{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,30]],"date-time":"2026-07-30T12:15:37Z","timestamp":1785413737111,"version":"3.56.0"},"reference-count":48,"publisher":"MDPI AG","issue":"14","license":[{"start":{"date-parts":[[2026,7,18]],"date-time":"2026-07-18T00:00:00Z","timestamp":1784332800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"\u2018Tianchi Talents\u2019 of XPCC","award":["230000339"],"award-info":[{"award-number":["230000339"]}]},{"name":"Guided Science and Technology Program Project of XPCC","award":["2023ZD108"],"award-info":[{"award-number":["2023ZD108"]}]},{"name":"President\u2019s Fund of Tarim University","award":["TDZKBS202302"],"award-info":[{"award-number":["TDZKBS202302"]}]},{"award":["TDZKBS202302"],"award-info":[{"award-number":["TDZKBS202302"]}],"id":[{"id":"https:\/\/ror.org\/05202v862","id-type":"ROR","asserted-by":"publisher"}]},{"name":"National College Student Innovation and Entrepreneurship Training Program","award":["202610757018"],"award-info":[{"award-number":["202610757018"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["IJMS"],"abstract":"<jats:p>This study elucidated the molecular, morphological and anatomical mechanisms underlying differential cold tolerance between two cotton cultivars, cold-tolerant Xinluzhong 61 (C61) and cold-sensitive Tahe 2 (C2). Seedlings were subjected to 0 \u00b0C cold stress for 12 and 24 h, followed by comparative transcriptomic (RNA-Seq) and comprehensive anatomical analyses of cotyledons, true leaves and stems. Transcriptomic profiling identified 8834 and 14,664 differentially expressed genes (DEGs) in C2 and C61 at 12 h, and 14,399 and 16,791 DEGs at 24 h, respectively. KEGG enrichment revealed prominent involvement of phenylpropanoid biosynthesis, photosynthesis, and cutin\/suberin\/wax biosynthesis at 12 h, shifting to phagosome, cysteine and methionine metabolism at 24 h. Phenotypic and anatomical observations confirmed that C61 developed dense stem glandular trichomes absent in C2, maintained significantly greater leaf thickness, palisade tissue thickness and palisade-to-spongy ratio in true leaves after 24 h stress, and exhibited thicker stem xylem. These findings highlight core adaptive traits and provide valuable genetic targets for improving cold tolerance in cotton.<\/jats:p>","DOI":"10.3390\/ijms27146401","type":"journal-article","created":{"date-parts":[[2026,7,20]],"date-time":"2026-07-20T09:32:54Z","timestamp":1784539974000},"page":"6401","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Adaptive Structural and Transcriptional Responses Contribute to Cold Tolerance Variation in Xinluzhong 61 and Tahe 2 Cotton Cultivars"],"prefix":"10.3390","volume":"27","author":[{"given":"Hemeng","family":"Wang","sequence":"first","affiliation":[{"name":"Key Laboratory of Conservation and Utilization of Biological Resources in the Tarim Basin, Alar 843300, China"},{"name":"College of Life Science and Technology, Tarim University, Alar 843300, China"},{"name":"College of Agriculture, Northeast Agricultural University, Harbin 150030, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yuhao","family":"Hu","sequence":"additional","affiliation":[{"name":"Key Laboratory of Conservation and Utilization of Biological Resources in the Tarim Basin, Alar 843300, China"},{"name":"College of Life Science and Technology, Tarim University, Alar 843300, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Fengjiao","family":"Wang","sequence":"additional","affiliation":[{"name":"Key Laboratory of Conservation and Utilization of Biological Resources in the Tarim Basin, Alar 843300, China"},{"name":"College of Life Science and Technology, Tarim University, Alar 843300, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0007-2874-4588","authenticated-orcid":false,"given":"Mengmeng","family":"Jia","sequence":"additional","affiliation":[{"name":"Third Division of the Xinjiang Production and Construction Corps Agricultural Research Institute, Tumushuke 843900, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ziyi","family":"Yang","sequence":"additional","affiliation":[{"name":"Key