{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T17:42:32Z","timestamp":1790185352545,"version":"4.1.0"},"reference-count":105,"publisher":"MDPI AG","issue":"16","license":[{"start":{"date-parts":[[2026,8,15]],"date-time":"2026-08-15T00:00:00Z","timestamp":1786752000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Plants"],"abstract":"<jats:p>Climate change is placing global agriculture under growing pressure, as plants must withstand extreme environmental conditions such as drought and high salinity, both inducing osmotic and oxidative stress. As part of their survival strategies, plants accumulate protective molecules (osmolytes), including the amino acid proline. For decades, plant biology has largely assumed that high proline accumulation under stress signals strong stress tolerance. However, this review challenges that \u201cproline-centric\u201d perspective. Analyses across a wide range of plant species reveal a more complex picture. Stress-induced proline accumulation is not universal: in some species, proline levels remain relatively unchanged, with other metabolites acting as functional osmolytes, or increase only in response to artificially applied severe stress conditions. Even when proline increases, its absolute concentrations may be too low to contribute significantly to osmotic adjustment. Nevertheless, proline may still be involved in stress tolerance mechanisms through its additional roles, detoxifying reactive oxygen species (ROS), directly stabilising proteins or acting as a stress signalling molecule. Comparative analyses of genetically related taxa with varying degrees of stress tolerance sometimes show negative correlations between proline accumulation and tolerance, with higher proline concentrations measured in the most sensitive genotypes. Overall, the evidence indicates that proline\u2018s role in plant survival is highly context-dependent and strongly influenced by genetic background and must therefore be evaluated on a case-by-case basis. Distinguishing whether proline acts as an adaptive defence or merely as a biochemical marker of physiological strain under stress is essential for accurately assessing plant stress tolerance.<\/jats:p>","DOI":"10.3390\/plants15162478","type":"journal-article","created":{"date-parts":[[2026,8,17]],"date-time":"2026-08-17T08:05:41Z","timestamp":1786953941000},"page":"2478","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":2,"title":["Proline: A Reliable Biochemical Marker of Plant Abiotic Stress Tolerance?"],"prefix":"10.3390","volume":"15","author":[{"given":"Delia Maria","family":"Luca","sequence":"first","affiliation":[{"name":"Faculty of Biology, \u201cAlexandru Ioan Cuza\u201d University, 20A Carol I Bd, 700506 Iasi, Romania"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Marius-Nicu\u015for","family":"Grigore","sequence":"additional","affiliation":[{"name":"Doctoral School of Biology, \u201cAlexandru Ioan Cuza\u201d University, 20A Carol I Bd, 700506 Iasi, Romania"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5076-3784","authenticated-orcid":false,"given":"Oscar","family":"Vicente","sequence":"additional","affiliation":[{"name":"Institute for the Conservation and Improvement of Valencian Agrodiversity (COMAV), Universitat Polit\u00e8cnica de Val\u00e8ncia, Camino de Vera s\/n, 46022 Valencia, Spain"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2026,8,15]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"13020","DOI":"10.15835\/nbha51113020","article-title":"Climate Change, Its Impact on Crop Production, Challenges, and Possible Solutions","volume":"51","author":"Alotaibi","year":"2023","journal-title":"Not. Bot. Horti Agrobot."},{"key":"ref_2","doi-asserted-by":"crossref","unstructured":"\u0160ola, I., Poljuha, D., Pavi\u010di\u0107, I., Jurinjak Tu\u0161ek, A., and \u0160amec, D. (2025). Climate Change and Plant Foods: The Influence of Environmental Stressors on Plant Metabolites and Future Food Sources. Foods, 14.","DOI":"10.3390\/foods14030416"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"578","DOI":"10.1111\/ppl.13185","article-title":"Ion Homeostasis for Salinity Tolerance in Plants: A Molecular Approach","volume":"171","author":"Amin","year":"2021","journal-title":"Physiol. Plant."