Integrated Analysis of Physiological, Transcriptomic, and Metabolomic Data Reveals the Drought Response Mechanism in Cabbage
Abstract
1. Introduction
2. Materials and Methods
2.1. Plant Materials and PEG Treatment
2.2. Plant Growth Conditions and Drought Stress Treatment
2.3. Determination of Physiological Parameters
2.4. Transcriptome Analysis Under Drought Stress
2.5. Metabolomic Analysis Under Drought Stress
2.6. Combined Transcriptomic and Metabolome Analysis
2.7. Data Analysis
3. Results
3.1. Screening of Drought-Tolerant and Drought-Sensitive Cabbage Genotypes
3.2. Effects of Drought Stress on Physiological Characteristics of Cabbage Leaves
3.3. Comparative Transcriptome Analysis of “ZG-628” and “ZG-21” in Response to Drought
3.4. Comparative Metabolome Analysis of “ZG-628” and “ZG-21” in Response to Drought
3.5. Integrated Analysis of Differentially Expressed Genes and Differentially Accumulated Metabolites Responding to Drought Stress
4. Discussion
5. Conclusions
Supplementary Materials
Author Contributions
Funding
Data Availability Statement
Conflicts of Interest
References
- Sato, H.; Mizoi, J.; Shinozaki, K.; Yamaguchi-Shinozaki, K. Complex plant responses to drought and heat stress under climate change. Plant J. 2024, 117, 1873–1892. [Google Scholar] [CrossRef]
- Vicente-Serrano, S.M.; Peña-Angulo, D.; Beguería, S.; Domínguez-Castro, F.; Tomás-Burguera, M.; Noguera, I.; Gimeno-Sotelo, L.; El Kenawy, A. Global drought trends and future projections. Philos. T. R. Soc. A 2022, 380, 20210285. [Google Scholar] [CrossRef]
- 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.; et al. Warming accelerates global drought severity. Nature 2025, 642, 628–635. [Google Scholar] [CrossRef] [PubMed]
- Yang, Y.; Ma, X.; Yan, L.; Li, Y.; Wei, S.; Teng, Z.; Zhang, H.; Tang, W.; Peng, S.; Li, Y. Soil–root interface hydraulic conductance determines responses of photosynthesis to drought in rice and wheat. Plant Physiol. 2023, 194, 376–390. [Google Scholar] [CrossRef] [PubMed]
- Chen, Q.; Cao, X.; Nie, X.; Li, Y.; Liang, T.; Ci, L. Alleviation role of functional carbon nanodots for tomato growth and soil environment under drought stress. J. Hazard. Mater. 2022, 423, 127260. [Google Scholar] [CrossRef] [PubMed]
- Raza, A.; Mubarik, M.S.; Sharif, R.; Habib, M.; Jabeen, W.; Zhang, C.; Chen, H.; Chen, Z.; Siddique, K.H.M.; Zhuang, W.; et al. Developing drought-smart, ready-to-grow future crops. Plant Genome 2023, 16, e20279. [Google Scholar] [CrossRef]
- Dzinyela, R.; Hwarari, D.; Opoku, K.N.; Yang, L.; Movahedi, A. Enhancing drought stress tolerance in horticultural plants through melatonin-mediated phytohormonal crosstalk. Plant Cell Rep. 2024, 43, 272. [Google Scholar] [CrossRef]
- Abbas, K.; Li, J.; Gong, B.; Lu, Y.; Wu, X.; Lü, G.; Gao, H. Drought stress tolerance in vegetables: The functional role of structural features, key gene pathways, and exogenous hormones. Int. J. Mol. Sci. 2023, 24, 13876. [Google Scholar] [CrossRef]
- Shawon, R.A.; Kang, B.S.; Lee, S.G.; Kim, S.K.; Ju Lee, H.; Katrich, E.; Gorinstein, S.; Ku, Y.G. Influence of drought stress on bioactive compounds, antioxidant enzymes and glucosinolate contents of Chinese cabbage (Brassica rapa). Food Chem. 2020, 308, 125657. [Google Scholar] [CrossRef]
