular mechanisms of CmMLO17 involved in Alternaria alternata infection[Ph. D. Dissertation]. Nanjing: China Agricultural University. (in Chinese)
	辛静静. 2020. 菊花CmMLO17参与链格孢菌侵染的分子机制[博士论文]. 南京: 南京农业大学.
[108]	Xing Li-li, Chen Fa-di, Miao Heng-bin. 2009. Meiosis and ISSR analysis on genetic variation of ⁶⁰Co γ-rays irradiated variants of cut chrysanthemum. Journal of Nuclear Agricultural Sciences, 23:587-591. (in Chinese)
	邢莉莉, 陈发棣, 缪恒彬. 2009. 切花菊'长紫'辐照后代减数分裂行为及ISSR遗传变异分析. 核农学报, 23:587-591. doi: 10.11869/hnxb.2009.04.0587
[109]	Xing Xiao-juan. 2019. The lateral bud development related candidate genes research and functional verfication of inhibition of high temperature on verification inhibition og high temperature on cut-flower chrysanthemum'Sei No Issei'[Ph. D. Dissertation]. Nanjing: Nanjing Agricultural University. (in Chinese)
	邢晓娟. 2019. 高温抑制切花菊'精の一世'侧枝发育相关候选基因挖掘与功能验证[博士论文]. 南京: 南京农业大学.
[110]	Xu Ting-ting, Ma-Jie, Chi Tian-hua, Ye Dan, Guan Zhi-yong, Fang Wei-min, Chen Fa-di, Zhang Fei. 2019. Genetic variation and molecular markers for cold tolerance of cut-chrysanthemum cultivars at seedling stage. Journal of Nuclear Agricultural Sciences, 33:199-207. (in Chinese)
	徐婷婷, 马杰, 迟天华, 叶丹, 管志勇, 房伟民, 陈发棣, 张飞. 2019. 切花菊苗期抗寒性评价及相关的分子标记挖掘. 核农学报, 33:199-207. doi: 10.11869/j.issn.100-8551.2019.01.0199
[111]	Xu Y J, Zhao X, Aiwaili P, Mu X Y, Zhao M, Zhao J, Cheng L N, Ma C, Gao J P, Hong B. 2020. A zinc finger protein BBX19 interacts with ABF3 to affect drought tolerance negatively in chrysanthemum. The Plant Journal, 103:1783-1795. doi: 10.1111/tpj.14863 URL
[112]	Xue Shou-ji. 2004. Coloratlas of the chrysanthemum of China. Beijing: China Forestry Press:1-5. (in Chinese)
	薛守纪. 2004. 中国菊花图谱. 北京: 中国林业出版社:1-5.
[113]	Yang Hui-ting. 2019. Induced ploidy variations in vitro of cutting white chrysanthemum and rapid propagation in vitro of yellow chrysanthemum 'Jinshan'[M. D. Dissertation]. Fuzhou: Fujian Agriculture and Forestry University. (in Chinese)
	杨惠婷. 2019. 单头切花白菊的离体倍性诱变与'金扇'的组培快繁[硕士论文]. 福州: 福建农林大学.
[114]	Yang Xin-cheng, Su Jiang-shuo, Sun Wei, Wu Yang-yang, Zhang Fei, Guan Zhi-yong, Chen Fa-di, Yao Jian-jun, Fang Wei-min. 2018. Combining ability analysis of the major branching traits in spray cut chrysanthemum. Acta Horticulturae Sinica, 45 (10):1952-1960. (in Chinese)
	杨信程, 苏江硕, 孙炜, 吴洋洋, 张飞, 管志勇, 陈发棣, 姚建军, 房伟民. 2018. 切花小菊主要分枝性状的配合力与遗传力分析. 园艺学报, 45 (10):1952-1960.
[115]	Yang X D, Fang X Q, Su J S, Ding L, Guan Z Y, Jiang J F, Chen S M, Chen F D, Fang W M, Zhang F. 2019a. Genetic dissection of floral traits in anemone-type chrysanthemum by QTL mapping. Molecular Breeding, 39:1-11. doi: 10.1007/s11032-018-0907-x URL
[116]	Yang X D, Wu Y Y, Su J S, Ao N, Guan Z Y, Jiang J F, Chen S M, Fang W M, Chen F D, Zhang F. 2019b. Genetic variation and development of a SCAR marker of anemone-type flower in chrysanthemum. Molecular Breeding, 39:1-12. doi: 10.1007/s11032-018-0907-x URL
[117]	Yang Y J, Ma C, Xu Y J, Wei Q, Imtiaz M, Lan H B, Gao S, Cheng L N, Wang M Y, Fei Z J, Hong B, Gao J P. 2014. A zinc finger protein regulates flowering time and abiotic stress tolerance in chrysanthemum by modulating gibberellin biosynthesis. The Plant Cell, 26:2038-2054. doi: 10.1105/tpc.114.124867 pmid: 24858937
[118]	Yang Yi-ru. 2020. Cloning and expression analysis of CgDREB22 gene from Chrysanthemum × grandiflora[M. D. Dissertation]. Harbin: Northeast Forestry University. (in Chinese)
	杨伊如. 2020. 露地菊CgDREB22基因克隆与表达分析[硕士论文]. 哈尔滨: 东北林业大学.
[119]	Yuan C Q, Ahmad S, Cheng T R, Wang J, Pan H T, Zhao L J, Zhang Q X. 2018. Red to far-red light ratio modulates hormonal and genetic control of axillary bud outgrowth in Chrysanthemum(Dendranthema grandiflorum'Jinba'). International Journal of Molecular Sciences, 19:1590. doi: 10.3390/ijms19061590 URL
[120]	Yuan C Q, Huang D, Yang Y, Sun M, Cheng T R, Wang J, Pan H, Zhang Q X. 2020. CmCYC2-like transcription factors may interact with each other or bind to the promoter to regulate floral symmetry development in Chrysanthemum morifolium'. Plant Molecular Biology, 103:159-171. doi: 10.1007/s11103-020-00981-5 URL
[121]	Yuan P S, Qian S Y, Zhai Z, FernánMartínez J, Xu H L. 2022. Study of chrysanthemum image phenotype on-line classification based on transfer learning and bilinear convolutional neural network. Computers and Electronics in Agriculture, 194:106679. doi: 10.1016/j.compag.2021.106679 URL
[122]	Yuan Pei-sen, Li Wei, Ren Shou-gang, Xu Huan-liang. 2018. Recognition for flower type and variety of chrysanthemum with convolutional neural network. Transactions of the Chinese Society of Agricultural Engineering, 34 (5):152-158. (in Chinese)
	袁培森, 黎薇, 任守纲, 徐焕良. 2018. 基于卷积神经网络的菊花花型和品种识别. 农业工程学报, 34 (5):152-158.
