Bai M Y, Fan M, Oh E, Wang Z Y. 2012. A triple helix-loop-helix/basic helix-loop-helix cascade controls cell elongation downstream of multiple hormonal and environmental signaling pathways in Arabidopsis. The Plant Cell, 24, 4917-4929.

Baumann M J, Eklöf J M, Michel G, Kallas A M, Teeri T T, Czjzek M, Brumer H III. 2007. Structural evidence for the evolution of xyloglucanase activity from xyloglucan endo-transglycosylases: biological implications for cell wall metabolism. The Plant Cell, 19, 1947-1963.

Betancur L, Singh B, Rapp R A, Wendel J F, Marks M D, Roberts A W, Haigler C H. 2010. Phylogenetically distinct cellulose synthase genes support secondary wall thickening in Arabidopsis shoot trichomes and cotton fiber. Journal of Integrative Plant Biology, 52, 205-220.

Behar H, Graham S W, Brumer H. 2018. Comprehensive cross-genome survey and phylogeny of glycoside hydrolase family 16 members reveals the evolutionary origin of EG16 and XTH proteins in plant lineages. The Plant Journal, 95, 1114-1128.

Bi H H, Liu Z L, Liu S S, Qiao W C, Zhang K P, Zhao M H, Wang D W. 2024. Genome-wide analysis of wheat xyloglucan endotransglucosylase/hydrolase (XTH) gene family revealed TaXTH17 involved in abiotic stress responses. BMC Plant Biology, 24, 640.

Chen Y, Qi H Y, Yang L J, Xu L, Wang J X, Guo J Z, Zhang L, Tan Y Y, Pan R H, Shu Q Y, Qian Q, Song S Y. 2023. The OsbHLH002/OsICE1-OSH1 module orchestrates secondary cell wall formation in rice. Cell Reports, 42, 112702.

Cho S K, Kim J E, Park J A, Eom T J, Kim W T. 2006. Constitutive expression of abiotic stress-inducible hot pepper CaXTH3, which encodes a xyloglucan endotransglucosylase /hydrolase homolog, improves drought and salt tolerance in transgenic Arabidopsis plants. Febs Letters, 580, 3136–3144.

Choi J Y, Seo Y S, Kim S J, Kim W T, Shin J S. 2011. Constitutive expression of CaXTH3, a hot pepper xyloglucan endotransglucosylase/hydrolase, enhanced tolerance to salt and drought stresses without phenotypic defects in tomato plants (Solanum lycopersicum cv. Dotaerang). Plant Cell Reports, 30, 867-877.

Coleman H D, Brunner A M, Tsai C J. 2021. Synergies and entanglement in secondary cell wall development and abiotic stress response in trees. Frontiers in Plant Science, 12, 639769.

Dabravolski S A, Isayenkov S V. 2023. The regulation of plant cell wall organisation under salt stress. Frontiers in Plant Science, 14, 1118313.

Fu W C, Fan D L, Zhang Y. 2024. Genome-wide identification and characterization of xyloglucan endotransglucosylase/hydrolase gene family in maize (Zea mays L.) and the function of ZmXTH30 in response to drought stress. Environmental and Experimental Botany, 222, 105744.

Gebrechorkos S H, Sheffield J, Vicente-Serrano S M, Funk C, Miralles D G, Peng J, Dyer E, Talib J, Beck H E, Singer M B, Dadson S J. 2025. Warming accelerates global drought severity. Nature, 642, 628-635.

Geitmann A, Bacic A T. 2023. Focus on cell walls. Plant Physiology, 194, 1-4.

Guo H Y, Wang Y C, Wang L Q, Hu P, Wang Y M, Jia Y Y, Zhang C R, Zhang Y, Zhang Y M, Wang C, Yang C P. 2017. Expression of the MYB transcription factor gene BplMYB46 affects abiotic stress tolerance and secondary cell wall deposition in Betula platyphylla. Plant Biotechnology Journal, 15, 107-121.