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"909","DOI":"10.1016\/j.plaphy.2010.08.016","article-title":"Reactive Oxygen Species and Antioxidant Machinery in Abiotic Stress Tolerance in Crop Plants","volume":"48","author":"Gill","year":"2010","journal-title":"Plant Physiol. Biochem."},{"key":"ref_5","doi-asserted-by":"crossref","unstructured":"Cannea, F.B., and Padiglia, A. (2025). Antioxidant Defense Systems in Plants: Mechanisms, Regulation, and Biotechnological Strategies for Enhanced Oxidative Stress Tolerance. Life, 15.","DOI":"10.3390\/life15081293"},{"key":"ref_6","doi-asserted-by":"crossref","unstructured":"Hasanuzzaman, M., Bhuyan, M.H.M., Zulfiqar, F., Raza, A., Mohsin, S., Mahmud, J., Fujita, M., and Fotopoulos, V. (2020). Reactive Oxygen Species and Antioxidant Defense in Plants under Abiotic Stress: Revisiting the Crucial Role of a Universal Defense Regulator. Antioxidants, 9.","DOI":"10.3390\/antiox9080681"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"4629","DOI":"10.1007\/s00344-022-10839-3","article-title":"Proline Alleviates Abiotic Stress Induced Oxidative Stress in Plants","volume":"42","author":"Zulfiqar","year":"2023","journal-title":"J. Plant Growth Regul."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"433","DOI":"10.1093\/aob\/mcu239","article-title":"Diversity, Distribution and Roles of Osmoprotective Compounds Accumulated in Halophytes under Abiotic Stress","volume":"115","author":"Slama","year":"2015","journal-title":"Ann. Bot."},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"59","DOI":"10.1007\/s44279-024-00070-x","article-title":"Role of Osmolytes Dynamics in Plant Metabolism to Cope with Salinity Induced Osmotic Stress","volume":"2","author":"Kaur","year":"2024","journal-title":"Discov. Agric."},{"key":"ref_10","doi-asserted-by":"crossref","unstructured":"Arteaga, S., Yabor, L., D\u00edez, M.J., Prohens, J., Boscaiu, M., and Vicente, O. (2020). The Use of Proline in Screening for Tolerance to Drought and Salinity in Common Bean (Phaseolus vulgaris L.) Genotypes. Agronomy, 10.","DOI":"10.3390\/agronomy10060817"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"167","DOI":"10.33545\/2618060X.2024.v7.i12b.5112","article-title":"Biochemical and Physiological Changes in Chickpea (Cicer arietinum) under Combined Drought and Salinity Stress","volume":"7","author":"Sun","year":"2024","journal-title":"Int. J. Res. Agron."},{"key":"ref_12","doi-asserted-by":"crossref","unstructured":"Dbira, S., Al Hassan, M., Gramazio, P., Ferchichi, A., Vicente, O., Prohens, J., and Boscaiu, M. (2018). Variable Levels of Tolerance to Water Stress (Drought) and Associated Biochemical Markers in Tunisian Barley Landraces. Molecules, 23.","DOI":"10.3390\/molecules23030613"},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"222","DOI":"10.1016\/j.plaphy.2023.01.038","article-title":"Individual and Combined Effects of Heat and Drought and Subsequent Recovery on Winter Wheat (Triticum aestivum L.) Photosynthesis, Nitrogen Metabolism, Cell Osmoregulation, and Yield Formation","volume":"196","author":"Ru","year":"2023","journal-title":"Plant Physiol. Biochem."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"e2133","DOI":"10.7717\/peerj.2133","article-title":"Screening for Drought Tolerance in Cultivars of the Ornamental Genus Tagetes (Asteraceae)","volume":"4","author":"Cicevan","year":"2016","journal-title":"PeerJ"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"plac015","DOI":"10.1093\/aobpla\/plac015","article-title":"How Chrysanthemum (Chrysanthemum \u00d7 grandiflorum) \u2018Palisade White\u2019 Deals with Long-Term Salt Stress","volume":"14","author":"Bandurska","year":"2022","journal-title":"AoB Plants"},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"949","DOI":"10.1071\/FP16007","article-title":"Stress Tolerance Mechanisms in Juncus: Responses to Salinity and Drought in Three Juncus Species Adapted to Different Natural Environments","volume":"43","author":"Boscaiu","year":"2016","journal-title":"Funct. Plant Biol."