[123]	Zhai Guo, Li Zhi-min, Lu Wen-chao, Zhao Yong, Wang Cheng. 2016. Study on varieties identification of ornamental chrysanthemum based on image processing. Journal of Chinese Agricultural Mechanization, 37:105-110. (in Chinese)
	翟果, 李志敏, 路文超, 赵勇, 王成. 2016. 基于图像处理技术的观赏菊品种识别方法研究. 中国农机化学报, 37:105-110.
[124]	Zhang L L, Xu Y J, Liu X N, Qin M Z, Li S L, Jiang T H, Yang Y J, Jiang C Z, Gao J P, Hong B, Ma C. 2022. The chrysanthemum DEAD-box RNA helicase CmRH 56 regulates rhizome outgrowth in response to drought stress. Journal of Experimental Botany,erac213. doi: https://doi.org/10.1093/jxb/erac213. doi: https://doi.org/10.1093/jxb/erac213 URL
[125]	Zhang Fei, Chen Fa-di, Fang Wei-min, Chen Su-mei, Li Feng-tong. 2010a. Heterosis and mixed genetic analysis of inflorescence traits of chrysanthemum. Scientia Agricultura Sinica, 43:2953-2961. (in Chinese)
	张飞, 陈发棣, 房伟民, 陈素梅, 李风童. 2010a. 菊花花器性状杂种优势与混合遗传分析. 中国农业科学, 43:2953-2961.
[126]	Zhang Fei, Fang Wei-min, Chen Fa-di, Chen Su-mei. 2010b. Combining ability analysis on ornamental characters of chrysanthemum. Acta Horticulturae Sinica, 37 (4):589-596. (in Chinese)
	张飞, 房伟民, 陈发棣, 陈素梅. 2010b. 菊花观赏性状的配合力分析. 园艺学报, 37 (4):589-596.
[127]	Zhang F, Chen S M, Chen F D, Fang W M, Chen Y, Li F T. 2011. SRAP-based mapping and QTL detection for inflorescence-related traits in chrysanthemum(Dendranthema morifolium). Molecular Breeding, 27:11-23. doi: 10.1007/s11032-010-9409-1 URL
[128]	Zhang F, Chen S M, Chen F D, Fang W M, Li F T. 2010. A preliminary genetic linkage map of chrysanthemum(Chrysanthemum morifolium) cultivars using RAPD,ISSR and AFLP markers. Scientia Horticulturae, 125:422-428. doi: 10.1016/j.scienta.2010.03.028 URL
[129]	Zhang F, Chen S M, Jiang J F, Guan Z Y, Fang W M, Chen F D. 2013. Genetic mapping of quantitative trait loci underlying flowering time in chrysanthemum(Chrysanthemum morifolium). PLoS ONE, 8:e83023.
[130]	Zhang Hong-wei. 2015. Chrysanthemum coloratlas. Zhengzhou: Zhongzhou Ancient Books Press:1-20. (in Chinese)
	章宏伟. 2015. 菊谱. 郑州: 中州古籍出版社:1-20.
[131]	Zhang Shu-lin, Dai Si-lan. 2013. The book of chrysanthemum in China. Beijing: China Forestry Publishing House. (in Chinese)
	张树林, 戴思兰. 2013. 中国菊花全书. 北京: 中国林业出版社.
[132]	Zhang Z H, Zhu L, Song A P, Wang H B, Chen S M, Jiang J F, Chen F D. 2020. Chrysanthemum (Chrysanthemum morifolium)CmICE2conferred freezing tolerance in Arabidopsis. Plant Physiology and Biochemistry, 146:31-41. doi: 10.1016/j.plaphy.2019.10.041 URL
[133]	Zhang Z X, Hu Q, Cheng H, Cheng P L, Liu Y N, Liu W X, Xing X J, Chen S M, Chen F D, Jiang J F. 2019. A single residue change in the product of the chrysanthemum gene TPL1-2 leads to a failure in its repression of flowering. Plant Science, 285:165-174. doi: 10.1016/j.plantsci.2019.04.027 URL
[134]	Zhao K K, Ding L, Xia W K, Zhao W Q, Zhang X, Jiang J F, Chen S M, Chen F D. 2020. Characterization of an APETALA1 and a FRUITFUL-like homolog in chrysanthemum. Scientia Horticulturae, 272:109518. doi: 10.1016/j.scienta.2020.109518 URL
[135]	Zhao Q, He L, Wang B, Liu Q L, Pan Y Z, Zhang F, Jiang B B, Zhang L. 2019. Overexpression of a multiprotein bridging factor 1 gene DgMBF1 improves the salinity tolerance of chrysanthemum. International Journal of Molecular Sciences, 20:2453. doi: 10.3390/ijms20102453 URL
[136]	Zhao Q, Zhong M, He L, Wang B, Liu Q L, Pan Y Z, Jiang B B, Zhang L. 2018. Overexpression of a chrysanthemum transcription factor gene DgNAC1 improves drought tolerance in chrysanthemum. Plant Cell,Tissue and Organ Culture(PCTOC), 135:119-132.
[137]	Zhao W Q, Ding L, Liu J Y, Zhang X, Li S, Zhao K K, Guan Y X, Song A P, Wang H B, Chen S M, Jiang J F, Chen F D. 2022. Regulation of lignin biosynthesis by an atypical bHLH protein CmHLB in Chrysanthemum. Journal of Experimental Botany, 73 (8):2403-2419. doi: 10.1093/jxb/erac015 URL
[138]	Zhou L J, Geng Z Q, Wang Y X, Wang Y G, Liu S H, Chen C W, Song A P, Jiang J F, Chen S M, Chen F D. 2021. A novel transcription factor CmMYB 012 inhibits flavone and anthocyanin biosynthesis in response to high temperatures in chrysanthemum. Horticulture Research, 8:248. doi: 10.1038/s41438-021-00675-z pmid: 34848687
[139]	Zhou L J, Wang Y X, Wang Y G, Song A P, Jiang J F, Chen S M, Ding B Q, Chen F D. 2022. Transcription factor CmbHLH 16 regulates petal anthocyanin homeostasis under different lights in Chrysanthemum. Plant Physiology,doi: https://doi.org/10.1093/plphys/kiac342. doi: https://doi.org/10.1093/plphys/kiac342 URL
[140]	Zhu Kai, Wu Jian, Yang Yan-fang, Chen Fa-di, Yu De-yue. 2017. Overexpression of DlNAC1 gene isolated from Dendranthema lavandulifolium in enhancing heat stress tolerance in transgenic tobacco. Bulletin of Botanical Research, 37:432-439. (in Chinese)
	朱凯, 武剑, 杨艳芳, 陈发棣, 喻德跃. 2017. 甘菊DlNAC1转录因子基因提高烟草耐高温能力. 植物研究, 37:432-439.

