{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,16]],"date-time":"2026-07-16T05:03:49Z","timestamp":1784178229365,"version":"3.55.0"},"reference-count":46,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,7,16]],"date-time":"2026-07-16T00:00:00Z","timestamp":1784160000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Plant Sci."],"abstract":"<jats:sec>\n                    <jats:title>Introduction<\/jats:title>\n                    <jats:p>\n                      Drought stress is a major environmental factor limiting plant growth and ecosystem stability in arid and semi-arid regions.\n                      <jats:italic>Hemerocallis minor<\/jats:italic>\n                      Mill., a perennial herb with strong drought tolerance, has potential applications in ecological restoration and marginal land utilization. However, the coordinated relationships among morphological, physiological, and metabolic responses, as well as the response characteristics under different intensities of drought stress, remain poorly understood.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Methods<\/jats:title>\n                    <jats:p>\n                      In this study, a 12-week gradient drought stress experiment was conducted. An integrated multi-parameter approach was applied to systematically evaluate the responses of\n                      <jats:italic>H. minor<\/jats:italic>\n                      to varying drought intensities at the levels of growth morphology, physiological traits, and root metabolism.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Results<\/jats:title>\n                    <jats:p>Morphologically, drought stress induced changes such as root thickening, whereas severe and extreme drought significantly inhibited plant height, root growth, and adventitious root formation. Physiologically, plants under moderate drought maintained relatively stable leaf water content and some physiological traits, while severe drought caused pronounced membrane damage and oxidative stress responses. Metabolically, drought stress substantially remodeled root metabolic networks, with numerous differential metabolites detected across comparison groups, primarily involving amino acids, flavonoids, and coumarins. Among them, 23 differential metabolites were annotated to the isoflavonoid biosynthesis pathway. Correlation analysis indicated significant associations between root morphological traits, physiological indicators, and key metabolites: proline, malondialdehyde (MDA), and peroxidase (POD) activity were negatively correlated with certain growth parameters, whereas some isoflavone glycosides and isoflavonoid metabolites were positively correlated.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Discussion<\/jats:title>\n                    <jats:p>\n                      This study provides a comprehensive \u201cmorphology-physiology -metabolism\u201d perspective on the responses of\n                      <jats:italic>H. minor<\/jats:italic>\n                      to drought stress. Results indicate that under mild to moderate drought, plants maintain relatively stable growth and physiological status, whereas severe drought induces pronounced stress responses and metabolic adjustments. These response patterns likely contribute to the adaptation of\n                      <jats:italic>H. minor<\/jats:italic>\n                      to drought environments. The findings offer references for the selection and application of drought-tolerant plants in ecological restoration and marginal land utilization, while further validation of the underlying response mechanisms and ecological significance is warranted through anatomical, molecular, targeted metabolomic, and field studies.\n                    <\/jats:p>\n                  <\/jats:sec>","DOI":"10.3389\/fpls.2026.1855949","type":"journal-article","created":{"date-parts":[[2026,7,16]],"date-time":"2026-07-16T04:20:45Z","timestamp":1784175645000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":0,"title":["Morphological, physiological, and root metabolomic responses of Hemerocallis minor Mill. to drought stress"],"prefix":"10.3389","volume":"17","author":[{"given":"Zhen","family":"Liu","sequence":"first","affiliation":[{"name":"School of Life Sciences, Hebei University","place":["Baoding, China"]},{"name":"Key Laboratory of Regional Energy and Environmental Systems Optimization, Ministry of Education, North China Electric Power University","place":["Beijing, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zichan","family":"Li","sequence":"additional","affiliation":[{"name":"School of Life Sciences, Hebei University","place":["Baoding, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Can","family":"Wang","sequence":"additional","affiliation":[{"name":"School of Life Sciences, Hebei University","place":["Baoding, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yikang","family":"Tian","sequence":"additional","affiliation":[{"name":"School