- Wang, X.; Zhao, W.; Wei, X.; Sun, Y.; Dong, S. Molecular mechanism of drought resistance in soybean roots revealed using physiological and multi-omics analyses. Plant Physiol. Biochem. 2024, 208, 108451. [Google Scholar] [CrossRef]
- Ayyaz, A.; Fang, R.; Ma, J.; Hannan, F.; Huang, Q.; Athar, H.-R.; Sun, Y.; Javed, M.; Ali, S.; Zhou, W.; et al. Calcium nanoparticles (ca-nps) improve drought stress tolerance in brassica napus by modulating the Photosystem II, nutrient acquisition and antioxidant performance. NanoImpact 2022, 28, 100423. [Google Scholar] [CrossRef]
- Liu, H.; Song, S.; Zhang, H.; Li, Y.; Niu, L.; Zhang, J.; Wang, W. Signaling Transduction of ABA, ROS, and Ca2+ in Plant Stomatal Closure in Response to Drought. Int. J. Mol. Sci. 2022, 23, 14824. [Google Scholar] [CrossRef]
- Zhao, Y.; Wang, S.; Ma, X.; He, Y.; Zhou, J.; Jiao, S.; Xun, J.; Kong, X.; Wu, X.; Bai, X. GmANKTM21 positively regulates drought tolerance and enhanced stomatal response through the MAPK signaling pathway in soybean. Int. J. Mol. Sci. 2024, 25, 6972. [Google Scholar] [CrossRef]
- Liu, J.; Shi, X.; Zhang, Z.; Cen, X.; Lin, L.; Wang, X.; Chen, Z.; Zhang, Y.; Zheng, X.; Wu, B.; et al. Deep neural network-mining of rice drought-responsive TF-TAG modules by a combinatorial analysis of ATAC-Seq and RNA-Seq. Plant Cell Environ. 2025, 48, 5217–5235. [Google Scholar] [CrossRef]
- Yu, Y.; He, L.; Wu, Y. Wheat WRKY Transcription factor TaWRKY24 confers drought and salt tolerance in transgenic plants. Plant Physiol. Biochem. 2023, 205, 108137. [Google Scholar] [CrossRef]
- Zhang, J.; Huang, D.; Zhao, X.; Zhang, M.; Wang, Q.; Hou, X.; Di, D.; Su, B.; Wang, S.; Sun, P. Drought-responsive WRKY transcription factor genes IgWRKY50 and IgWRKY32 from iris germanica enhance drought resistance in transgenic arabidopsis. Front. Plant Sci. 2022, 13, 983600. [Google Scholar] [CrossRef]
- Debnath, S.; Kant, A.; Bhowmick, P.; Malakar, A.; Purkaystha, S.; Jena, B.K.; Mudgal, G.; Rahimi, M.; Helal, M.M.U.; Hasan, R.; et al. The enhanced affinity of WRKY reinforces drought tolerance in Solanum Lycopersicum L.: An innovative bioinformatics study. Plants 2023, 12, 762. [Google Scholar] [CrossRef]
- Li, C.; Yan, C.; Sun, Q.; Wang, J.; Yuan, C.; Mou, Y.; Shan, S.; Zhao, X. The bHLH transcription factor AhbHLH112 improves the drought tolerance of peanut. BMC Plant Biol. 2021, 21, 540. [Google Scholar] [CrossRef]
- Gu, X.; Gao, S.; Li, J.; Song, P.; Zhang, Q.; Guo, J.; Wang, X.; Han, X.; Wang, X.; Zhu, Y.; et al. The bHLH transcription factor regulated gene OsWIH2 is a positive regulator of drought tolerance in rice. Plant Physiol. Biochem. 2021, 169, 269–279. [Google Scholar] [CrossRef]
- Liang, Y.; Ma, F.; Li, B.; Guo, C.; Hu, T.; Zhang, M.; Liang, Y.; Zhu, J.; Zhan, X. A bHLH Transcription factor, SlbHLH96, promotes drought tolerance in tomato. Hortic. Res. 2022, 9, uhac198. [Google Scholar] [CrossRef]
- Yang, C.; Huang, Y.; Lv, P.; Antwi-Boasiako, A.; Begum, N.; Zhao, T.; Zhao, J. NAC Transcription factor GmNAC12 improved drought stress tolerance in soybean. Int. J. Mol. Sci. 2022, 23, 12029. [Google Scholar] [CrossRef]