},{"key":"ref_17","doi-asserted-by":"crossref","unstructured":"Mir, R., Romero, I., Gonz\u00e1lez-Orenga, S., Ferrer-Gallego, P.P., Laguna, E., Boscaiu, M., Opric\u0103, L., Grigore, M.-N., and Vicente, O. (2022). Constitutive and Adaptive Traits of Environmental Stress Tolerance in the Threatened Halophyte Limonium angustebracteatum Erben (Plumbaginaceae). Plants, 11.","DOI":"10.3390\/plants11091137"},{"key":"ref_18","doi-asserted-by":"crossref","unstructured":"Behr, J.H., Bouchereau, A., Berardocco, S., Seal, C.E., Flowers, T.J., and Z\u00f6rb, C. (2017). Metabolic and Physiological Adjustment of Suaeda maritima to Combined Salinity and Hypoxia. Ann. Bot., mcw282.","DOI":"10.1093\/aob\/mcw282"},{"key":"ref_19","doi-asserted-by":"crossref","unstructured":"Schiop, S.T., Al Hassan, M., Sestras, A.F., Boscaiu, M., Sestras, R.E., and Vicente, O. (2015). Identification of Salt Stress Biomarkers in Romanian Carpathian Populations of Picea abies (L.) Karst. PLoS ONE, 10.","DOI":"10.1371\/journal.pone.0135419"},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"1479","DOI":"10.1007\/s00468-017-1563-1","article-title":"Biochemical Responses to Drought, at the Seedling Stage, of Several Romanian Carpathian Populations of Norway Spruce (Picea abies L. Karst)","volume":"31","author":"Schiop","year":"2017","journal-title":"Trees"},{"key":"ref_21","doi-asserted-by":"crossref","unstructured":"Kuang, Y., Xu, Y., Zhang, L., Hou, E., and Shen, W. (2017). Dominant Trees in a Subtropical Forest Respond to Drought Mainly via Adjusting Tissue Soluble Sugar and Proline Content. Front. Plant Sci., 8.","DOI":"10.3389\/fpls.2017.00802"},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"89","DOI":"10.1016\/j.tplants.2009.11.009","article-title":"Proline: A Multifunctional Amino Acid","volume":"15","author":"Szabados","year":"2010","journal-title":"Trends Plant Sci."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"39","DOI":"10.1016\/j.tplants.2021.07.009","article-title":"Proline Metabolism as Regulatory Hub","volume":"27","author":"Alvarez","year":"2022","journal-title":"Trends Plant Sci."},{"key":"ref_24","doi-asserted-by":"crossref","unstructured":"Renzetti, M., Funck, D., and Trovato, M. (2024). Proline and ROS: A Unified Mechanism in Plant Development and Stress Response?. Plants, 14.","DOI":"10.3390\/plants14010002"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"998","DOI":"10.1089\/ars.2012.5074","article-title":"Proline Mechanisms of Stress Survival","volume":"19","author":"Liang","year":"2013","journal-title":"Antioxid. Redox Signal."},{"key":"ref_26","doi-asserted-by":"crossref","unstructured":"Hosseinifard, M., Stefaniak, S., Ghorbani Javid, M., Soltani, E., Wojtyla, \u0141., and Garnczarska, M. (2022). Contribution of Exogenous Proline to Abiotic Stresses Tolerance in Plants: A Review. Int. J. Mol. Sci., 23.","DOI":"10.3390\/ijms23095186"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"300","DOI":"10.1111\/pce.12157","article-title":"Is Proline Accumulation per Se Correlated with Stress Tolerance or Is Proline Homeostasis a More Critical Issue?","volume":"37","author":"Sreenivasulu","year":"2014","journal-title":"Plant Cell Environ."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"503","DOI":"10.1007\/s11101-021-09786-w","article-title":"Plant Metabolomics in Biotic and Abiotic Stress: A Critical Overview","volume":"21","author":"Anzano","year":"2022","journal-title":"Phytochem. Rev."