{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,20]],"date-time":"2026-08-20T04:26:33Z","timestamp":1787199993051,"version":"build-2736575974"},"reference-count":35,"publisher":"MDPI AG","issue":"16","license":[{"start":{"date-parts":[[2026,8,18]],"date-time":"2026-08-18T00:00:00Z","timestamp":1787011200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001809","name":"National Natural Science Foundation of China","doi-asserted-by":"crossref","award":["31902094"],"award-info":[{"award-number":["31902094"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"crossref"}]},{"award":["31902094"],"award-info":[{"award-number":["31902094"]}],"id":[{"id":"https:\/\/ror.org\/01h0zpd94","id-type":"ROR","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Plants"],"abstract":"<jats:p>Drought stress is a primary abiotic constraint limiting potato productivity. While polyphenol oxidase (PPO) is known to participate in stress responses, the specific role of StuPPO9 in drought tolerance remains poorly understood. In this study, we generated StuPPO9-overexpressing (OE) and CRISPR\/Cas9-mediated knockout (C4) lines in the potato cultivar \u2018Atlantic\u2019. Under sustained drought stress, OE lines exhibited significantly superior growth phenotypes compared to wild-type (WT) and C4 plants, characterized by increased leaf and root relative water content, root number and enhanced photosynthetic efficiency (Pn and Gs). OE plants also maintained lower levels of MDA and ROS through elevated antioxidant enzyme activities. Notably, transcriptomic analysis revealed that StuPPO9 triggers a global reprogramming of metabolic pathways. Key drought-responsive genes associated with phenylpropanoid biosynthesis (e.g., anthocyanin acyltransferase), terpenoid metabolism, and hormone signaling (e.g., HPt protein) were significantly upregulated in OE plants. These findings suggest that StuPPO9 confers drought resilience through a multi-layered network involving optimized carbon allocation, reinforced cell wall integrity, and enhanced ROS scavenging capacity. This study provides a promising genetic target and theoretical foundation for breeding drought-resistant potato varieties.<\/jats:p>","DOI":"10.3390\/plants15162495","type":"journal-article","created":{"date-parts":[[2026,8,18]],"date-time":"2026-08-18T08:08:41Z","timestamp":1787040521000},"page":"2495","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["StuPPO9 Enhances Drought Tolerance in Potato (Solanum tuberosum) by Coordinating Photosynthetic Stability and Secondary Metabolism Reprogramming"],"prefix":"10.3390","volume":"15","author":[{"given":"Ming","family":"Chi","sequence":"first","affiliation":[{"name":"College of Horticulture and Landscape, Tianjin Agricultural University, No. 22 Jinjing Road, Tianjin 300392, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Bo","family":"Liu","sequence":"additional","affiliation":[{"name":"College of Horticulture and Landscape, Tianjin Agricultural University, No. 22 Jinjing Road, Tianjin 300392, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hengzhao","family":"Yang","sequence":"additional","affiliation":[{"name":"College of Horticulture and Landscape, Tianjin Agricultural University, No. 22 Jinjing Road, Tianjin 300392, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jingting","family":"Chen","sequence":"additional","affiliation":[{"name":"College of Horticulture and Landscape, Tianjin Agricultural University, No. 22 Jinjing Road, Tianjin 300392, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yue","family":"Liu","sequence":"additional","affiliation":[{"name":"College of Horticulture and Landscape, Tianjin Agricultural University, No. 22 Jinjing Road, Tianjin 300392, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,8,18]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"2875","DOI":"10.1111\/pce.15306","article-title":"Proteome Reprogramming and Acquired Stress Tolerance in Potato Cells Exposed to Acute or Stepwise Water Deficit","volume":"48","author":"Cappetta","year":"2025","journal-title":"Plant Cell Environ."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"kiaf126","DOI":"10.1093\/plphys\/kiaf126","article-title":"Integration of multi-omics data and deep phenotyping provides insights into responses to single and combined abiotic stress in potato","volume":"197","author":"Zagorscak","year":"2025","journal-title":"Plant Physiol."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"109","DOI":"10.1007\/s00299-025-03496-9","article-title":"Transcriptomics-proteomics analysis reveals StCOMT1 regulates drought, alkali and combined stresses in potato","volume":"44","author":"Zhang","year":"2025","journal-title":"Plant Cell Rep."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"109782","DOI":"10.1016\/j.plaphy.2025.109782","article-title":"MicroRNA408 negatively regulates drought tolerance by controlling lignin synthesis in potato","volume":"222","author":"Liu","year":"2025","journal-title":"Plant Physiol. Biochem."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"109195","DOI":"10.1016\/j.plaphy.2024.109195","article-title":"Contribution of the regulatory miR156-SPL9 module to the drought stress response in pigmented potato (Solanum tuberosum L.)","volume":"217","author":"Parisi","year":"2024","journal-title":"Plant Physiol. Biochem."