- Sun, S.; Li, X.; Nie, N.; Chen, Y.; Gao, S.; Zhang, H.; He, S.; Liu, Q.; Zhai, H. Sweet potato NAC transcription factor NAC43 negatively regulates plant growth by causing leaf curling and reducing photosynthetic efficiency. Front. Plant Sci. 2023, 14, 1095977. [Google Scholar] [CrossRef]
- Zhang, F.; Wu, J.; Sade, N.; Wu, S.; Egbaria, A.; Fernie, A.R.; Yan, J.; Qin, F.; Chen, W.; Brotman, Y.; et al. Genomic basis underlying the metabolome-mediated drought adaptation of maize. Genome Biol. 2021, 22, 260. [Google Scholar] [CrossRef]
- Yadav, A.K.; Carroll, A.J.; Estavillo, G.M.; Rebetzke, G.J.; Pogson, B.J. Wheat drought tolerance in the field is predicted by amino acid responses to glasshouse-imposed drought. J. Exp. Bot. 2019, 70, 4931–4948. [Google Scholar] [CrossRef]
- Dwivedi, A.K.; Singh, V.; Anwar, K.; Pareek, A.; Jain, M. Integrated transcriptome, proteome and metabolome analyses revealed secondary metabolites and auxiliary carbohydrate metabolism augmenting drought tolerance in rice. Plant Physiol. Biochem. 2023, 201, 107849. [Google Scholar] [CrossRef]
- Hu, H.; Fei, X.; He, B.; Luo, Y.; Qi, Y.; Wei, A. Integrated analysis of metabolome and transcriptome data for uncovering flavonoid components of zanthoxylum bungeanum maxim. Leaves Under Drought Stress. Front. Nutr. 2022, 8, 801244. [Google Scholar] [CrossRef]
- Guo, X.; Lv, L.; Zhao, A.; Zhao, W.; Liu, Y.; Li, Z.; Li, H.; Chen, X. Integrated transcriptome and metabolome analysis revealed differential drought stress response mechanisms of wheat seedlings with varying drought tolerance. BMC Plant Biol. 2025, 25, 571. [Google Scholar] [CrossRef]
- Xu, D.; Ni, Y.; Zhang, X.; Guo, Y. Multiomic analyses of two sorghum cultivars reveals the change of membrane lipids in their responses to water deficit. Plant Physiol. Biochem. 2022, 176, 44–56. [Google Scholar] [CrossRef]
- Mukherjee, A.; Dwivedi, S.; Bhagavatula, L.; Datta, S. Integration of light and aba signaling pathways to combat drought stress in plants. Plant Cell Rep. 2023, 42, 829–841. [Google Scholar] [CrossRef]
- Liu, R.; Liang, G.; Gong, J.; Wang, J.; Zhang, Y.; Hao, Z.; Li, G. a potential aba analog to increase drought tolerance in arabidopsis thaliana. Int. J. Mol. Sci. 2023, 42, 8783. [Google Scholar] [CrossRef]
- Fidler, J.; Graska, J.; Gietler, M.; Nykiel, M.; Prabucka, B.; Rybarczyk-Płońska, A.; Muszyńska, E.; Morkunas, I.; Labudda, M. PYR/PYL/RCAR receptors play a vital role in the abscisic-acid-dependent responses of plants to external or internal stimuli. Cells 2022, 11, 1352. [Google Scholar] [CrossRef]
- Soma, F.; Takahashi, F.; Kidokoro, S.; Kameoka, H.; Suzuki, T.; Uga, Y.; Shinozaki, K.; Yamaguchi-Shinozaki, K. Constitutively active B2 raf-like kinases are required for drought-responsive gene expression upstream of ABA-Activated SnRK2 kinases. Proc. Natl. Acad. Sci. USA 2023, 120, e2221863120. [Google Scholar] [CrossRef]
- Brunetti, C.; Sebastiani, F.; Tattini, M. Review: ABA, flavonols, and the evolvability of land plants. Plant Sci. 2019, 280, 448–454. [Google Scholar] [CrossRef]