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"e02952","DOI":"10.1016\/j.heliyon.2019.e02952","article-title":"Regulation of L-Proline Biosynthesis, Signal Transduction, Transport, Accumulation and Its Vital Role in Plants during Variable Environmental Conditions","volume":"5","author":"Meena","year":"2019","journal-title":"Heliyon"},{"key":"ref_30","doi-asserted-by":"crossref","unstructured":"Roychoudhury, A., and Tripathi, D.K. (2020). Role of Proline and Glycine Betaine in Overcoming Abiotic Stresses. Protective Chemical Agents in the Amelioration of Plant Abiotic Stress, Wiley.","DOI":"10.1002\/9781119552154"},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"227","DOI":"10.1111\/plb.13363","article-title":"Proline, a Multifaceted Signalling Molecule in Plant Responses to Abiotic Stress: Understanding the Physiological Mechanisms","volume":"24","author":"Ghosh","year":"2022","journal-title":"Plant Biol. J."},{"key":"ref_32","doi-asserted-by":"crossref","unstructured":"Zhang, L., and Becker, D.F. (2015). Connecting Proline Metabolism and Signaling Pathways in Plant Senescence. Front. Plant Sci., 6.","DOI":"10.3389\/fpls.2015.00552"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"917","DOI":"10.1093\/jxb\/erad406","article-title":"Pyrroline-5-Carboxylate Metabolism Protein Complex Detected in Arabidopsis thaliana Leaf Mitochondria","volume":"75","author":"Zheng","year":"2024","journal-title":"J. Exp. Bot."},{"key":"ref_34","doi-asserted-by":"crossref","unstructured":"Vahdati, K. (2013). Abiotic Stress in Plants and Metabolic Responses. Abiotic Stress\u2014Plant Responses and Applications in Agriculture, InTechOpen.","DOI":"10.5772\/45842"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"901","DOI":"10.1111\/pce.14529","article-title":"Pyrroline-5-carboxylate Dehydrogenase Is an Essential Enzyme for Proline Dehydrogenase Function during Dark-induced Senescence in Arabidopsis thaliana","volume":"46","author":"Zheng","year":"2023","journal-title":"Plant Cell Environ."},{"key":"ref_36","doi-asserted-by":"crossref","unstructured":"Rizzi, Y.S., Monteoliva, M.I., Fabro, G., Grosso, C.L., Lar\u00f3vere, L.E., and Alvarez, M.E. (2015). P5CDH Affects the Pathways Contributing to Pro Synthesis after ProDH Activation by Biotic and Abiotic Stress Conditions. Front. Plant Sci., 6.","DOI":"10.3389\/fpls.2015.00572"},{"key":"ref_37","doi-asserted-by":"crossref","unstructured":"Ingrisano, R., Tosato, E., Trost, P., Gurrieri, L., and Sparla, F. (2023). Proline, Cysteine and Branched-Chain Amino Acids in Abiotic Stress Response of Land Plants and Microalgae. Plants, 12.","DOI":"10.3390\/plants12193410"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"6279","DOI":"10.1038\/s41598-023-33398-3","article-title":"Genome-Wide Transcriptional Profiling Provides Clues to Molecular Mechanisms Underlying Cold Tolerance in Chickpea","volume":"13","author":"Akbari","year":"2023","journal-title":"Sci. Rep."},{"key":"ref_39","doi-asserted-by":"crossref","unstructured":"Zarattini, M., and Forlani, G. (2017). Toward Unveiling the Mechanisms for Transcriptional Regulation of Proline Biosynthesis in the Plant Cell Response to Biotic and Abiotic Stress Conditions. Front. Plant Sci., 8.","DOI":"10.3389\/fpls.2017.00927"},{"key":"ref_40","doi-asserted-by":"crossref","unstructured":"Radani, Y., Li, R., Korboe, H.M., Ma, H., and Yang, L. (2023). Transcriptional and Post-Translational Regulation of Plant bHLH Transcription Factors during the Response to Environmental Stresses. Plants, 12.","DOI":"10.3390\/plants12112113"},{"key":"ref_41","doi-asserted-by":"crossref","unstructured":"Cao, X., Wu, L., Wu, M., Zhu, C., Jin, Q., and Zhang, J. (2020). Abscisic Acid Mediated Proline Biosynthesis and Antioxidant Ability in Roots of Two Different Rice Genotypes under Hypoxic Stress. BMC Plant Biol., 20.","DOI":"10.1186\/s12870-020-02414-3"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"3060","DOI":"10.1016\/S2095-3119(20)63485-2","article-title":"Melatonin Treatment Induces Chilling Tolerance by Regulating the Contents of Polyamine, \u03b3-Aminobutyric Acid, and Proline in Cucumber Fruit","volume":"20","author":"Madebo","year":"2021","journal-title":"J. Integ. Agric."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"1456","DOI":"10.4161\/psb.21949","article-title":"Role of Proline under Changing Environments: A Review","volume":"7","author":"Hayat","year":"2012","journal-title":"Plant Signal. Behav."