性状 Trait	基因 Gene	供体 Used for transgene isolated	受体 For genetic transformation	获得性状 Changed trait	文献 Reference
花色 Flower color	CmMYB#7	'神马''Jinba'	'神马''Jinba'	负调控花色苷合成Negative regulator of anthocyanin biosynthesis	Xiang et al.,2019
	CmMYB012	'南农粉翠' 'Nannong Fencui'	'南农粉翠' 'Nannong Fencui'	花色变浅 Pink flower color becomes lighter	Zhou et al.,2021
	CmMYB21	'粉安娜' 'Anastasia Pink'	烟草 Tobacco	花色变浅Pink flower color becomes lighter	Wang et al.,2022b
	CmMYB9a	'粉安娜' 'Anastasia Pink'	烟草 Tobacco	花色加深 Pink flower color becomes darker	Wang et al.,2022c
	CmbHLH16	'南农粉翠' 'Nannong Fencui'	'南农粉翠' 'Nannong Fencui'	在不同红光和远红光比例下调控花色 Regulates flower color under different red and far-red lights	Zhou et al.,2022
花型 Flower shape	CmXTH1-4	'杭白菊''Hangbaiju'	'杭白菊''Hangbaiju'	促进舌状花花瓣伸长 Promote elongation of ray florets	Wen et al.,2017
	CmTCP20	'神马''Jinba'	'神马''Jinba'	促进舌状花花瓣伸长 Promote elongation of ray florets	Wang et al.,2019
	CmYAB1	'神马''Jinba'	'神马''Jinba'	改变舌状花翻卷程度和花型 Changes in petal curvature and inflorescence morphology	Ding et al.,2019
	CmCYC2a-2f	'毛香玉''Maoxiangyu'	甘菊 Chrysanthemun lavandulifolium	调控舌状花形成和花对称性 Regulation of ray florets formation and flower symmetry	Yuan et al.,2020
	CmBES1;CmCUC2/ 3	'神马''Jinba'	'神马''Jinba'	影响舌状花融合程度Affection in merged degree of the ray florets	Cheng et al.,2020; 夏伟康,2020
株型 Plant	CmERF053	'神马''Jinba'	拟南芥 Arabidopsis	促进分枝和侧根形成 Promote branching and lateral root formation	Nie et al.,2018
architecture	CmMAX1	'神马''Jinba'	拟南芥 Arabidopsis	减少分枝数Reduce branch numbers	Dong et al.,2018
	CmSWEET17	'神马''Jinba'	'神马''Jinba'	促进腋芽生长Promote axillary bud outgrowth	刘伟鑫,2019
	CmHLB,CmKNAT7	'神马''Jinba'	'神马''Jinba'	植株矮化,茎秆增粗 Dwarf plants,thickened stems	Zhao et al.,2022
	CmDIV	'毛香玉''Maoxiangyu'	拟南芥 Arabidopsis	植株较低矮,叶片小且圆 Plants become shorter,with smaller and rounder leaves	刘轶奇,2016