of Life Sciences, Hebei University","place":["Baoding, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Qiqi","family":"Han","sequence":"additional","affiliation":[{"name":"School of Life Sciences, Hebei University","place":["Baoding, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiaoming","family":"Wan","sequence":"additional","affiliation":[{"name":"School of Life Sciences, Hebei University","place":["Baoding, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhongkuan","family":"Liu","sequence":"additional","affiliation":[{"name":"Institute of Agro-resources and Environment, Hebei Academy of Agriculture and Forestry Sciences","place":["Shijiazhuang, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhenyu","family":"Liu","sequence":"additional","affiliation":[{"name":"Institute of Agro-resources and Environment, Hebei Academy of Agriculture and Forestry Sciences","place":["Shijiazhuang, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Guixia","family":"Liu","sequence":"additional","affiliation":[{"name":"School of Life Sciences, Hebei University","place":["Baoding, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1965","published-online":{"date-parts":[[2026,7,16]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2025.1565635","article-title":"Exploring physiological and molecular dynamics of drought stress responses in plants: challenges and future directions","volume":"16","author":"Ali","year":"2025","journal-title":"Front. Plant Sci."},{"key":"B2","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1007\/s10343-025-01110-2","article-title":"Physiological, biochemical and molecular response of pepper genotypes exposed to water deficit","volume":"77","author":"Araz","year":"2025","journal-title":"J. Crop Health"},{"key":"B3","volume-title":"Study on Hybrid Breeding of Hemerocallis Minor","author":"Bai","year":"2023"},{"key":"B4","doi-asserted-by":"publisher","first-page":"44","DOI":"10.3969\/j.issn.1000-2006.201803030","article-title":"Effects of drought and re-watering on physiological indexes of four Paeonia lactiflora cultivars and evaluation of their drought resistance","volume":"42","author":"Chang","year":"2018","journal-title":"J. Nanjing. For. Univ. (Nat. Sci. Ed.)"},{"key":"B5","doi-asserted-by":"publisher","first-page":"5169","DOI":"10.5846\/stxb2013\/33\/13","article-title":"Root morphological development and its response to drought stress in peanut cultivars with different drought resistance","volume":"33","author":"Ding","year":"2013","journal-title":"Acta Ecol. Sin."},{"key":"B6","doi-asserted-by":"publisher","first-page":"843","DOI":"10.3969\/j.issn.1674-8530.24.0076","article-title":"Effects of different water and fertilizer equipment on growth of Hemerocallis citrina and soil properties","volume":"43","author":"Fan","year":"2025","journal-title":"J. Drain. Irrig. Mach. Eng."},{"key":"B7","doi-asserted-by":"publisher","DOI":"10.1111\/ppl.70332","article-title":"Advancements in water-saving strategies and crop adaptation to drought: a comprehensive review","volume":"177","author":"Franco-Navarro","year":"2025","journal-title":"Physiol. Plant"},{"key":"B8","doi-asserted-by":"publisher","first-page":"13465","DOI":"10.1038\/s41598-025-94166-z","article-title":"Disparities in the impact of drought on agriculture across countries","volume":"15","author":"Freedman","year":"2025","journal-title":"Sci. Rep."},{"key":"B9","doi-asserted-by":"publisher","first-page":"62","DOI":"10.14067\/j.cnki.1003-8981.2024.04.006","article-title":"Relationship between soluble sugar and soluble protein contents in buds, leaves and fruits of Idesia polycarpa and its alternate bearing","volume":"42","author":"Geng","year":"2024","journal-title":"J. Econ. For."},{"key":"B10","first-page":"930","article-title":"Effects of NaCl stress on growth and related physiological indicators of Hemerocallis minor Mill","volume":"41","author":"Geng","year":"2021","journal-title":"Guihaia"},{"key":"B11","doi-asserted-by":"publisher","DOI":"10.3390\/plants13212962","article-title":"Drought tolerance in plants: physiological and molecular responses","volume":"13","author":"Haghpanah","year":"2024","journal-title":"Plants."},{"key":"B12","doi-asserted-by":"publisher","first-page":"5","DOI":"10.11924\/j.issn.1000-6850.2012-2514","article-title":"Effect of PEG-induced drought stress on physiological and biochemical indicators of resistance in red clover","volume":"29","author":"He","year":"2013","journal-title":"Chin. Agric. Sci. Bull."},{"key":"B13","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2020.597554","article-title":"Morphophysiology of potato (Solanum tuberosum) in response to drought stress: paving the way forward","volume":"11","author":"Hill","year":"2020","journal-title":"Front. Plant Sci."