Hacke U G, Sperry J S, Pockman W T, Davis S D, McCulloh K A. 2001. Trends in wood density and structure are linked to prevention of xylem implosion by negative pressure. Oecologia, 126, 457-461.

Haghir S, Yamada K, Kato M, Tsuge T, Wada T, Tominaga R, Ohashi Y, Aoyama T. 2025. The Arabidopsis basic-helix-loop-helix transcription factor LRL1 activates cell wall-related genes during root hair development. Plant Cell Physiology, 66, 384-399.

Han Y, Luo F, Liang A, Zhang H, Liu T, Qi H. 2024. Aquaporin CmPIP2;3 links H₂O₂ signal and antioxidation to modulate trehalose-induced cold tolerance in melon seedlings. Plant Physiology, 197, kiae477.

Hao Y Q, Zong X M, Ren P, Qian Y Q, Fu A G. 2021. Basic helix-loop-helix (bHLH) transcription factors regulate a wide range of functions in Arabidopsis. International Journal of Molecular Sciences, 22, 7152.

Hu T X, Wang Y Q, Wang Q Q, Dang N N, Wang L, Liu C C, Zhu J H, Zhan X Q. 2019. The tomato 2-oxoglutarate-dependent dioxygenase gene SlF3HL is critical for chilling stress tolerance. Horticulture Research, 6, 45.

Li S T, Zhang J J, Liu L, Wang Z, Li Y, Guo L, Li Y, Zhang X, Ren S, Zhao B, Zhang N, Guo Y D. 2020. SlTLFP8 reduces water loss to improve water-use efficiency by modulating cell size and stomatal density via endoreduplication. Plant Cell Environment, 43, 2666-2679.

Li X, Wang Z, Sun S F, Dai Z, Zhang J, Wang W, Peng K, Geng W, Xia S, Liu Q, Zhai H, Gao S, Zhao N, Tian F, Zhang H, He S. 2024. IbNIEL-mediated degradation of IbNAC087 regulates jasmonic acid-dependent salt and drought tolerance in sweet potato. Journal of Integrative Plant Biology, 66, 176-195.

Liang Y F, Ma F, Li B Y, Guo C, Hu T X, Zhang M K, Liang Y, Zhu J H, Zhan X Q. 2022. A bHLH transcription factor, SlbHLH96, promotes drought tolerance in tomato. Horticulture Research, 9, uhac198.

Liu D C, Guo W F, Guo X Y, Hu W, Kuang L Q, Huang Y J, Xie J H, Liu Y. 2022. Ectopic overexpression of CsECR from navel orange increases cuticular wax accumulation in tomato and enhances its tolerance to drought stress. Frontiers in Plant Science, 13, 924552.

Liu T, Jiao X C, Yang S C, Zhang Z D, Ye L., Li J M, Qi H Y, Hu X H. 2020. Crosstalk between GABA and ALA to improve antioxidation and cell expansion of tomato seedling under cold stress. Environmental and Experimental Botany, 180, 104228.

Liu X Y, Shang C Y, Duan P Y, Yang J, Wang J, Sui D, Chen G, Li X, Li G, Hu S, Hu X H. 2025. The SlWRKY42-SlMYC2 module synergistically enhances tomato saline–alkali tolerance by activating the jasmonic acid signaling and spermidine biosynthesis pathway. Journal of Integrative Plant Biology, 67, 1254-1273.

Ma J H, Geng Y K, Liu H, Zhang M, Liu S, Hao C, Hou J, Zhang Y, Zhang D, Zhang W, Zhang X, Li T. 2023. TaTIP41 and TaTAP46 positively regulate drought tolerance in wheat by inhibiting PP2A activity. Journal of Integrative Plant Biology, 65, 2056-2070.

Miedes E, Lorences E P. 2009. Xyloglucan endotransglucosylase/hydrolases (XTHs) during tomato fruit growth and ripening. Journal of Plant Physiology, 166, 489-498.