性状 Trait	基因 Gene	供体 Used for transgene isolated	受体 For genetic transformation	获得性状 Changed trait	文献 Reference
花色 Flower color	CmMYB#7	'神马''Jinba'	'神马''Jinba'	负调控花色苷合成Negative regulator of anthocyanin biosynthesis	Xiang et al.,2019
	CmMYB012	'南农粉翠' 'Nannong Fencui'	'南农粉翠' 'Nannong Fencui'	花色变浅 Pink flower color becomes lighter	Zhou et al.,2021
	CmMYB21	'粉安娜' 'Anastasia Pink'	烟草 Tobacco	花色变浅Pink flower color becomes lighter	Wang et al.,2022b
	CmMYB9a	'粉安娜' 'Anastasia Pink'	烟草 Tobacco	花色加深 Pink flower color becomes darker	Wang et al.,2022c
	CmbHLH16	'南农粉翠' 'Nannong Fencui'	'南农粉翠' 'Nannong Fencui'	在不同红光和远红光比例下调控花色 Regulates flower color under different red and far-red lights	Zhou et al.,2022
花型 Flower shape	CmXTH1-4	'杭白菊''Hangbaiju'	'杭白菊''Hangbaiju'	促进舌状花花瓣伸长 Promote elongation of ray florets	Wen et al.,2017
	CmTCP20	'神马''Jinba'	'神马''Jinba'	促进舌状花花瓣伸长 Promote elongation of ray florets	Wang et al.,2019
	CmYAB1	'神马''Jinba'	'神马''Jinba'	改变舌状花翻卷程度和花型 Changes in petal curvature and inflorescence morphology	Ding et al.,2019
	CmCYC2a-2f	'毛香玉''Maoxiangyu'	甘菊 Chrysanthemun lavandulifolium	调控舌状花形成和花对称性 Regulation of ray florets formation and flower symmetry	Yuan et al.,2020
	CmBES1;CmCUC2/ 3	'神马''Jinba'	'神马''Jinba'	影响舌状花融合程度Affection in merged degree of the ray florets	Cheng et al.,2020; 夏伟康,2020
株型 Plant	CmERF053	'神马''Jinba'	拟南芥 Arabidopsis	促进分枝和侧根形成 Promote branching and lateral root formation	Nie et al.,2018
architecture	CmMAX1	'神马''Jinba'	拟南芥 Arabidopsis	减少分枝数Reduce branch numbers	Dong et al.,2018
	CmSWEET17	'神马''Jinba'	'神马''Jinba'	促进腋芽生长Promote axillary bud outgrowth	刘伟鑫,2019
	CmHLB,CmKNAT7	'神马''Jinba'	'神马''Jinba'	植株矮化,茎秆增粗 Dwarf plants,thickened stems	Zhao et al.,2022
	CmDIV	'毛香玉''Maoxiangyu'	拟南芥 Arabidopsis	植株较低矮,叶片小且圆 Plants become shorter,with smaller and rounder leaves	刘轶奇,2016