Laboratory of Conservation and Utilization of Biological Resources in the Tarim Basin, Alar 843300, China"},{"name":"College of Life Science and Technology, Tarim University, Alar 843300, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhijie","family":"Wang","sequence":"additional","affiliation":[{"name":"Key Laboratory of Conservation and Utilization of Biological Resources in the Tarim Basin, Alar 843300, China"},{"name":"College of Life Science and Technology, Tarim University, Alar 843300, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Fuling","family":"Wang","sequence":"additional","affiliation":[{"name":"Key Laboratory of Conservation and Utilization of Biological Resources in the Tarim Basin, Alar 843300, China"},{"name":"College of Life Science and Technology, Tarim University, Alar 843300, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yanqin","family":"Wang","sequence":"additional","affiliation":[{"name":"Key Laboratory of Conservation and Utilization of Biological Resources in the Tarim Basin, Alar 843300, China"},{"name":"College of Life Science and Technology, Tarim University, Alar 843300, China"},{"name":"College of Agriculture, Northeast Agricultural University, Harbin 150030, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,7,18]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","unstructured":"Jan, M., Liu, Z., Guo, C., and Sun, X. (2022). Molecular Regulation of Cotton Fiber Development: A Review. Int. J. Mol. Sci., 23.","DOI":"10.3390\/ijms23095004"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"109320","DOI":"10.1016\/j.plaphy.2024.109320","article-title":"Gene expression modules during the emergence stage of upland cotton under low-temperature stress and identification of the GhSPX9 cold-tolerance gene","volume":"218","author":"Lin","year":"2025","journal-title":"Plant Physiol. Biochem."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"9","DOI":"10.1186\/s42397-021-00086-4","article-title":"Review of oxidative stress and antioxidative defense mechanisms in Gossypium hirsutum L. in response to extreme abiotic conditions","volume":"4","author":"Qamer","year":"2021","journal-title":"J. Cotton Res."},{"key":"ref_4","doi-asserted-by":"crossref","unstructured":"Li, Y., Zhu, J., Xu, J., Zhang, X., Xie, Z., and Li, Z. (2024). Effect of cold stress on photosynthetic physiological characteristics and molecular mechanism analysis in cold-resistant cotton (ZM36) seedlings. Front. Plant Sci., 15.","DOI":"10.3389\/fpls.2024.1396666"},{"key":"ref_5","doi-asserted-by":"crossref","unstructured":"Tang, S., Xian, Y., Wang, F., Luo, C., Song, W., Xie, S., Chen, X., Cao, A., Li, H., and Liu, H. (2021). Comparative transcriptome analysis of leaves during early stages of chilling stress in two different chilling-tolerant brown-fiber cotton cultivars. PLoS ONE, 16.","DOI":"10.1371\/journal.pone.0246801"},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"e70246","DOI":"10.1111\/ppl.70246","article-title":"Photoprotective Responses of Cotton Non-Leaf Green Tissues to Short-Term Low-Temperature Stress","volume":"177","author":"Yang","year":"2025","journal-title":"Physiol. Plant."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"444","DOI":"10.1016\/j.tplants.2007.07.002","article-title":"Cold stress regulation of gene expression in plants","volume":"12","author":"Chinnusamy","year":"2007","journal-title":"Trends Plant Sci."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"5615","DOI":"10.1093\/jxb\/eraa254","article-title":"The \u03b2-ketoacyl-CoA synthase KCS13 regulates the cold response in cotton by modulating lipid and oxylipin biosynthesis","volume":"71","author":"Wang","year":"2020","journal-title":"J. Exp. Bot."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"154324","DOI":"10.1016\/j.jplph.2024.154324","article-title":"An insight into heat stress response and adaptive mechanism in cotton","volume":"302","author":"Dev","year":"2024","journal-title":"J. Plant Physiol."