Abstract:Mulberry (Morus alba) is an important economically and ecologically significant tree species. Leaf development directly influences mulberry leaves yield and quality, which is also one of the key indexes for identification and evaluation of mulberry varieties. The TCP transcription factor family plays pivotal roles in plant organogenesis and stress responses; however, its systematic characterization in mulberry has been limited to date. In this study, 22 TCP genes (MnTCPs) were identified in the mulberry genome, and their structural characteristics and expression patterns were systematically analyzed. Phylogenetic analysis classified the MnTCPs into 3 subfamilies—PCF, CIN and CYC/TB1. Genes within each subfamily exhibited conserved exon structures and motif compositions, whereas inter-subfamily differences were evident, including a characteristic amino acid deletion in the BASIC domain of CYC/TB1 subfamily members. Analysis of promoter cis-elements revealed that MnTCPs promoters contained abundant hormone-, light- and stress-responsive cis-elements. Expression analysis further revealed significant temporal and varietal divergence: in 'Luoyu 1', most MnTCPs showed peak expression at early developmental stages or exhibited minimal variation, whereas in 'Fengyuan1', the majority reached maximal expression in mature leaves. These findings suggested that MnTCPs might play critical roles in late-stage leaf development and adaptive responses. Overall, this study presents a genome-wide characterization of the TCP gene family in mulberry and provides a theoretical basis for elucidating their regulatory functions and advancing molecular breeding efforts.
李茹霞, 董亚茹, 付娆, 陈传杰, 陈丽琛, 顾寅钰, 李萌, 李东阳, 张海洋. 桑树TCP基因家族全基因组鉴定及其在叶片发育中的表达分析[J]. 农业生物技术学报, 2026, 34(7): 1441-1451.
LI Ru-Xia, DONG Ya-Ru, FU Rao, CHEN Chuan-Jie, CHEN Li-Chen, GU Yin-Yu, LI Meng, LI Dong-Yang, ZHANG Hai-Yang. Genome-wide Identification of the TCP Gene Family in Mulberry (Morus alba) and Its Expression Analysis During Leaf Development. 农业生物技术学报, 2026, 34(7): 1441-1451.
[1] 李仕培, 黎尔纳, 周东来, 等. 2024.桑树资源功能性饲料开发的研究进展[J]. 中国饲料, (03):156-164. (Li S P, Li E N, Zhou D L, et al. 2024. Research progress on the development of functional feed from mulberry resources[J]. China Feed, (3): 156-164.) [2] 曾玉理, 史建国, 常风云, 等. 2024. 桑树响应盐碱胁迫的研究进展[J]. 果树学报, 41(9):1862-1874. (Zeng Y L, Shi J G, Chang F Y, et al.2024. Research progress on mulberry response to saline-alkali stress[J]. Journal of Fruit Science, 41(9): 1862-1874.) [3] 张凤, 于翠, 董朝霞, 等. 2024. 桑树代谢组学研究进展[J].蚕业科学, 50(3): 567-576. (Zhang F, Yu C, Dong Z X, et al.2024. Research progress on metabolomics of mulberry[J]. Science of Sericulture, 50(3): 567-576.) [4] Aguilar-Martínez J A, Sinha N.2013. Analysis of the role of Arabidopsis classⅠ TCP genes AtTCP7, AtTCP8, AtTCP22, and AtTCP23 in leaf development[J]. Frontiers in Plant Science, 4: 406. [5] Challa K R, Rath M, Nath U.2019. The CIN-TCP transcription factors promote commitment to differentiation in Arabidopsis leaf pavement cells via both auxin-dependent and independent pathways[J]. PLoS Genetics, 15(2): e1007988. [6] Cubas P, Lauter N, Doebley J, et al.1999. The TCP domain: A motif found in proteins regulating plant growth and development[J]. Plant Journal, 18(2): 215-222. [7] Doebley J, Stec A, Hubbard L.1997. The evolution of apical dominance in maize[J]. Nature, 386(6624): 485-488. [8] Efroni I, Blum E, Goldshmidt A, et al.2008. A protracted and dynamic maturation schedule underlies Arabidopsis leaf development[J]. Plant Cell, 20(9): 2293-2306. [9] Koyama T, Furutani M, Tasaka M, et al.2007. TCP transcription factors control the morphology of shoot lateral organs via negative regulation of boundary-specific genes in Arabidopsis[J]. Plant Cell, 19(2): 473-484. [10] Koyama T, Mitsuda N, Seki M, et al.2010. TCP transcription factors regulate the activities of ASYMMETRIC LEAVES1 and miR164, as well as the auxin response, during differentiation of leaves in Arabidopsis[J]. Plant Cell, 22(11): 3574-3588. [11] Koyama T, Sato F, Ohme-Takagi M.2017. Roles of miR319 and TCP transcription factors in leaf development[J]. Plant Physiology, 175(2): 874-885. [12] Lan J, Qin G.2020. The regulation of CIN-like TCP transcription factors[J]. International Journal of Molecular Sciences, 21(12): 4498. [13] Liu D H, Luo Y, Han H, et al.2022. Genome-wide analysis of citrus TCP transcription factors and their responses to abiotic stresses[J]. BMC Plant Biology, 22: 325. [14] Navaud O, Dabos P, Carnus E, et al.2007. TCP transcription factors predate the emergence of land plants[J]. Journal of Molecular Evolution, 65(1): 23-33. [15] Palatnik J F, Allen E, Wu X, et al.2003. Control of leaf morphogenesis by microRNAs[J]. Nature, 425(6955): 257-263. [16] Schommer C, Palatnik J F, Aggarwal P, et al.2008. Control of jasmonate biosynthesis and senescence by miR319 targets[J]. PLoS Biology, 6(9): e230. [17] Shankar N, Sunkara P, Nath U.2023. A double-negative feedback loop between miR319c and JAW-TCPs establishes growth pattern in incipient leaf primordia in Arabidopsis thaliana[J]. PLoS Genetics, 19: e1010978. [18] Viola I L, Alem A L, Jure R M, et al.2023. Physiological roles and mechanisms of action of ClassⅠ TCP transcription factors[J]. International Journal of Molecular Sciences, 24: 5437. [19] Wang K, Zhang H, Wei H, et al.2025. Roles of TCP transcription factors in plant growth and development[J]. Physiologia Plantarum, 177: e70357. [20] Yang C, Li D, Mao D, et al.2013. Overexpression of microRNA319 impacts leaf morphogenesis and leads to enhanced cold tolerance in rice (Oryza sativa L.)[J]. Plant, Cell & Environment, 36(10): 2207-2218. [21] Zhang Y, Xu Y P, Nie J K, et al.2023. DNA-TCP complex structures reveal a unique recognition mechanism for TCP transcription factor families[J]. Nucleic Acids Research, 51(1): 434-448. [22] Zhou H, Hwarari D, Ma H, et al.2022. Genomic survey of TCP transcription factors in plants: Phylogenomics, evolution and their biology[J]. Frontiers in Genetics, 13: 1060546