},{"key":"B14","doi-asserted-by":"publisher","first-page":"2062","DOI":"10.9734\/ijecc\/2022\/v12i1131196","article-title":"Morpho-physiological responses of sorghum cultivars to drought stress","volume":"12","author":"Jeevitha","year":"2022","journal-title":"Int. J. Environ. Clim. Change"},{"key":"B15","doi-asserted-by":"publisher","first-page":"68","DOI":"10.11924\/j.issn.1000-6850.casb16070109","article-title":"Responses of desert herbaceous plant growth to drought stress","volume":"33","author":"Jiang","year":"2017","journal-title":"Chin. Agric. Sci. Bull."},{"key":"B16","doi-asserted-by":"publisher","first-page":"115","DOI":"10.1111\/pce.13195","article-title":"UPLC-HRMS-based untargeted metabolic profiling reveals changes in chickpea (Cicer arietinum) metabolome following long\u2010term drought stress","volume":"42","author":"Khan","year":"2019","journal-title":"Plant Cell Environ."},{"key":"B17","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2022.1085409","article-title":"Responses of root system architecture to water stress at multiple scales: A meta-analysis of trials under controlled conditions","volume":"13","author":"Kou","year":"2022","journal-title":"Front. Plant Sci."},{"key":"B18","doi-asserted-by":"publisher","first-page":"1979","DOI":"10.3969\/j.issn.1674-1749.2025.10.001","article-title":"Identification of chemical components in water extract and alcohol extract of Poria cocos by ultra-high performance liquid chromatography coupled with quadrupole time-of-flight mass spectrometry","volume":"18","author":"Lei","year":"2025","journal-title":"Global Tradit. Chin. Med."},{"key":"B19","doi-asserted-by":"publisher","first-page":"680","DOI":"10.1007\/s00344-016-9571-8","article-title":"Drought stress memory in sugar beet: mismatch between biochemical and physiological parameters","volume":"35","author":"Leufen","year":"2016","journal-title":"J. Plant Growth Regul."},{"key":"B20","volume-title":"Principles and Techniques of Plant Physiological and Biochemical Experiments","author":"Li","year":"2000"},{"key":"B21","doi-asserted-by":"publisher","first-page":"214","DOI":"10.3390\/su16104173","article-title":"Effects of different drip irrigation amounts on growth and yield of Hemerocallis citrina","volume":"51","author":"Li","year":"2023","journal-title":"J. Anhui. Agric. Sci."},{"key":"B22","volume-title":"Physiological and Molecular Mechanisms of Lotus Corniculatus Seedlings in Response to Drought","author":"Liu","year":"2024"},{"key":"B23","doi-asserted-by":"publisher","first-page":"14698","DOI":"10.1021\/acs.jafc.0c05987","article-title":"UPLC-QTof-MSE metabolomics reveals changes in leaf primary and secondary metabolism of hop (Humulus lupulus L.) plants under drought stress","volume":"68","author":"Morcol","year":"2020","journal-title":"J. Agric. Food. Chem."},{"key":"B24","doi-asserted-by":"publisher","first-page":"15","DOI":"10.1007\/s10021-024-00946-5","article-title":"Plant functional diversity shapes soil respiration response to soil moisture availability","volume":"28","author":"Podzikowski","year":"2025","journal-title":"Ecosystems."},{"key":"B25","first-page":"1240","article-title":"Transmission pattern of superoxide anion (O2-) signal among tissue cells of winter wheat","volume":"45","author":"Qi","year":"2025","journal-title":"Guihaia"},{"key":"B26","doi-asserted-by":"publisher","first-page":"26","DOI":"10.19349\/j.cnki.issn1006-7949.2026.03.004","article-title":"Inversion model of tobacco chlorophyll content based on hyperspectral feature extraction","volume":"35","author":"Qi","year":"2026","journal-title":"Surv. Mapp. Eng."},{"key":"B27","doi-asserted-by":"publisher","DOI":"10.3390\/plants14071082","article-title":"Transcriptome analysis reveals the role of plant hormone signal transduction pathways in the drought stress response of Hemerocallis middendorffii","volume":"14","author":"Qian","year":"2025","journal-title":"Plants."},{"key":"B28","doi-asserted-by":"publisher","first-page":"304","DOI":"10.16688\/j.zwbh.2025218","article-title":"Effects of soil relative water content and waterlogging stress on the emergence of Spodoptera frugiperda under relatively low temperature conditions","volume":"52","author":"Qiu","year":"2026","journal-title":"Plant Prot."},{"key":"B29","doi-asserted-by":"publisher","DOI":"10.1111\/ppl.13651","article-title":"Shaping the root system architecture in plants for adaptation to drought stress","volume":"174","author":"Ranjan","year":"2022","journal-title":"Physiol. Plant"},{"key":"B30","doi-asserted-by":"publisher","first-page":"2583","DOI":"10.16213\/j.cnki.scjas.2016.11.013","article-title":"Variation patterns of chlorophyll, water saturation deficit, and malondialdehyde in different mulberry cultivars under drought stress","volume":"29","author":"Ren","year":"2016","journal-title":"Southwest. China J. Agric. Sci."},{"key":"B31","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2025.1601094","article-title":"Exogenous strigolactones modulate antioxidant metabolism via CsD27 to enhance drought tolerance in tea plants","volume":"16","author":"Shen","year":"2025","journal-title":"Front. Plant Sci."