Identification of Cowpea Mild Mottle Virus Variants in Zhejiang and Localization of TGB Proteins

doi:10.16420/j.issn.0513-353x.2023-0801

Acta Horticulturae Sinica ›› 2025, Vol. 52 ›› Issue (1): 123-135.doi: 10.16420/j.issn.0513-353x.2023-0801

• Genetic & Breeding·Germplasm Resources·Molecular Biology • Previous Articles Next Articles

Identification of Cowpea Mild Mottle Virus Variants in Zhejiang and Localization of TGB Proteins

ZHANG Zixin, CHEN Shuting, ZHOU Weixin, ZHANG Yihan, LU Qian, LI Zhongyi, WU Xinyang, XU Pei^**()

Key Laboratory of Specialty Agri-Product Quality and Hazard Controlling Technology of Zhejiang，College of Life Sciences，China Jiliang University，Hangzhou 310018，China

Received:2024-08-21 Revised:2024-10-18 Online:2025-01-25 Published:2025-01-19
Contact: XU Pei

Abstract

Abstract:

During the period from October to November 2022，an outbreak of suspected virus-infected common bean occurred in the Qiantang District of Hangzhou City，Zhejiang Province. Subsequent viral genome sequencing and PCR confirmed the presence of cowpea mild mottle virus（CPMMV），cucumber mosaic virus（CMV），milk vetch dwarf virus（MDV）. CPMMV was designated as CPMMV-ZJ（GenBank：OR667247）. By virus identification，genome sequence amplification，phylogenetic analysis and subcellular localization analysis，the sequence structure of CPMMV-ZJ and the localization of the triple gene block proteins（TGB）were elucidated. The genome of CPMMV-ZJ encompassed a total length of 8 226 bp，consisting of six open reading frames. Additionally，both the 3′ and 5′ regions contained non-coding regions（UTRs）of 147 and 75 bp，respectively. Phylogenetic analysis of CPMMV-ZJ revealed the highest sequence identity（98.03%）with the CPMMV Anhui isolate，whereas lower identities（63.71%-82.49%）were observed when comparing to foreign isolates from the United States，India，and Brazil，indicating virus evolution is related to geographical factors. Subcellular localization analysis demonstrated similar localization of TGBs between the Zhejiang and Jiangsu isolates. Notably，overexpression TGB2 of the Zhejiang isolate leads to a clustering of chloroplasts around the nuclei，suggesting its potential involvement in plant defense responses.

Key words: common bean, cowpea mild mottle virus, sequence analysis, subcellular localization, clustering of chloroplasts around the nuclei

ZHANG Zixin, CHEN Shuting, ZHOU Weixin, ZHANG Yihan, LU Qian, LI Zhongyi, WU Xinyang, XU Pei. Identification of Cowpea Mild Mottle Virus Variants in Zhejiang and Localization of TGB Proteins[J]. Acta Horticulturae Sinica, 2025, 52(1): 123-135.

Figures/Tables 9

Table 1 Primers sequences used in this study

用途 Usage	引物 Primer	引物序列（5′-3′） Primers sequence
5′端快速cDNA扩增 5′RACE	5AP	GGTCTCAAGGGCTCTAAACATTT
	GSP1	CAACTCATGACCAGCTCAGC
	GSP2	TTCCCTTGGCAGAACGAGAC
序列扩增 Sequence amplification	CPMMV-F1	GTTTACCCACCGGAGTTGCT
	CPMMV-R2	AATCACTTTGGCAGGGATCC
	CPMMV-F2	TCAAAGCTAGAATGTCTAGG
序列扩增；病毒检测 Sequence amplification；Virus detection	CPMMV-R2	CTGTCAATACAGTTTTGGAG
序列扩增；病毒检测 Sequence amplification；Virus detection	CPMMV-F3	AGATGAAGGAATTAGAGAGG
序列扩增 Sequence amplification	CPMMV-R3	AGAACCTCAGCTACGTGCTC
3′端快速cDNA扩增 3′RACE	3AP	GTTTTCCCAGTCACGACAC
	GSP3	GCCCTTCCAGACATTGATGC
	GSP4	TCAAATAACATGGCCACAGCTG
亚细胞定位载体构建 Subcellular localization vector construction	TGB1-F	ATGAATGAACTGATCAGTAA
	TGB1-R	CTCAGAGTTTGGATAGGTTG
	TGB2-F	ATGCCACTGACTCCACCA
	TGB2-R	GTGAACCCTATTGCAGA
	TGB3-F	ATGTCTGCAATAGGGTTCAC
	TGB3-R	CAACCTACAACTTAGGCTA
病毒检测 Virus detection	MDV-F	ACTCAAGGAGAGGCAAGAGC
	MDV-R	ATGTCGGCTGTCTTTCCACC
	CMV-F	AAGTGGTTTGCAGCGTTGAC
	CMV-R	TGTTGCAAATCACGCACTCG