基因 Gene	供体Used for transgene isolated		受体For genetic transformation	获得性状 Changed trait		途径 Pathway		文献 Reference
DenFUL	甘菊Chrysanthemun lavandulifolium		拟南芥 Arabidopsis	提前开花 Early-flowering		不详Unknown		王翊,2013
CmBBX24	'Fall Color'		'Fall Color'	延迟开花Late-flowering		光周期和赤霉素途径 Photoperiod and GA pathways		Yang et al.,2014
AtCO	拟南芥 Arabidopsis		'火焰''Huoyan'	提前开花Early-flowering		光周期途径 Photoperiod pathway		宋凤艳,2016
CmFTL1	'神马''Jinba'		'神马''Jinba'	提前开花Early-flowering		光周期途径 Photoperiod pathway		Mao et al.,2016
CmFTL2	'优香''Yuuka'		'优香''Yuuka'	提前开花Early-flowering		光周期途径 Photoperiod pathway		Sun et al.,2017
CmNF-YB8	'Fall Color'		'Fall Color'	延迟开花Late-flowering		年龄途径 Aging pathway		Wei et al.,2017
ClPHYB	甘菊Chrysanthemun lavandulifolium		拟南芥 Arabidopsis	推迟开花Late-flowering		光周期途径 Photoperiod pathway		马朝峰,2019
CmBBX13	'优香''Yuuka'		拟南芥 Arabidopsis	推迟开花Late-flowering		独立于光周期途径 Independently of the photoperiod pathway		Ping et al.,2019
CmTPL1-2	'神马''Jinba'		'神马''Jinba'	推迟开花Late-flowering		不详Unknown		Zhang et al.,2019
CmBBX8	'优香''Yuuka'		'优香''Jinba'	提前开花Early-flowering		光周期途径 Photoperiod pathway		Wang et al.,2020b
CmAP1L1	'神马''Jinba'		'神马''Jinba'	提前开花Early-flowering		不详Unknown		Zhao et al.,2020
CmJAZ1-like	'神马''Jinba'	'神马''Jinba'			延迟开花Late-flowering		不详Unknown	Guan et al.,2021
CmERF110,CmFLK	'神马''Jinba'	'神马''Jinba'			提前开花Early-flowering		自主途径Autonomous pathway	Huang et al.,2022a