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"e14362","DOI":"10.1111\/ppl.14362","article-title":"StMAPK10 gene functional identification and analysis in drought resistance of potato crop (Solanum tuberosum L.)","volume":"176","author":"Majeed","year":"2024","journal-title":"Physiol. Plant."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"e70081","DOI":"10.1111\/tpj.70081","article-title":"Comprehensive transcriptional regulatory networks in potato through chromatin accessibility and transcriptome under drought and salt stresses","volume":"121","author":"Wen","year":"2025","journal-title":"Plant J. Cell Mol. Biol."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"163","DOI":"10.1007\/978-1-4939-3393-8_15","article-title":"Knockdown of Polyphenol Oxidase Gene Expression in Potato (Solanum tuberosum L.) with Artificial MicroRNAs","volume":"1405","author":"Chi","year":"2016","journal-title":"Methods Mol. Biol."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"102310","DOI":"10.1016\/j.pmpp.2024.102310","article-title":"Genome-wide analysis of polyphenol oxidase gene family in Solanum tuberosum and functional identification of StuPPO9 in regulating tolerance to Phytophthora infestans","volume":"132","author":"Chi","year":"2024","journal-title":"Physiol. Mol. Plant Pathol."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"e14258","DOI":"10.1111\/ppl.14258","article-title":"Halotolerant Pseudomonas koreensis S4T10 mitigate salt and drought stress in Arabidopsis thaliana","volume":"176","author":"Kalleku","year":"2024","journal-title":"Physiol. Plant."},{"key":"ref_11","doi-asserted-by":"crossref","unstructured":"Ullah, Z., Iqbal, J., Abbasi, B., Akhtar, W., Kanwal, S., Ali, I., Chalgham, W., El-Sheikh, M., and Mahmood, T. (2023). Assessment of Gus Expression Induced by Anti-Sense OsPPO Gene Promoter and Antioxidant Enzymatic Assays in Response to Drought and Heavy Metal Stress in Transgenic Arabidopsis thaliana. Sustainability, 15.","DOI":"10.3390\/su151712783"},{"key":"ref_12","first-page":"21","article-title":"Response of rice polyphenol oxidase promoter to drought and salt stress","volume":"49","author":"Akhtar","year":"2017","journal-title":"Pak. J. Bot."},{"key":"ref_13","doi-asserted-by":"crossref","unstructured":"Liu, D., Meng, S., Xiang, Z., Yang, G., and He, N. (2019). An R1R2R3 MYB Transcription Factor, MnMYB3R1, Regulates the Polyphenol Oxidase Gene in Mulberry (Morus notabilis). Int. J. Mol. Sci., 20.","DOI":"10.3390\/ijms20102602"},{"key":"ref_14","doi-asserted-by":"crossref","unstructured":"Chi, M., Bhagwat, B., Lane, W.D., Tang, G., Su, Y., Sun, R., Oomah, B.D., Wiersma, P.A., and Xiang, Y. (2014). Reduced polyphenol oxidase gene expression and enzymatic browning in potato (Solanum tuberosum L.) with artificial microRNAs. BMC Plant Biol., 14.","DOI":"10.1186\/1471-2229-14-62"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"3992","DOI":"10.1021\/acs.jafc.0c07810","article-title":"Drought Resistance in Qingke Involves a Reprogramming of the Phenylpropanoid Pathway and UDP-Glucosyltransferase Regulation of Abiotic Stress Tolerance Targeting Flavonoid Biosynthesis","volume":"69","author":"Xu","year":"2021","journal-title":"J. Agric. Food Chem."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"5391","DOI":"10.1021\/acs.jafc.2c07163","article-title":"Multi-Omics Analysis Provides Crucial Insights into the Drought Adaptation of Glycyrrhiza uralensis Fisch","volume":"71","author":"Li","year":"2023","journal-title":"J. Agric. Food Chem."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"144402","DOI":"10.1016\/j.ijbiomac.2025.144402","article-title":"Terpene synthase TgHPTPS2 from Torreya grandis modulates terpenoid profiles to balance ROS and confer drought tolerance in plants","volume":"314","author":"Zhang","year":"2025","journal-title":"Int. J. Biol. Macromol."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"e14277","DOI":"10.1111\/ppl.14277","article-title":"Feedback regulation of the isoprenoid pathway by SsdTPS overexpression has the potential to enhance plant tolerance to drought stress","volume":"176","author":"Yao","year":"2024","journal-title":"Physiol. Plant."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"1393","DOI":"10.1111\/nph.18443","article-title":"Comparative transcriptomics and metabolomics reveal specialized metabolite drought stress responses in switchgrass (Panicum virgatum)","volume":"236","author":"Tiedge","year":"2022","journal-title":"New Phytol."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"e14050","DOI":"10.1111\/ppl.14050","article-title":"Drought stress-responsive abscisic acid and salicylic acid crosstalk with the phenylpropanoid pathway in soybean seeds","volume":"175","author":"La","year":"2023","journal-title":"Physiol. Plant."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"1732","DOI":"10.1111\/tpj.15920","article-title":"The histidine phosphotransfer AHP4 plays a negative role in Arabidopsis plant response to drought","volume":"111","author":"Ha","year":"2022","journal-title":"Plant J. Cell Mol. Biol."