Fig. 1 Filed symptoms of common bean infected with virus diseases

Fig. 2 RT-PCR detection of CPMMV，CMV and MDV from samples randomly selected in field of common bean M：DL2 000 DNA marker；1-10：Ten randomly selected common bean leaf samples from virus-infected field plants

Fig. 3 Symptoms of common bean leaves mechanically inoculated with CPMMV-ZJ at 7 days post inoculation

Fig. 4 RT-PCR detection of CPMMV in common bean leaves mechanically inoculated with CPMMV-ZJ M：DL2 000 DNA marker；1-10：Common bean leaf samples inoculated with CPMMV；11：Positive control；12：Negative control

Fig. 5 Genome segmentation amplification strategies for CPMMV-ZJ

Fig. 6 Genomic structure of the CPMMV Zhejiang isolate（CPMMV-ZJ）from common bean

Fig. 7 Phylogenetic analysis of CPMMV-ZJ The numbers on the branches of the tree represent bootstraps

Fig. 8 Subcellular localization analysis of TGBs of CPMMV Zhejiang（CPMMV-ZJ）and Jiangsu（CPMMV-JS）isolate（A）and the clustering of chloroplasts around the nuclei caused by TGB2-ZJ（B） White arrows represents perinuclear aggregation of chloroplasts