- Hussain, Q.; Asim, M.; Zhang, R.; Khan, R.; Farooq, S.; Wu, J. Transcription factors interact with ABA through gene expression and signaling pathways to mitigate drought and salinity stress. Biomolecules 2021, 11, 1159. [Google Scholar] [CrossRef]
- Wang, L.; Tian, T.; Liang, J.; Li, R.; Xin, X.; Qi, Y.; Zhou, Y.; Fan, Q.; Ning, G.; Becana, M.; et al. A transcription factor of the NAC family regulates nitrate-induced legume nodule senescence. New Phytol. 2023, 238, 2113–2129. [Google Scholar] [CrossRef]
- Collin, A.; Daszkowska-Golec, A.; Szarejko, I. Updates on the role of ABSCISIC ACID INSENSITIVE 5 (ABI5) and ABSCISIC ACID-RESPONSIVE ELEMENT BINDING FACTORs (ABFs) in ABA signaling in different developmental stages in plants. Cells 2021, 10, 1996. [Google Scholar] [CrossRef]
- Marusig, D.; Tombesi, S. Abscisic acid mediates drought and salt stress responses in vitis vinifera—A Review. Int. J. Mol. Sci. 2020, 21, 8648. [Google Scholar] [CrossRef]
- Wang, P.; Cao, W.; Yang, L.; Zhang, Y.; Fang, Z.; Zhuang, M.; Lv, H.; Wang, Y.; Cheng, S.; Ji, J. Glucosinolate biosynthetic genes of cabbage: Genome-wide identification, evolution, and expression analysis. Genes 2023, 14, 476. [Google Scholar] [CrossRef]
- Barber, A.; Müller, C. Drought and subsequent soil flooding affect the growth and metabolism of savoy cabbage. Int. J. Mol. Sci. 2021, 22, 13307. [Google Scholar] [CrossRef]
- Zhang, W.; Wang, L.; Zhang, L.; Kong, X.; Zhang, J.; Wang, X.; Pei, Y.; Jin, Z. H2S-mediated balance regulation of stomatal and non-stomatal factors responding to drought stress in Chinese cabbage. Hortic. Res. 2023, 10, uhac284. [Google Scholar] [CrossRef]
- Eom, S.; Baek, S.-A.; Kim, J.; Hyun, T. Transcriptome analysis in chinese cabbage (Brassica rapa ssp. pekinensis) provides the role of glucosinolate metabolism in response to drought stress. Molecules 2018, 23, 1186. [Google Scholar] [CrossRef]
- Hasanuzzaman, M.; Nahar, K.; Anee, T.I.; Khan, M.I.; Fujita, M. Silicon-mediated regulation of antioxidant defense and glyoxalase systems confers drought stress tolerance in Brassica napus L. S. Afr. J. Bot. 2018, 115, 50–57. [Google Scholar]
- Li, L.; Li, H.; Wu, L.; Qi, H. Sulfur dioxide improves drought tolerance through activating Ca2+ signaling pathways in wheat seedlings. Ecotoxicology 2022, 31, 852–859. [Google Scholar] [CrossRef]
- Zhang, X.; Cheng, Z.; Yao, W.; Gao, Y.; Fan, G.; Guo, Q.; Zhou, B.; Jiang, T. Overexpression of PagERF072 from poplar improves salt tolerance. Int. J. Mol. Sci. 2022, 23, 10707. [Google Scholar] [CrossRef]
- Jing, D.; Liu, F.; Li, S.; Dong, Y. Synergistic effects of SAP and PGPR on physiological characteristics of leaves and soil enzyme activities in the rhizosphere of poplar seedlings under drought stress. Front. Plant Sci. 2024, 15, 1485362. [Google Scholar] [CrossRef]
- Mohagheghian, B.; Saeidi, G.; Arzani, A. Phenolic compounds, antioxidant enzymes, and oxidative stress in barley (Hordeum Vulgare L.) genotypes under field drought-stress conditions. BMC Plant Biol. 2025, 25, 709. [Google Scholar] [CrossRef]
- Stefanov, M.; Rashkov, G.; Borisova, P.; Apostolova, E. Sensitivity of the photosynthetic apparatus in maize and sorghum under different drought levels. Plants 2023, 12, 1863. [Google Scholar] [CrossRef]