},{"key":"B32","doi-asserted-by":"publisher","DOI":"10.1016\/j.stress.2025.101153","article-title":"Plant drought stress: physiological, biochemical and molecular mechanisms","volume":"19","author":"Song","year":"2025","journal-title":"Plant Stress"},{"key":"B33","doi-asserted-by":"publisher","first-page":"38","DOI":"10.13817\/j.cnki.cyycp.2023.01.005","article-title":"Effects of drought stress on morphology, physiology and survival rate of Callisia repens","volume":"43","author":"Sun","year":"2023","journal-title":"Grassl. Turf."},{"key":"B34","volume-title":"Study on Shade Tolerance and Drought Tolerance of Three Hemerocallis Species","author":"Wang","year":"2018"},{"key":"B35","doi-asserted-by":"publisher","DOI":"10.3390\/ijms252413733","article-title":"Studies on the physiological response of Hemerocallis middendorffii to two types of drought stresses","volume":"25","author":"Wang","year":"2024","journal-title":"Int. J. Mol. Sci."},{"key":"B36","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2024.1341826","article-title":"Metabolomics reveal root differential metabolites of different root-type alfalfa under drought stress","volume":"15","author":"Wang","year":"2024","journal-title":"Front. Plant Sci."},{"key":"B37","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2024.1371895","article-title":"Current views of drought research: experimental methods, adaptation mechanisms and regulatory strategies","volume":"15","author":"Wang","year":"2024","journal-title":"Front. Plant Sci."},{"key":"B38","doi-asserted-by":"publisher","DOI":"10.3390\/ijms23126869","article-title":"Transcriptomic and metabolomic analysis of seedling-stage soybean responses to PEG-simulated drought stress","volume":"23","author":"Wang","year":"2022","journal-title":"Int. J. Mol. Sci."},{"key":"B39","doi-asserted-by":"publisher","DOI":"10.1016\/j.plaphy.2023.107723","article-title":"Integrative transcriptome and metabolome analysis reveals the mechanism of exogenous melatonin alleviating drought stress in maize roots","volume":"199","author":"Wang","year":"2023","journal-title":"Plant Physiol. Biochem."},{"key":"B40","first-page":"105","article-title":"Changes of relative electrical conductivity in leaves of Bauhinia purpurea and Bauhinia variegata families under low temperature stress","volume":"37","author":"Wang","year":"2021","journal-title":"For. Environ. Sci."},{"key":"B41","doi-asserted-by":"publisher","first-page":"44","DOI":"10.14088\/j.cnki.issn0439-8114.2026.04.008","article-title":"Effects of copper sulfate stress on antioxidant enzyme activities and malondialdehyde content in wheat","volume":"65","author":"Wen","year":"2026","journal-title":"Hubei. Agric. Sci."},{"key":"B42","doi-asserted-by":"publisher","first-page":"4931","DOI":"10.1093\/jxb\/erz224","article-title":"Wheat drought tolerance in the field is predicted by amino acid responses to glasshouse-imposed drought","volume":"70","author":"Yadav","year":"2019","journal-title":"J. Exp. Bot."},{"key":"B43","doi-asserted-by":"publisher","DOI":"10.3390\/plants13172512","article-title":"The small auxin-up RNA 50 (SAUR50) gene from Ammopiptanthus nanus negatively regulates drought tolerance","volume":"13","author":"Zhang","year":"2024","journal-title":"Plants (Basel)."},{"key":"B44","doi-asserted-by":"publisher","DOI":"10.1016\/j.ecolind.2025.114124","article-title":"The impact of climate warming on biomanipulation effectiveness in mitigating eutrophication in shallow small water ecosystems","volume":"178","author":"Zhang","year":"2025","journal-title":"Ecol. Indic."},{"key":"B45","doi-asserted-by":"publisher","first-page":"1276","DOI":"10.13866\/j.azr.2026.06.13","article-title":"Effects of simulated drought on soil moisture and community structure of temperate steppe in the northeastern margin of Qinghai-Xizang Plateau","volume":"43","author":"Zhao","year":"2025","journal-title":"Arid. Zone. Res."},{"key":"B46","doi-asserted-by":"publisher","first-page":"224","DOI":"10.13718\/j.cnki.xdzk.1995.03.013","article-title":"Annual changes of root activity and nutrients in Hemerocallis citrina","volume":"3","author":"Zhou","year":"1995","journal-title":"J. Southwest. Agric. Univ."}],"container-title":["Frontiers in Plant Science"],"original-title":[],"link":[{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fpls.2026.1855949\/full","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,7,16]],"date-time":"2026-07-16T04:20:56Z","timestamp":1784175656000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.frontiersin.org\/articles\/10.3389\/fpls.2026.1855949\/full"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,7,16]]},"references-count":46,"alternative-id":["10.3389\/fpls.2026.1855949"],"URL":"https:\/\/doi.org\/10.3389\/fpls.2026.1855949","relation":{},"ISSN":["1664-462X"],"issn-type":[{"value":"1664-462X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,7,16]]},"article-number":"1855949"}}