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However, despite their significant ecological importance, limited research has been performed on shrub drought tolerance strategies at the morphological, physiological, and molecular levels. Therefore, this study focused on the typical desert shrub, Calligonum leucocladum, and analyzed its morphology, physiology, and protein expression under two different habitats: moist low-salt and arid low-salt. The results indicate that drought stress inhibited the growth of C. leucocladum, leading to significant reductions in its plant height, base diameter, and crown width by 14.93%, 49.57%, and 48.49%, respectively. Drought stress triggered a 30% decline in stomatal conductance, whereas homeostasis was observed in net photosynthesis, intercellular CO\u2082, and transpiration. The soluble sugar content significantly increased by 13.43%, while the starch, soluble protein, and proline content significantly decreased by 20.32%, 10.67%, and 55.61%, respectively. In addition, under drought stress, membrane peroxidation products, reactive oxygen species metabolites, and antioxidant enzyme activities significantly increased. Weighted gene co-expression network analysis revealed 40 proteins that were significantly enriched in the photosynthesis and oxidative phosphorylation pathways through KEGG enrichment analysis. In addition, C. leucocladum maintains photosynthetic homeostasis by enhancing PSII repair (PsbE, PsbL, PsbH) and electron transfer chain efficiency (PetD, nad 2, nad 9), thereby compensating for the insufficient carbon dioxide supply caused by stomatal limitation. This study integrated multidimensional data from morphology, physiology, and proteomics to reveal that C. leucocladum drives a coupled network of photosynthesis, antioxidant, and carbon metabolism through chloroplast translation reprogramming. It maintains photosynthetic homeostasis and osmotic balance under a 30% decrease in stomatal conductance, elucidating the cross-scale regulatory strategy of desert shrubs adapting to extreme drought.<\/jats:p>","DOI":"10.3390\/ijms26094403","type":"journal-article","created":{"date-parts":[[2025,5,6]],"date-time":"2025-05-06T05:54:43Z","timestamp":1746510883000},"page":"4403","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":9,"title":["Responses of Calligonum leucocladum to Prolonged Drought Stress Through Antioxidant System Activation, Soluble Sugar Accumulation, and Maintaining Photosynthetic Homeostasis"],"prefix":"10.3390","volume":"26","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-7963-8418","authenticated-orcid":false,"given":"Fang","family":"Yang","sequence":"first","affiliation":[{"name":"School of Ecology and Environment, Xinjiang University, Urumqi 830017, China"},{"name":"Key Laboratory of Oasis Ecology, Ministry of Education, Urumqi 830017, China"},{"name":"Xinjiang Jinghe Observation and Research Station of Temperate Desert Ecosystem, Ministry of Education, Jinghe 833300, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2382-776X","authenticated-orcid":false,"given":"Guanghui","family":"Lv","sequence":"additional","affiliation":[{"name":"School of Ecology and Environment, Xinjiang University, Urumqi 830017, China"},{"name":"Key Laboratory of Oasis Ecology, Ministry of Education, Urumqi 830017, China"},{"name":"Xinjiang Jinghe Observation and Research Station of Temperate Desert Ecosystem, Ministry of Education, Jinghe 833300, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2025,5,6]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"5615","DOI":"10.5194\/acp-21-5615-2021","article-title":"Drought-induced biomass burning as a source of black carbon to the central himalaya since 1781 CE as reconstructed from the dasuopu ice core","volume":"21","author":"Barker","year":"2021","journal-title":"Atmos. 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