References 41

[1]	Barreto da Silva F, Muller C, Bello V H, Watanabe L F M, Rossitto de Marchi B, Fusco L M, Ribeiro-Junior M R, Minozzi G B, Vivan L M, Tamai M A, Farias J R, Nogueira A M, Sartori M M P, Krause-Sakate R. 2020. Effects of cowpea mild mottle virus on soybean cultivars in Brazil. PeerJ, 8:e9828.
[2]	Brito M, Fernández-Rodríguez T, Garrido M J, Mejías A, Romano M, Marys E. 2012. First report of cowpea mild mottle Carlavirus on Yardlong Bean(Vigna unguiculata subsp. sesquipedalis)in Venezuela. Viruses, 4 (12):3804-3811.
[3]	Brunt A A, Kenten R H. 1973. Cowpea mild mottle,a newly recognized virus infecting cowpeas(Vigna unguiculata)in Ghana. Annals of Applied Biology, 74 (1):67-74. pmid: 4701691
[4]	Caplan J L, Mamillapalli P, Burch-Smith T M, Czymmek K, DineshKumar S P. 2008. Chloroplastic protein NRIP 1 mediates innate immune receptor recognition of a viral effector. Cell, 132 (3):449-462.
[5]	Caplan J L, Kumar A S, Park E, Padmanabhan M S, Hoban K, Modla S, Dinesh-Kumar S P. 2015. Chloroplast Stromules function during innate immunity. Developmental Cell, 34 (1):45-57. doi: 10.1016/j.devcel.2015.05.011 pmid: 26120031
[6]	Carvalho S L, Tilsner J, Figueira A R, Carvalho C M. 2022. Subcellular localization and interactions among TGB proteins of cowpea mild mottle virus. Archives of Virology, 167 (12):2555-2566. doi: 10.1007/s00705-022-05576-7 pmid: 36269413
[7]	Chang C A, Chien L Y, Tsai C F, Lin Y Y, Cheng Y H. 2013. First report of cowpea mild mottle virus in cowpea and French bean in Taiwan. Plant Disease, 97 (7):1001.
[8]	Chiquito-Almanza E, Caballero-Pérez J, Guevara-Olvera L, Acosta-García G, Pérez-Pérez M C I, Acosta-Gallegos J A, Anaya-López J L. 2018. First report of cowpea mild mottle virus infecting cultivated and wild phaseolus in the central-western region of Mexico. Plant Disease, 102 (5):1047-1047.
[9]	Cui Tiantian, Wang Yanjiao, Li Zhongan, Zhou Changyong, Song Zhen. 2017. Bioinformatics and subcellular localization on citrus yellow vein clearing virus-Triple Gene Block. Acta Horticulturae Sinica, 44 (8):1579-1588. (in Chinese) doi: 10.16420/j.issn.0513-353x.2017-0165 URL
	崔甜甜, 王艳娇, 李中安, 周常勇, 宋震. 2017. 柑橘黄化脉明病毒的TGB基因生物信息学分析及亚细胞定位. 园艺学报, 44 (8):1579-1588.
[10]	Ding X, Jimenez-Gongora T, Krenz B, Lozano-Duran R. 2019. Chloroplast clustering around the nucleus is a general response to pathogen perception in Nicotiana benthamiana. Molecular Plant Pathology, 20 (9):1298-1306. doi: 10.1111/mpp.12840 pmid: 31257720
[11]	Gambley C, Nimmo P, Persley D, Steele V, Sharman M, Campbell P. 2022. Cowpea mild mottle virus, a sometimes problem for French bean crops. Australasian Plant Pathology, 51 (6):565-576.
[12]	Han K, Zheng H, Ji M, Cui W, Hu S, Peng J, Zhao J, Lu Y, Lin L, Liu Y, Chen J, Yan F. 2020. A single amino acid in coat protein of Pepper mild mottle virus determines its subcellular localization and the chlorosis symptom on leaves of pepper. The Journal of General Virology, 101 (5):565-570.
[13]	Hanson M R, Conklin P L. 2020. Stromules,functional extensions of plastids within the plant cell. Current Opinion in Plant Biology, 58:25-32.
[14]	Iizuka N, Rajeshwari R, Reddy D V R, Goto T, Muniyappa V, Bharathan N, Ghanekar A M. 1984. Natural occurrence of a strain of cowpea mild mottle virus on groundnut(Arachis hypogaea)in India. Journal of Phytopathology, 109 (3):245-253.
[15]	Ji Ying, Sun Feng, Wu Shuhua, Li Shuo, Tu Liqin, Gao Danna, Cui Xiaoyan, Chen Xin, Ji Yinghua, Guo Qingyun. 2022. Genomic sequence cloning and the vharacteristics analysis of cowpea mild mottle virus infecting soybean from Jiangsu Province. Acta Agriculturae Boreali-Sinica, 37 (3):200-205.. (in Chinese) doi: 10.7668/hbnxb.20193080