Mittler R. 2017. ROS are good. Trends in Plant Science, 22, 11-19.

Nakagawa A, Sepuru K M, Yip S J, Seo H, Coffin CM, Hashimoto K, Li Z, Segawa Y, Iwasaki R, Kato H, Kurihara D, Aihara Y, Kim S, Kinoshita T, Itami K, Han SK, Murakami K, Torii KU. 2024. Chemical inhibition of stomatal differentiation by perturbation of the master-regulatory bHLH heterodimer via an ACT-Like domain. Nature Communications, 15, 8996.

Pei Y G, Xue Q H, Shu P, Xu W, Du X, Wu M, Liu K, Pirrello J, Bouzayen M, Hong Y, Liu M. 2024. Bifunctional transcription factors SlERF.H5 and H7 activate cell wall and repress gibberellin biosynthesis genes in tomato via a conserved motif. Developmental Cell, 59, 1345-1359.e6.

Quadrana L, Almeida J, Asís R, Duffy T, Dominguez P G, Bermúdez L, Conti G, Corrêa da Silva J V, Peralta I E, Colot V, Asurmendi S, Fernie A R, Rossi M, Carrari F. 2014. Natural occurring epialleles determine vitamin E accumulation in tomato fruits. Nature Communications, 5, 4027.

Ren Y R, Yang Y Y, Zhao Q, Zhang T E, Wang C K, Hao Y J, You C X. 2021. MdCIB1, an apple bHLH transcription factor, plays a positive regulator in response to drought stress. Environmental and Experimental Botany, 188, 104523.

Roig-Oliver M, Fullana-Pericàs M, Bota J, Flexas J. 2021. Adjustments in photosynthesis and leaf water relations are related to changes in cell wall composition in Hordeum vulgare and Triticum aestivum subjected to water deficit stress. Plant Science, 311, 111015.

Schwartz A R, Morbitzer R, Lahaye T, Staskawicz B J. 2017. TALE-induced bHLH transcription factors that activate a pectate lyase contribute to water soaking in bacterial spot of tomato. Proceedings of the National Academy of Sciences USA, 114, E897-E903.

Seo J S, Joo J, Kim M J, Kim Y K, Nahm B H, Song S I, Cheong J J, Lee J S, Kim J K, Choi Y D. 2011. OsbHLH148, a basic helix-loop-helix protein, interacts with OsJAZ proteins in a jasmonate signaling pathway leading to drought tolerance in rice. The Plant Journal, 65, 907-921.

Shao L, Zhang Z, Dong L, Zhang Y, Khan A, Han Y, Xu D, Liang A, Ye X, Hu X H, Liu T, Qi H Y. 2025. SlMYB86-SlXTH23 module positively assists ALA-alleviated growth inhibition in tomato seedlings under cold stress. The Plant Journal, 124, e70577.

Somerville C, Bauer S, Brininstool G, Facette M, Hamann T, Milne J, Osborne E, Paredez A, Persson S, Raab T, Vorwerk S, Youngs H. 2004. Toward a systems approach to understanding plant cell walls. Science, 306, 2206-2211.

Taheri S, Gantait S, Azizi P, Mazumdar P. 2022. Drought tolerance improvement in Solanum lycopersicum: an insight into "OMICS" approaches and genome editing. 3 Biotech, 12, 63.

Tian S Y, Li L J, Wei M, Yang F J. 2019. Genome-wide analysis of basic helix-loop-helix superfamily members related to anthocyanin biosynthesis in eggplant (Solanum melongena L.). PeerJ, 7, e7768.

Tiika R J, Wei J, Cui G X, Ma Y J, Yang H S, Duan H R. 2021. Transcriptome-wide characterization and functional analysis of xyloglucan endo-transglycosylase/hydrolase (XTH) gene family of Salicornia europaea L. under salinity and drought stress. BMC Plant Biology, 21, 491.