- Wang, G.; Xu, X.; Gao, Z.; Liu, T.; Li, Y.; Hou, X. Genome-wide identification of LEA gene family and cold response mechanism of BcLEA4-7 and BcLEA4-18 in non-heading chinese cabbage [Brassica campestris (Syn. Brassica rapa) Ssp. Chinensis]. Plant Sci. 2022, 321, 111291. [Google Scholar] [CrossRef]
- Lee, Y.R.; Ko, K.S.; Lee, H.E.; Lee, E.S.; Han, K.; Yoo, J.Y.; Vu, B.N.; Choi, H.N.; Lee, Y.N.; Hong, J.C.; et al. CRISPR/Cas9-mediated HY5 gene editing reduces growth inhibition in Chinese cabbage (Brassica rapa) under ER stress. Int. J. Mol. Sci. 2023, 24, 13105. [Google Scholar] [CrossRef]
- Saadaoui, W.; Tarchoun, N.; Msetra, I.; Pavli, O.; Falleh, H.; Ayed, C.; Amami, R.; Ksouri, R.; Petropoulos, S.A. Effects of drought stress induced by d-mannitol on the germination and early seedling growth traits, physiological parameters and phytochemicals content of tunisian squash (Cucurbita maxima Duch.) landraces. Front. Plant Sci. 2023, 14, 1215394. [Google Scholar] [CrossRef]
- Ren, K.; Tang, T.; Kong, W.; Su, Y.; Wang, Y.; Cheng, H.; Yang, Y.; Zhao, X. Response of watermelon to drought stress and its drought-resistance evaluation. Plants 2025, 14, 1289. [Google Scholar] [CrossRef]
- Chegini, S.N.; Jafarinia, M.; Ghotbi-Ravandi, A.A. Unraveling the impacts of progressive drought stress on the photosynthetic lightreaction of tomato: Assessed by chlorophyll-a fluorescence and gene expression analysis. Cell Mol. Biol. 2024, 70, 176–184. [Google Scholar] [CrossRef]
- Gupta, A.; Rico-Medina, A.; Caño-Delgado, A.I. The physiology of plant responses to drought. Science 2020, 368, 266–269. [Google Scholar] [CrossRef]
- Nolan, T.M.; Vukašinović, N.; Liu, D.; Russinova, E.; Yin, Y. Brassinosteroids: Multidimensional regulators of plant growth, development, and stress responses. Plant Cell 2020, 32, 295–318. [Google Scholar] [CrossRef]
- Hsu, P.; Dubeaux, G.; Takahashi, Y.; Schroeder, J.I. Signaling mechanisms in abscisic acid-mediated stomatal closure. Plant J. 2021, 105, 307–321. [Google Scholar] [CrossRef]
- Wang, Z.; Li, Z.; Zhou, X.; Lu, M.; Ma, Y.; Zhang, M.; Liu, Y.; Gai, Z.; Yang, K.; Ren, M.; et al. Saline-alkaline stress alters the drought resistance of maize through the ABA-PYL-SnRK2s signaling axis. Plant Physiol. Biochem. 2025, 229, 110659. [Google Scholar] [CrossRef]
- González-Guzmán, M.; Rodríguez, L.; Lorenzo-Orts, L.; Pons, C.; Sarrión-Perdigones, A.; Fernández, M.A.; Peirats-Llobet, M.; Forment, J.; Moreno-Alvero, M.; Cutler, S.R.; et al. Tomato PYR/PYL/RCAR abscisic acid receptors show high expression in root, differential sensitivity to the abscisic acid agonist quinabactin, and the capability to enhance plant drought resistance. J. Exp. Bot. 2014, 65, 4451–4464. [Google Scholar] [CrossRef]
- Li, T.; Li, B.; Wang, Y.; Xu, J.; Li, W.; Chen, Z.; Mou, W.; Xue, D. WRKY transcription factors in rice: Key regulators orchestrating development and stress resilience. Plant Cell Environ. 2025, 48, 8388–8406. [Google Scholar] [CrossRef]