基因 Gene	供体Used for transgene isolated		受体For genetic transformation	获得性状 Changed trait		途径 Pathway		文献 Reference
DenFUL	甘菊Chrysanthemun lavandulifolium		拟南芥 Arabidopsis	提前开花 Early-flowering		不详Unknown		王翊,2013
CmBBX24	'Fall Color'		'Fall Color'	延迟开花Late-flowering		光周期和赤霉素途径 Photoperiod and GA pathways		Yang et al.,2014
AtCO	拟南芥 Arabidopsis		'火焰''Huoyan'	提前开花Early-flowering		光周期途径 Photoperiod pathway		宋凤艳,2016
CmFTL1	'神马''Jinba'		'神马''Jinba'	提前开花Early-flowering		光周期途径 Photoperiod pathway		Mao et al.,2016
CmFTL2	'优香''Yuuka'		'优香''Yuuka'	提前开花Early-flowering		光周期途径 Photoperiod pathway		Sun et al.,2017
CmNF-YB8	'Fall Color'		'Fall Color'	延迟开花Late-flowering		年龄途径 Aging pathway		Wei et al.,2017
ClPHYB	甘菊Chrysanthemun lavandulifolium		拟南芥 Arabidopsis	推迟开花Late-flowering		光周期途径 Photoperiod pathway		马朝峰,2019
CmBBX13	'优香''Yuuka'		拟南芥 Arabidopsis	推迟开花Late-flowering		独立于光周期途径 Independently of the photoperiod pathway		Ping et al.,2019
CmTPL1-2	'神马''Jinba'		'神马''Jinba'	推迟开花Late-flowering		不详Unknown		Zhang et al.,2019
CmBBX8	'优香''Yuuka'		'优香''Jinba'	提前开花Early-flowering		光周期途径 Photoperiod pathway		Wang et al.,2020b
CmAP1L1	'神马''Jinba'		'神马''Jinba'	提前开花Early-flowering		不详Unknown		Zhao et al.,2020
CmJAZ1-like	'神马''Jinba'	'神马''Jinba'			延迟开花Late-flowering		不详Unknown	Guan et al.,2021
CmERF110,CmFLK	'神马''Jinba'	'神马''Jinba'			提前开花Early-flowering		自主途径Autonomous pathway	Huang et al.,2022a