Velikova V, Yordanov I, Edreva A. 2000. Oxidative stress and some antioxidant systems in acid rain-treated bean plants: protective role of exogenous polyamines. Plant Science, 151, 59-66.

Wang D D, Lu Q H, Wang X M, Ling H, Huang N. 2023. Elucidating the role of SlXTH5 in tomato fruit softening. Horticultural Plant Journal, 9, 777-788.

Wang H M, Zhou L, Fu Y P, Cheng M Y, Wong F L, Phang T H, Sun Z X, Lam H M. 2012. Expression of an apoplast-localized BURP-domain protein from soybean (GmRD22) enhances tolerance towards abiotic stress. Plant Cell and Environment, 35, 1932-1947.

Wang P T, Liu W C, Han C, Wang S T, Bai M Y, Song C P. 2024c. Reactive oxygen species: multidimensional regulators of plant adaptation to abiotic stress and development. Journal of Integrative Plant Biology, 66, 330-367.

Wang Q, Wang M, Chen J, Qi W, Lai J, Ma Z, Song R. 2022. ENB1 encodes a cellulose synthase 5 that directs synthesis of cell wall ingrowths in maize basal endosperm transfer cells. Plant Cell, 34, 1054-1074.

Wang X, Diao Z, Cao C, Liu Y, Xia N, Zhang Y, Lu L, Kong F, Zhou H, Chen L, Zhang J, Wang B, Huang R, Tang D, Li S. 2024a. The receptor-like cytoplasmic kinase OsBSK1-2 regulates immunity via an HLH/bHLH complex. Journal of Integrative Plant Biology, 66, 2754-2771.

Wang X, Ren Z B, Xie S P, Li Z H, Zhou Y Y, Duan L S. 2024b. Jasmonate mimic modulates cell elongation by regulating antagonistic bHLH transcription factors via brassinosteroid signaling. Plant Physiology, 195, 2712-2726.

Wei J W, Liu M H, Zhao D, Du P M, Yan L, Liu D R, Shi Q H, Yang C X, Qin G C, Gong B. 2025. Melatonin confers saline-alkali tolerance in tomato by alleviating nitrosative damage and S-nitrosylation of H⁺-ATPase 2. The Plant Cell, 37, koaf035.

Xu P P, Fang S, Chen H Y, Cai W M. 2020. The brassinosteroid-responsive xyloglucan endotransglucosylase/hydrolase 19 (XTH19) and XTH23 genes are involved in lateral root development under salt stress in Arabidopsis. The Plant Journal, 104, 59-75.

Xu Z J, Wang J C, Ma Y B, Wang F, Wang J R, Zhang Y, Hu X H. 2023. The bZIP transcription factor SlAREB1 regulates anthocyanin biosynthesis in response to low temperature in tomato. The Plant Journal, 115, 205-219.

Xue L Y, Wei Z H, Zhai H, Xing S H, Wang Y X, He S Z, Gao S P, Zhao N, Zhang H, Liu Q C. 2022. The IbPYL8-IbbHLH66-IbbHLH118 complex mediates the abscisic acid-dependent drought response in sweet potato. New Phytologist, 236, 2151-2171.

Yan J W, Huang Y, He H, Han T, Di P C, Sechet J, Fang L, Liang Y, Scheller H V, Mortimer J C, Ni L, Jiang M Y, Hou X L, Zhang A Y. 2019. Xyloglucan endotransglucosylase-hydrolase 30 negatively affects salt tolerance in Arabidopsis. Journal of Experimental Botany, 70, 5495-5506.

Yang X Y, Lu M Q, Wang Y F, Wang Y R, Liu Z J, Chen S. 2021. Response mechanism of plants to drought stress. Horticulturae, 7, 50.

Yuan L Q, Dang J, Zhang J Y, Wang L Y, Zheng H, Li G B, Li J X, Zhou F, Khan A, Zhang Z D, Hu X H. 2024. A glutathione S-transferase regulates lignin biosynthesis and enhances salt tolerance in tomato. Plant Physiology, 196, 2989-3006.