	吉颖, 孙枫, 吴淑华, 李硕, 涂丽琴, 高丹娜, 崔晓艳, 陈新, 季英华, 郭青云. 2022. 江苏大豆上分离的豇豆轻斑驳病毒基因组序列克隆及特征分析. 华北农学报, 37 (3):200-205. doi: 10.7668/hbnxb.20193080
[16]	Jiang C, Shan S Q, Huang Y, Mao C Y, Zhang H H, Li Y J, Chen J P, Wei Z Y, Sun Z T. 2022. The c-terminal transmembrane domain of cowpea mild mottle virus TGBp 2 is critical for plasmodesmata localization and for its interaction with TGBp1 and TGBp3. Frontiers in Microbiology, 13:860695.
[17]	Joanna S, Anna U, Marek F, Jozef J B. 2011. RNA-RNA recombination in plant virus replication and evolution. Annual Review of Phytopathology, 49:415-443.
[18]	Kumar A S, Park E, Nedo A, Alqarni A, Ren L, Hoban K, Caplan J L. 2018. Stromule extension along microtubules coordinated with actin-mediated anchoring guides perinuclear chloroplast movement during innate immunity. Elife, 7:e23625.
[19]	Lamas N S, Matos V O R L, Alves-Freitas D M T, Melo F L, Costa A F, Faria J C, Ribeiro S G. 2017. Occurrence of cowpea mild mottle virus in common bean and associated weeds in Northeastern Brazil. Plant Disease, 101 (10):1828. doi: 10.1094/PDIS-04-17-0562-PDN
[20]	Liu Y H, Priyadarshani S V G N, Chi M R, Yan M K, Mohammadi M A, Zhang M, Zhou Qiao, Wang L L, Luo T T, Wai M H, Wang X M, Cai H Y, Wang H F, Qin Y. 2023. Epigenetic modification mechanisms of chloroplasts mutants in pineapple leaves during somatic regeneration. Horticultural Plant Journal, 9 (3):509-522.
[21]	Liu Yong, Li Fan, Li Yueyue, Zhang Songbai, Gao Xiwu, Xie Yan, Yan Fei, Zhang Ansheng, Dai Liangying, Cheng Zhaobang, Ding Ming, Niu Yanbing, Wang Shengji, Che Haiyan, Jiang Tong, Shi Xiaobin, He Zifu, Wu Yunfeng, Zhang Deyong, Qing Ling, Yan Wanrong, Yang Xuehui, Tang Yafei, Zheng Hongying, Tang Qianjun, Zhang Songbai, Zhang Dongfang, Cai Li, Tao Xiaorong. 2019. Identification,distribution and occurrence of viruses in the main vegetables of China. Scientia Agricultura Sinica, 52 (2):239-261.. (in Chinese) doi: 10.3864/j.issn.0578-1752.2019.02.005
	刘勇, 李凡, 李月月, 张松柏, 高希武, 谢艳, 燕飞, 张安盛, 戴良英, 程兆榜, 丁铭, 牛颜冰, 王升吉, 车海彦, 江彤, 史晓斌, 何自福, 吴云锋, 张德咏, 青玲, 严婉荣, 杨学辉, 汤亚飞, 郑红英, 唐前君, 章松柏, 章东方, 蔡丽, 陶小荣. 2019. 侵染我国主要蔬菜作物的病毒种类、分布与发生趋势. 中国农业科学, 52 (2):239-261. doi: 10.3864/j.issn.0578-1752.2019.02.005
[22]	Mansour A, Al-Musa A, Vetten H J, Lesemann D E. 1998. Properties of a cowpea mild mottle mirus(CPMMV)isolate from eggplant in Jordan and evidence for biological and serological differences between CPMMV isolates from leguminons and solanaceous hosts. Journal of Phytopathology, 146 (11-12):539-547.
[23]	McLeish M J, Fraile A, García-Arenal F. 2019. Evolution of plant-virus interactions:host range and virus emergence. Current Opinion in Virology, 34:50-55.
[24]	Menzel W, Winter S, Vetten H J. 2010. Complete nucleotide sequence of the type isolate of cowpea mild mottle virus from Ghana. Archives of Virology, 155 (12):2069-2073. doi: 10.1007/s00705-010-0821-y pmid: 20890716
[25]	Naidu R A, Gowda S, Satyanarayana T, Boyko V, Reddy A S, Dawson W O, Reddy D V R. 1998. Evidence that whitefly-transmitted cowpea mild mottle virus belongs to the genus carlavirus. Archives of Virology, 143 (4):769-780. pmid: 9638146
[26]	Pardina P E R, Arneodo J D, Truol G A, Herrera P S, Laguna I G. 2004. First report of cowpea mild mottle virus in bean crops in argentina. Australasian Plant Pathology, 33 (1):129-130.
[27]	Samuels T D, Ju H J, Ye C M, Motes C M, Blancaflor E B, Verchot-Lubicz J. 2007. Subcellular targeting and interactions among the potato virus X TGB proteins. Virology, 367:375-389. pmid: 17610926