类型 Type	基因 Gene		供体Used for transgene isolated	受体For genetic transformation		获得性状 Changed trait	文献 Reference
非生物胁迫	DgNAC1	'神马''Jinba'			'神马''Jinba'	耐旱性Drought tolerance	Zhao et al.,2018
Abiotic stress	CmBBX19,CmABF3	'Fall Color'			'Fall Color'	耐旱性Drought tolerance	Xu et al.,2020
	CgDREB22	'纽9717''Niu9717'			烟草 Tobacco	耐旱性Drought tolerance	杨伊如,2020
	CmRH56	'Fall Color'			菊花C. morifolium	耐旱性Drought tolerance	Zhang et al.,2022
	CmSOS1,CmRCD1	'神马''Jinba'			'神马''Jinba'	耐涝性Waterlogging tolerance	Wang et al.,2020b
	DlNAC1		甘菊 Chrysanthemun lavandulifolium	烟草 Tobacco		耐热性Heat tolerance	朱凯等,2017
	eIF5B1		拟南芥 Arabidopsis	菊花 C. morifolium		耐热性Heat tolerance	Wei et al.,2019
	DgMYB1		'神马''Jinba'	'神马''Jinba'		耐寒性Cold tolerance	何玲,2019
	CmICE2		'神马''Jinba'	拟南芥 Arabidopsis		耐寒性Cold tolerance	Zhang et al.,2020
	DgC3H1		菊花 C. morifolium	菊花 C. morifolium		耐寒性Cold tolerance	Bai et al.,2021
	CmHSFA4;DgWRKY2; DgMBF1,DgJAZ1		'神马''Jinba'	'神马''Jinba'		耐盐性Salt tolerance	Li et al.,2018;He et al.,2018;Zhao et al.,2019
	ClSND1.1,ClVND1,ClBRN1		甘菊 Chrysanthemun lavandulifolium	拟南芥 Arabidopsis		耐盐性Salt tolerance	王海等,2021
	CmBBX24		'Fall Color'	'Fall Color'		耐干旱和低温 Drought and cold tolerance	Yang et al.,2014
	CdICE1		异色菊 C. dichrum	'神马''Jinba'		干旱、低温、盐 Drought,cold and salt tolerance	Chen et al.,2012
	DmDREBa/b		菊花C. morifolium	烟草 Tobacco		干旱、低温、盐 Drought,cold and salt tolerance	武剑等,2016
	CmDREBa-2		'C029'	'C029'		耐旱性和抗白锈病 Drought and white rust resistance	毕蒙蒙等,2019
	CgbZIP		'纽9717' 'Niu9717'	烟草 Tobacco		耐盐性和耐旱性 Salt and drought tolerance	Liu et al.,2022
生物胁迫 Biotic stress	CmWRKY15;CmWRKY33.1; CmMLO17,CmKIC		'神马''Jinba'	'神马''Jinba'		黑斑病抗性 Black spot disease resistance	Fan et al.,2015;Liu et al.,2020;辛静静,2020
	CmMYB19;CmTPS2/3;CmMYB15		'神马''Jinba'	'神马''Jinba'		抗蚜虫性Aphid resistance	Wang et al.,2017;王威姣,2019;An et al.,2019