},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"plz045","DOI":"10.1093\/aobpla\/plz045","article-title":"Comparative transcriptome, physiological and biochemical analyses reveal response mechanism mediated by CBF4 and ICE2 in enhancing cold stress tolerance in Gossypium thurberi","volume":"11","author":"Cai","year":"2019","journal-title":"AoB Plants"},{"key":"ref_11","doi-asserted-by":"crossref","unstructured":"Geng, Z., Liu, J., Zhao, G., Zhou, Y., Feng, T., Geng, X., Liu, X., Liu, X., Zhang, H., and Wang, Y. (2026). Global profiling of alternative splicing landscapes in response to cold stress during the cotyledon stage in cotton. BMC Genom., 27.","DOI":"10.1186\/s12864-026-12631-z"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"3516","DOI":"10.1111\/nph.70855","article-title":"An evolutionarily conserved histone modification H3K37ac activates gene transcription in response to salt stress in rice","volume":"250","author":"Ma","year":"2026","journal-title":"New Phytol."},{"key":"ref_13","doi-asserted-by":"crossref","unstructured":"Zhu, F., Xu, Q., Fan, J., Meng, L., Wang, R., Niu, J., Wang, J., Zhang, G., Shi, S., and Wang, F. (2025). Genome-Wide Identification of BPC Gene Family in Ten Cotton Species and Function Analysis of GhBPC4 Involved in Cold Stress Response. Int. J. Mol. Sci., 26.","DOI":"10.3390\/ijms26167978"},{"key":"ref_14","doi-asserted-by":"crossref","unstructured":"Cheng, G., Zhang, L., Wei, H., Wang, H., Lu, J., and Yu, S. (2020). Transcriptome Analysis Reveals a Gene Expression Pattern Associated with Fuzz Fiber Initiation Induced by High Temperature in Gossypium barbadense. Genes, 11.","DOI":"10.21203\/rs.3.rs-18232\/v1"},{"key":"ref_15","first-page":"1103","article-title":"Gene Regulation and Signal Transduction in the ICE-CBF-COR Signaling Pathway during Cold Stress in Plants","volume":"82","author":"Wang","year":"2017","journal-title":"Biochemistry"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"433","DOI":"10.1046\/j.1365-313x.1998.00310.x","article-title":"Low temperature regulation of the Arabidopsis CBF family of AP2 transcriptional activators as an early step in cold-induced COR gene expression","volume":"16","author":"Gilmour","year":"1998","journal-title":"Plant J."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"112390","DOI":"10.1016\/j.plantsci.2025.112390","article-title":"Molecular mechanisms of cold stress response in cotton: Transcriptional reprogramming and genetic strategies for tolerance","volume":"352","author":"Dev","year":"2025","journal-title":"Plant Sci."},{"key":"ref_18","doi-asserted-by":"crossref","unstructured":"Shen, Q., Zhang, S., Liu, S., Chen, J., Ma, H., Cui, Z., Zhang, X., Ge, C., Liu, R., and Li, Y. (2020). Comparative Transcriptome Analysis Provides Insights into the Seed Germination in Cotton in Response to Chilling Stress. Int. J. Mol. Sci., 21.","DOI":"10.3390\/ijms21062067"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"199","DOI":"10.1007\/s00484-023-02583-4","article-title":"Temperature impacts on cotton yield superposed by effects on plant growth and verticillium wilt infection in China","volume":"68","author":"Zhang","year":"2024","journal-title":"Int. J. Biometeorol."},{"key":"ref_20","doi-asserted-by":"crossref","unstructured":"Zhu, H., Song, J., Dhar, N., Shan, Y., Ma, X.Y., Wang, X.L., Chen, J.Y., Dai, X.F., Li, R., and Wang, Z.S. (2021). Transcriptome Analysis of a Cotton Cultivar Provides Insights into the Differentially Expressed Genes Underlying Heightened Resistance to the Devastating Verticillium Wilt. Cells, 10.","DOI":"10.3390\/cells10112961"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"153920","DOI":"10.1016\/j.jplph.2023.153920","article-title":"Transcriptome and anatomical studies reveal alterations in leaf thickness under long-term drought stress in tobacco","volume":"281","author":"Khan","year":"2023","journal-title":"J. Plant Physiol."