- Arroyo-Álvarez, E.; Chan-León, A.; Girón-Ramírez, A.; Fuentes, G.; Estrella-Maldonado, H.; Santamaría, J.M. Genome-wide analysis of WRKY and NAC transcription factors in Carica papaya L. and their possible role in the loss of drought tolerance by recent cultivars through the domestication of their wild ancestors. Plants 2023, 12, 2775. [Google Scholar] [CrossRef]
- Ma, X.; Wang, W.; Zhang, J.; Jiang, Z.; Xu, C.; Zhu, W.; Shi, B.; Yang, W.; Su, H.; Wang, X.; et al. NRT1.1B acts as an abscisic acid receptor in integrating compound environmental cues for plants. Cell 2025, 188, 5231–5248.e20. [Google Scholar] [CrossRef]
- Rao, M.J.; Feng, B.; Ahmad, M.H.; Tahir Ul Qamar, M.; Aslam, M.Z.; Khalid, M.F.; Hussain, S.; Zhong, R.; Ali, Q.; Xu, Q.; et al. LC-MS/MS-based metabolomics approach identified novel antioxidant flavonoids associated with drought tolerance in citrus species. Front. Plant Sci. 2023, 14, 1150854. [Google Scholar] [CrossRef]
- Cao, X.; Hu, Y.; Song, J.; Feng, H.; Wang, J.; Chen, L.; Wang, L.; Diao, X.; Wan, Y.; Liu, S.; et al. Transcriptome sequencing and metabolome analysis reveals the molecular mechanism of drought stress in millet. Int. J. Mol. Sci. 2022, 23, 10792. [Google Scholar] [CrossRef]
- Li, Y.; Chen, Y.; Chen, J.; Shen, C. Flavonoid metabolites in tea plant (Camellia sinensis) stress response: Insights from bibliometric analysis. Plant Physiol. Biochem. 2023, 202, 107934. [Google Scholar] [CrossRef]
- Chen, Y.; Wu, J.; Ma, C.; Zhang, D.; Zhou, D.; Zhang, J.; Yan, M. metabolome and transcriptome analyses reveal changes of rapeseed in response to ABA signal during early seedling development. BMC Plant Biol. 2024, 24, 245. [Google Scholar] [CrossRef]
- Kim, T.J.; Hwang, Y.J.; Park, Y.J.; Lee, J.S.; Kim, J.K.; Lee, M.-H. Metabolomics Reveals Lysinibacillus capsici TT41-Induced Metabolic Shifts Enhancing Drought Stress Tolerance in Kimchi Cabbage (Brassica rapa L. subsp. pekinensis). Metabolites 2024, 14, 87. [Google Scholar] [CrossRef]
- Han, C.; Chen, G.; Zheng, D.; Feng, N. Transcriptomic and metabolomic analyses reveal that ABA increases the salt tolerance of rice significantly correlated with jasmonic acid biosynthesis and flavonoid biosynthesis. Sci. Rep. 2023, 13, 20365. [Google Scholar] [CrossRef]









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Wang, H.; Gao, Y.; Cao, Y.; Li, J. Integrated Analysis of Physiological, Transcriptomic, and Metabolomic Data Reveals the Drought Response Mechanism in Cabbage. Horticulturae 2026, 12, 239. https://doi.org/10.3390/horticulturae12020239
Wang H, Gao Y, Cao Y, Li J. Integrated Analysis of Physiological, Transcriptomic, and Metabolomic Data Reveals the Drought Response Mechanism in Cabbage. Horticulturae. 2026; 12(2):239. https://doi.org/10.3390/horticulturae12020239
Chicago/Turabian StyleWang, Huiru, Yanming Gao, Yune Cao, and Jianshe Li. 2026. "Integrated Analysis of Physiological, Transcriptomic, and Metabolomic Data Reveals the Drought Response Mechanism in Cabbage" Horticulturae 12, no. 2: 239. https://doi.org/10.3390/horticulturae12020239
APA StyleWang, H., Gao, Y., Cao, Y., & Li, J. (2026). Integrated Analysis of Physiological, Transcriptomic, and Metabolomic Data Reveals the Drought Response Mechanism in Cabbage. Horticulturae, 12(2), 239. https://doi.org/10.3390/horticulturae12020239