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"112719","DOI":"10.1016\/j.foodres.2023.112719","article-title":"Integrating metabolomics and metatranscriptomics to explore the formation pathway of aroma-active volatile phenolics and metabolic profile during industrial radish paocai fermentation","volume":"167","author":"Zhao","year":"2023","journal-title":"Food Res. Int."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"2561","DOI":"10.1111\/nph.18734","article-title":"DNA hypomethylation of the host tree impairs interaction with mutualistic ectomycorrhizal fungus","volume":"238","author":"Vigneaud","year":"2023","journal-title":"New Phytol."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"124917","DOI":"10.1016\/j.envpol.2024.124917","article-title":"Nitric oxide and ascorbic acid confer cadmium (Cd) tolerance by improving plant terpenoid metabolism and epigenetically modifying DNA methylation","volume":"362","author":"Farahani","year":"2024","journal-title":"Environ. Pollut."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"110340","DOI":"10.1016\/j.plaphy.2025.110340","article-title":"Expression profiling, phenomics, and metabolomics revealed biochemical pathways regulating silicon induced tolerance to metalloid (As, Sb, Ge) toxicity in mungbean (Vigna radiata L. Wilczek)","volume":"229","author":"Thakral","year":"2025","journal-title":"Plant Physiol. Biochem."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"108080","DOI":"10.1016\/j.plaphy.2023.108080","article-title":"Metabolome and transcriptome reprogramming underlying tomato drought resistance triggered by a Pseudomonas strain","volume":"203","author":"Papadopoulou","year":"2023","journal-title":"Plant Physiol. Biochem."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"109171","DOI":"10.1016\/j.plaphy.2024.109171","article-title":"Xyloglucan endotransglucosylase-hydrolase 1 is a negative regulator of drought tolerance in barley via modulating lignin biosynthesis and stomatal closure","volume":"216","author":"Fu","year":"2024","journal-title":"Plant Physiol. Biochem."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"108451","DOI":"10.1016\/j.plaphy.2024.108451","article-title":"Molecular mechanism of drought resistance in soybean roots revealed using physiological and multi-omics analyses","volume":"208","author":"Wang","year":"2024","journal-title":"Plant Physiol. Biochem."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"377","DOI":"10.1111\/tpj.15294","article-title":"Molecular insights into plant desiccation tolerance: Transcriptomics, proteomics and targeted metabolite profiling in Craterostigma plantagineum","volume":"107","author":"Xu","year":"2021","journal-title":"Plant J. Cell Mol. Biol."},{"key":"ref_30","first-page":"154808","article-title":"Erchen decoction to reduce oxidative stress in dyslipidemia phlegm-dampness retention syndrome mice: In vivo mechanism revealed by metabolomics (liquid chromatography-mass spectrometry)","volume":"115","author":"Chen","year":"2023","journal-title":"Phytomed. Int. J. Phytother. Phytopharm."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"54","DOI":"10.1111\/tpj.15775","article-title":"Cross-inoculation of rhizobiome from a congeneric ruderal plant imparts drought tolerance in maize (Zea mays) through changes in root morphology and proteome","volume":"111","author":"Zhang","year":"2022","journal-title":"Plant J. Cell Mol. Biol."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"138600","DOI":"10.1016\/j.jhazmat.2025.138600","article-title":"Phytodegradation and phytotoxicity of N-ethyl perfluorooctane sulfonamidoethanol (N-EtFOSE) in pakchoi (Brassica chinensis L.) via root and foliar uptake from physio-biochemical and transcriptomic perspectives","volume":"494","author":"Han","year":"2025","journal-title":"J. Hazard. Mater."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"413","DOI":"10.1071\/BI9620413","article-title":"A Re-Examination of the Relative Turgidity Technique for Estimating Water Deficits in Leaves","volume":"15","author":"Barrs","year":"1962","journal-title":"Aust. J. Biol. Sci."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"107656","DOI":"10.1016\/j.plaphy.2023.107656","article-title":"Physiological, biochemical and structural changes in tomato plants by vermicompost application in different exposure periods under glass house conditions","volume":"197","author":"Tikoria","year":"2023","journal-title":"Plant Physiol. Biochem."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"550","DOI":"10.1186\/s13059-014-0550-8","article-title":"Moderated Estimation of Fold Change and Dispersion for RNA-Seq Data with DESeq2","volume":"15","author":"Love","year":"2014","journal-title":"Genome Biol."}],"container-title":["Plants"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2223-7747\/15\/16\/2495\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,20]],"date-time":"2026-08-20T04:19:51Z","timestamp":1787199591000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2223-7747\/15\/16\/2495"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,18]]},"references-count":35,"journal-issue":{"issue":"16","published-online":{"date-parts":[[2026,8]]}},"alternative-id":["plants15162495"],"URL":"https:\/\/doi.org\/10.3390\/plants15162495","relation":{},"ISSN":["2223-7747"],"issn-type":[{"value":"2223-7747","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,18]]}}}