Yuan S, Gou X, Hu J, Xiao C W, Du J. 2024. Mutation of tomato xyloglucan transglucosylase/hydrolase 5 increases fruit firmness and contributes to prolonged shelf life. Journal of Plant Physiology, 303, 154350.

Zeng R, Shi Y, Guo L, Fu D Y, Li M Z, Zhang X Y, Li Z Y, Zhuang J H, Yang X H, Zuo J R, Gong Z Z, Tian F, Yang S H. 2025. A natural variant of COOL1 gene enhances cold tolerance for high-latitude adaptation in maize. Cell, 188, 1315-1329.e13.

Zhang B C, Gao Y H, Zhang L J, Zhou Y H. 2021. The plant cell wall: biosynthesis, construction, and functions. Journal of Integrative Plant Biology, 63, 251-272.

Zhang H, Zhao Y, Zhu J K. 2020. Thriving under stress: how plants balance growth and the stress response. Developmental Cell, 55, 529-543.

Zhang H M, Zhu J H, Gong Z Z, Zhu J K. 2022. Abiotic stress responses in plants. Nature Reviews Genetics, 23, 104-119.

Zhang J L, Liu L, Dong D H, Xu J Y, Li H X, Deng Q L, Zhang Y, Huang W, Zhang H J, Guo Y D. 2025. The transcription factor SlLBD40 regulates seed germination by inhibiting cell wall remodeling enzymes during endosperm weakening. Plant Physiology, 197, kiaf022.

Zhang L C, Zhu M K, Ren L J, Li A Z, Chen G P, Hu Z L. 2018. The SlFSR gene controls fruit shelf-life in tomato. Journal of Experimental Botany, 69, 2897-2909.

Zhang Y, Zhang H Z, Fu J Y, Du Y Y, Qu J, Song Y, Wang P W. 2021. The GmXTH1 gene improves drought stress resistance of soybean seedlings. Molecular Breeding, 42, 3.

Zhang Z D, Dang J, Yuan L Q, Zhang Y H, Zhou F, Li T L, Hu X H. 2024a. Exogenous 5-Aminolevulinic acid improved low-temperature tolerance tomato seedling by regulating starch content and phenylalanine metabolism. Plant Physiology and Biochemistry, 210, 108083.

Zhang Z D, Yuan L Q, Dang J, Zhang YH, Wen YS, Du Y, Liang Y F, Wang Y, Liu T, Li T L, Hu X H. 2024b. 5-Aminolevulinic acid improves cold resistance through regulation of SlMYB4/SlMYB88-SlGSTU43 module to scavenge reactive oxygen species in tomato. Horticulture Research, 11, uhae026.

Zhao Q, Fan Z H, Qiu L, Che Q Q, Wang T, Li Y Y, Wang Y Z. 2020. MdbHLH130, an apple bHLH transcription factor, confers water stress resistance by regulating stomatal closure and ROS homeostasis in transgenic tobacco. Frontiers in Plant Science, 11, 543696.

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, 2403-2419.

Zhu C L, Lin Z M, Liu Y, Li H, Di X L, Li T K, Wang J F, Gao Z M. 2024. A bamboo bHLH transcription factor PeRHL4 has dual functions in enhancing drought and phosphorus starvation tolerance. Plant Cell and Environment, 47, 3015-3029.

Zhu X F, Shi Y Z, Lei G J, Fry S C, Zhang B C, Zhou Y H, Braam J, Jiang T, Xu X Y, Mao C Z, Pan Y J, Yang J L, Wu P, Zheng S J. 2012. XTH31, encoding an in vitro XEH/XET-active enzyme, regulates aluminum sensitivity by modulating in vivo XET action, cell wall xyloglucan content, and aluminum binding capacity in Arabidopsis. The Plant Cell, 24, 4731-4747.