[28]	Torrance L, Cowan G H, Gillespie T, Ziegler A, Lacomme C. 2006. Barley stripe mosaic virus-encoded proteins triple-gene block 2 and γb localize to chloroplasts in virus-infected monocot and dicot plants,revealing hitherto-unknown roles in virus replication. Journal of General Virology, 87 (8):2403-2411.
[29]	Wan S, Liang B, Yang L, Hu W, Kuang L Q, Song J, Xie J H, Huang Y J, Liu D C, Liu Y. 2024. The MADS-box family gene PtrANR1 encodes a transcription activator promoting root growth and enhancing plant tolerance to drought stress. Plant Cell Reports, 43:16.
[30]	Wang L P, Ding X, Xiao J J, Jiménez-Gόngora T, Liu R Y, LozanoDurán R. 2017. Inference of a Geminivirus-host protein-protein interaction network through affinity purification and mass spectrometry analysis. Viruses, 9 (10):275.
[31]	Wang X W, Shao C X, Liu L B, Wang Y, An Y Q, Li H, Ding Y W, Jing Y P, Li X J, Xiao J W. 2023. PtrDJ1C,an atypical member of the DJ-1 superfamily, is essential for early chloroplast development and lignin deposition in poplar. Horticultural Plant Journal, 9 (5):1039-1054.
[32]	Wei Tian, Li Hongwei, Su Jiankun, Liu Jianfeng, Liu Hongxia. 2016. Screening bacteria for biological control of cucumber green mottle mosaic virus disease. Acta Horticulturae Sinica, 43 (12):2391-2400. (in Chinese) doi: 10.16420/j.issn.0513-353x.2016-0401 URL
	卫甜, 李宏伟, 苏建坤, 刘建凤, 刘红霞. 2016. 黄瓜绿斑驳花叶病毒病生防菌株的分离与筛选. 园艺学报, 43 (12):2391-2400. doi: 10.16420/j.issn.0513-353x.2016-0401 URL
[33]	Wu X Y, Zhang P P, Chen S T, Zhang Z X, Zhang Y H, Fang P P, Ning K, Sun T, Xu P. 2023a. A molecular toolkit to boost functional genomic studies in transformation-recalcitrant vegetable legumes. Horticulture Research, 10 (5):uhad064.
[34]	Wu X Y, Chen S T, Zhang Z X, Zhang Y H, Li P M, Chen X Y, Liu M M, Lu Q, Li Z Y, Wei Z Y, Xu P. 2023b. Development of recombinase polymerase amplification combined with lateral flow strips for rapid detection of cowpea mild mottle virus. Plant Pathology Journal, 39 (5):486-493.
[35]	Wu X Y, Chen S T, Zhang Z X, Zhou W X, Sun T, Ning K, Xu M, Ke X, Xu P. 2024. A viral small interfering RNA-host plant mRNA pathway modulates virus-induced drought tolerance by enhancing autophagy. The Plant Cell, 36 (9):3219-3236.
[36]	Xu M J, Ai H, Yan R, Li P B, Cheng R X, Xu Y. 2024. Natural Occurrence of cowpea mild mottle virus infecting sesame(Sesamum indicum L.)in Anhui Province,China. Plant Disease,https://doi.org/10.1094/PDIS-06-24-1182-PDN.
[37]	Yadav M K, Biswas K K, Lal S K, Baranwal V K, Jain R K. 2013. A distinct strain of cowpea mild mottle virus infecting soybean in India. Journal of Phytopathology, 161 (10):739-744.
[38]	Zanardo L G, Carvalho C M. 2017. Cowpea mild mottle virus(Carlavirus,Betaflexiviridae):a review. Tropical Plant Pathology, 42 (6):417-430.
[39]	Zanardo L G, Silva F N, Lima A T M, Milanesi D F, Castilho-Urquiza G P, Almeida A M R, Zerbini F M, Carvalho C M. 2014. Molecular variability of cowpea mild mottle virus infecting soybean in Brazil. Archives of Virology, 159 (4):727-737. doi: 10.1007/s00705-013-1879-0 pmid: 24142270
[40]	Zhai Y S, Yuan Q, Qiu S Y, Li S S, Li M M, Zheng H Y, Wu G W, Lu Y W, Peng J J, Rao S F, Chen J P, Yan F. 2021. Turnip mosaic virus impairs perinuclear chloroplast clustering to facilitate viral infection. Plant,Cell & Environment, 44 (11):3681-3699.
[41]	Zhao J P, Zhang X, Hong Y G, Liu Y L. 2016. Chloroplast in plant-virus interaction. Frontiers in Microbiology, 7:1565. pmid: 27757106

Identification of Cowpea Mild Mottle Virus Variants in Zhejiang and Localization of TGB Proteins

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