Retrospection and Prospect of Chrysanthemum Genetic Breeding for Last Six Decades in China

RichHTML

PDF

Knowledge

Abstract

Cite this article

share this article

Figures/Tables 5

References 140

Related Articles 15

Recommended Articles

Metrics

Comments

[1]	YE Zhiqin and YANG Juan. A New Chrysanthemum Cultivar‘Zibinyun’ [J]. Acta Horticulturae Sinica, 2022, 49(S2): 173-174.
[2]	ZHAO Yanli, LI Zhanhong, CAO Qin, DAI Miaofei, GAO Mengmeng, SUN Zhenzhu, LI Huikuan, LI Yonghua, BIAN Shuxun, and HUANG Gan. A New Chrysanthemum Cultivar‘Bianjing Qingdianhuang’ [J]. Acta Horticulturae Sinica, 2022, 49(S2): 175-176.
[3]	CAO Qin, ZHAO Yanli, LI Zhanhong, GAO Mengmeng, DAI Miaofei, LI Yonghua, LI Huikuan, SUN Zhenzhu, BIAN Shuxun, and LI Fei. A New Muitiflora Chrysanthemum Cultivar‘Bianjing Zijingling’ [J]. Acta Horticulturae Sinica, 2022, 49(S2): 177-178.
[4]	CHU Zhuannan, LI Weiwen, DONG Ling, PENG Xingxing, CUI Guangsheng, TANG Maogui, ZHANG Qinghua, SHEN Wensheng, ZHAO Wei, XIONG Rui, HAN Piao, WU Jiqiu, LI Yulong, YANG Yueying, and ZHANG Qiling. A New Chrysanthemum morifolium Cultivar‘Gongju 1’ [J]. Acta Horticulturae Sinica, 2022, 49(S2): 179-180.
[5]	WU Fan, JIANG Junhao, LU Shan, ZHANG Nan, QIU Shuai, WEI Jianfen, SHEN Baichun. Research Progress on Germplasm Resources and Utilization of Hydrangea in China [J]. Acta Horticulturae Sinica, 2022, 49(9): 2037-2050.
[6]	YANG Xiuwei, SU Jiangshuo, ZHANG Fei, GUAN Zhiyong, FANG Weimin, CHEN Fadi. Quantitative Evaluation and Genetic Variation of Flower Opening Angles in Spray Cut Chrysanthemums [J]. Acta Horticulturae Sinica, 2022, 49(8): 1723-1734.
[7]	LÜ Zhengxin, HE Yanqun, JIA Dongfeng, HUANG Chunhui, ZHONG Min, LIAO Guanglian, ZHU Yi, YUAN Kaichang, LIU Chuanhao, XU Xiaobiao. Genetic Diversity Analysis of Phenotypic Traits for Kiwifruit Germplasm Resources [J]. Acta Horticulturae Sinica, 2022, 49(7): 1571-1581.
[8]	LI Pingping, ZHANG Xiang, LIU Yuting, XIE Zhihe, ZHANG Ruihao, ZHAO Kai, LÜ Junheng, WANG Ziran, WEN Jinfen, ZOU Xuexiao, DENG Minghua. Studies on the Relationship Between Pigment Composition and Fruit Coloration of 63 Peppers [J]. Acta Horticulturae Sinica, 2022, 49(7): 1589-1601.
[9]	LI Qiong, LI Lili, HOU Juan, LUO Renren, WANG Ruidan, HU Jianbin, HUANG Song. Advances on Mechanism of Cucurbit Crops in Response to Low- temperature Stress [J]. Acta Horticulturae Sinica, 2022, 49(6): 1382-1394.
[10]	LIU Xiaowei, XIA Bin, LI Ziwei, YANG Yujia, CHEN Bin, ZHOU Yunwei, HE Miao. Overexpression of Chrysanthemum indicum miR396a Gene in Arabidopsis Enhances Its Salt Tolerance [J]. Acta Horticulturae Sinica, 2022, 49(4): 816-826.
[11]	WANG Ying, AI Penghui, LI Shuailei, KANG Dongru, LI Zhongai, WANG Zicheng. Identification and Expression Analysis of Genes Related to DNA Methylation in Chrysanthemum × morifolium and C. nankingense [J]. Acta Horticulturae Sinica, 2022, 49(4): 827-840.
[12]	ZHANG Shicai, LI Yifei, WANG Chunping, YANG Xiaomiao, HUANG Qizhong, HUANG Renzhong. Resistance Identification and Evaluation of Pepper Germplasm to Colletotrichum acutatum [J]. Acta Horticulturae Sinica, 2022, 49(4): 885-892.
[13]	LI Junzhang, QIN Yuan, XIAO Qiang, AN Chang, LIAO Jingyi, ZHENG Ping. Recent Advances in Molecular Biology of Crassulacean Acid Metabolism Plants and the Application Potential of CAM Engineering [J]. Acta Horticulturae Sinica, 2022, 49(12): 2597-2610.
[14]	YANG Sichao, ZHANG Meng, ZHANG Qinglin, LUO Zhengrong. Advances on Natural Deastringency Characteristics and Molecular Mechanism of Chinese PCNA Persimmon [J]. Acta Horticulturae Sinica, 2022, 49(12): 2659-2668.
[15]	XU Wan, LIN Yajun, ZHAO Zhuang, ZHOU Zhuang. Advances in Genetic Resources and Breeding Research of Cymbidium [J]. Acta Horticulturae Sinica, 2022, 49(12): 2722-2742.