},{"key":"ref_22","doi-asserted-by":"crossref","unstructured":"Yao, X.C., Meng, L.F., Zhao, W.L., and Mao, G.L. (2023). Changes in the morphology traits, anatomical structure of the leaves and transcriptome in Lycium barbarum L. under salt stress. Front. Plant Sci., 14.","DOI":"10.3389\/fpls.2023.1090366"},{"key":"ref_23","doi-asserted-by":"crossref","unstructured":"Xu, H., Huang, C., Jiang, X., Zhu, J., Gao, X., and Yu, C. (2022). Impact of Cold Stress on Leaf Structure, Photosynthesis, and Metabolites in Camellia weiningensis and C. oleifera Seedlings. Horticulturae, 8.","DOI":"10.3390\/horticulturae8060494"},{"key":"ref_24","doi-asserted-by":"crossref","unstructured":"Corti, E., Falsini, S., Patrussi, G., Bazihizina, N., Gonnelli, C., and Papini, A. (2025). Rapamycin-Reactivated Lipid Catabolism in Eruca sativa Mill. Exposed to Salt Stress. Cells, 14.","DOI":"10.3390\/cells14141083"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"197","DOI":"10.1038\/s41597-019-0210-7","article-title":"Transcriptomic profiling of cotton Gossypium hirsutum challenged with low-temperature gradients stress","volume":"6","author":"Li","year":"2019","journal-title":"Sci. Data"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"922","DOI":"10.1016\/j.tplants.2022.01.008","article-title":"Transcriptional regulatory network of plant cold-stress responses","volume":"27","author":"Kidokoro","year":"2022","journal-title":"Trends Plant Sci."},{"key":"ref_27","doi-asserted-by":"crossref","unstructured":"Dai, Y., Zhou, J., Zhang, B., Zheng, D., Wang, K., and Han, J. (2025). Time-course transcriptome analysis reveals gene co-expression networks and transposable element responses to cold stress in cotton. BMC Genom., 26.","DOI":"10.1186\/s12864-025-11433-z"},{"key":"ref_28","doi-asserted-by":"crossref","unstructured":"Feng, Y., Li, Z., Kong, X., Khan, A., Ullah, N., and Zhang, X. (2025). Plant Coping with Cold Stress: Molecular and Physiological Adaptive Mechanisms with Future Perspectives. Cells, 14.","DOI":"10.3390\/cells14020110"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"e20005","DOI":"10.7717\/peerj.20005","article-title":"Identification of key pathways and genes underlying melatonin-enhanced drought tolerance in cotton","volume":"13","author":"Zhong","year":"2025","journal-title":"PeerJ"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"31","DOI":"10.1007\/s11120-013-9849-7","article-title":"May photoinhibition be a consequence, rather than a cause, of limited plant productivity?","volume":"117","author":"Adams","year":"2013","journal-title":"Photosynth Res."},{"key":"ref_31","doi-asserted-by":"crossref","unstructured":"Adams, W.W., Stewart, J.J., Cohu, C.M., Muller, O., and Demmig-Adams, B. (2016). Habitat temperature and precipitation of Arabidopsis thaliana ecotypes determine the response of foliar vasculature, photosynthesis, and transpiration to growth temperature. Front. Plant Sci., 7.","DOI":"10.3389\/fpls.2016.01026"},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"45527","DOI":"10.1007\/s11356-022-19066-5","article-title":"Taurine modulates dynamics of oxidative defense, secondary metabolism, and nutrient relation to mitigate boron and chromium toxicity in Triticum aestivum L. plants","volume":"29","author":"Ashraf","year":"2022","journal-title":"Environ. Sci. Pollut. Res. Int."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"46","DOI":"10.1186\/s40659-018-0195-2","article-title":"The physiological and molecular mechanism of brassinosteroid in response to stress: A review","volume":"51","author":"Anwar","year":"2018","journal-title":"Biol. Res."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"117184","DOI":"10.1016\/j.envpol.2021.117184","article-title":"Excretion from long glandular trichomes contributes to alleviation of cadmium toxicity in Nicotiana tabacum","volume":"285","author":"Zhang","year":"2021","journal-title":"Environ. Pollut."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"112285","DOI":"10.1016\/j.phytochem.2020.112285","article-title":"Peltate glandular trichomes of Colquhounia vestita harbor diterpenoid acids that contribute to plant adaptation to UV radiation and cold stresses","volume":"172","author":"Tang","year":"2020","journal-title":"Phytochemistry"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"533","DOI":"10.1093\/aob\/mcz196","article-title":"Leaf trichomes in Metrosideros polymorpha can contribute to avoiding extra water stress by impeding gall formation","volume":"125","author":"Amada","year":"2020","journal-title":"Ann. Bot."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"109","DOI":"10.1007\/s11099-013-0005-6","article-title":"Leaf morphological and ultrastructural performance of eggplant (Solanum melongena L.) in response to water stress","volume":"51","author":"Fu","year":"2013","journal-title":"Photosynthetica"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"1550","DOI":"10.1093\/treephys\/tpw078","article-title":"Limited acclimation in leaf anatomy to experimental drought in Tropical Rainforest Trees","volume":"36","author":"Binks","year":"2016","journal-title":"Tree Physiol."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"1362","DOI":"10.1111\/pce.14287","article-title":"Desiccation of the leaf mesophyll and its implications for CO2 diffusion and light processing","volume":"45","author":"Momayyezi","year":"2022","journal-title":"Plant Cell Environ."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"1517","DOI":"10.15244\/pjoes\/174476","article-title":"Drought-Induced Changes in Leaf Morphology and Anatomy: Overview, Implications and Perspectives","volume":"33","author":"Yavas","year":"2024","journal-title":"Pol. J. Environ. Stud."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"8166","DOI":"10.1002\/ece3.6519","article-title":"Variation in leaf traits at different altitudes reflects the adaptive strategy of plants to environmental changes","volume":"10","author":"Liu","year":"2020","journal-title":"Ecol. Evol."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"231","DOI":"10.1093\/aob\/mcac086","article-title":"Leaf trait covariation and controls on leaf mass per area (LMA) following cotton domestication","volume":"130","author":"Lei","year":"2022","journal-title":"Ann. Bot."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"1283","DOI":"10.1093\/treephys\/tpv091","article-title":"Global poplar root and leaf transcriptomes reveal links between growth and stress responses under nitrogen starvation and excess","volume":"35","author":"Luo","year":"2015","journal-title":"Tree Physiol."},{"key":"ref_44","doi-asserted-by":"crossref","unstructured":"Cavender-Bares, J. (2005). Impacts of Freezing on Long-Distance Transport in Woody Plants. [Master\u2019s Thesis, College of Biological Sciences, University of Minnesota].","DOI":"10.1016\/B978-012088457-5\/50021-6"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"102","DOI":"10.1111\/nph.12973","article-title":"Comparison of phloem and xylem hydraulic architecture in Picea abies stems","volume":"205","author":"Jyske","year":"2015","journal-title":"New Phytol."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"1143","DOI":"10.1111\/nph.12224","article-title":"Concurrent measurements of change in the bark and xylem diameters of trees reveal a phloem-generated turgor signal","volume":"198","author":"Mencuccini","year":"2013","journal-title":"New Phytol."},{"key":"ref_47","first-page":"652","article-title":"Location of the low temperature water flow barrier in stems","volume":"43","author":"Molz","year":"1968","journal-title":"Plant Physiol."},{"key":"ref_48","unstructured":"Adams, W.W., and Terashima, I. (2018). Leaf vasculature and the upper limit of photosynthesis. Photosynthesis: Plastid Biology, Energy Conversion and Carbon Assimilation, Springer."}],"container-title":["International Journal of Molecular Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1422-0067\/27\/14\/6401\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,7,20]],"date-time":"2026-07-20T09:38:25Z","timestamp":1784540305000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1422-0067\/27\/14\/6401"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,7,18]]},"references-count":48,"journal-issue":{"issue":"14","published-online":{"date-parts":[[2026,7]]}},"alternative-id":["ijms27146401"],"URL":"https:\/\/doi.org\/10.3390\/ijms27146401","relation":{},"ISSN":["1422-0067"],"issn-type":[{"value":"1422-0067","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,7,18]]}}}