},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"205","DOI":"10.1007\/BF00018060","article-title":"Rapid Determination of Free Proline for Water-Stress Studies","volume":"39","author":"Bates","year":"1973","journal-title":"Plant Soil"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"255","DOI":"10.1046\/j.1365-3040.1999.00421.x","article-title":"Subcellular Compartmentation of Proline in the Leaves of the Subantarctic Kerguelen Cabbage Pringlea antiscorbutica R. Br. In vivo 13C-NMR Study","volume":"22","author":"Aubert","year":"1999","journal-title":"Plant Cell Environ."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"227","DOI":"10.1104\/pp.97.1.227","article-title":"Amino Acid and Sucrose Content Determined in the Cytosolic, Chloroplastic, and Vacuolar Compartments and in the Phloem Sap of Spinach Leaves","volume":"97","author":"Riens","year":"1991","journal-title":"Plant Physiol."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"180","DOI":"10.1007\/BF00199748","article-title":"Subcellular Volumes and Metabolite Concentrations in Barley Leaves","volume":"191","author":"Winter","year":"1993","journal-title":"Planta"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"1598","DOI":"10.1104\/pp.107.099820","article-title":"A Reassessment of the Function of the So-Called Compatible Solutes in the Halophytic Plumbaginaceae Limonium latifolium","volume":"144","author":"Gagneul","year":"2007","journal-title":"Plant Physiol."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"19","DOI":"10.1016\/j.envexpbot.2007.10.009","article-title":"Synthesis of Organic Osmolytes and Salt Tolerance Mechanisms in Paspalum vaginatum","volume":"63","author":"Lee","year":"2008","journal-title":"Environ. Exp. Bot."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"13639","DOI":"10.1038\/srep13639","article-title":"Contribution and Distribution of Inorganic Ions and Organic Compounds to the Osmotic Adjustment in Halostachys caspica Response to Salt Stress","volume":"5","author":"Zeng","year":"2015","journal-title":"Sci. Rep."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"945","DOI":"10.1111\/j.1469-8137.2008.02531.x","article-title":"Salinity Tolerance in Halophytes","volume":"179","author":"Flowers","year":"2008","journal-title":"New Phytol."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"407","DOI":"10.1515\/BMC.2011.032","article-title":"Ion Transport and Osmotic Adjustment in Plants and Bacteria","volume":"2","author":"Shabala","year":"2011","journal-title":"Biomol. Concepts"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"89","DOI":"10.1146\/annurev.pp.28.060177.000513","article-title":"The Mechanism of Salt Tolerance in Halophytes","volume":"28","author":"Flowers","year":"1977","journal-title":"Annu. Rev. Plant. Physiol."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"75","DOI":"10.1071\/PP9860075","article-title":"Ion Relations of Plants Under Drought and Salinity","volume":"13","author":"Flowers","year":"1986","journal-title":"Aust. J. Plant Physiol."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"530","DOI":"10.1007\/BF02411558","article-title":"Subcellular Volumes and Metabolite Concentrations in Spinach Leaves","volume":"193","author":"Winter","year":"1994","journal-title":"Planta"},{"key":"ref_56","doi-asserted-by":"crossref","unstructured":"Al Hassan, M., Estrelles, E., Soriano, P., L\u00f3pez-Gresa, M.P., Bell\u00e9s, J.M., Boscaiu, M., and Vicente, O. (2017). Unraveling Salt Tolerance Mechanisms in Halophytes: A Comparative Study on Four Mediterranean Limonium Species with Different Geographic Distribution Patterns. Front. Plant Sci., 8.","DOI":"10.3389\/fpls.2017.01438"},{"key":"ref_57","doi-asserted-by":"crossref","unstructured":"Gonz\u00e1lez-Orenga, S., Al Hassan, M., Llinares, J.V., Lis\u00f3n, P., L\u00f3pez-Gresa, M.P., Verdeguer, M., Vicente, O., and Boscaiu, M. (2019). Qualitative and Quantitative Differences in Osmolytes Accumulation and Antioxidant Activities in Response to Water Deficit in Four Mediterranean Limonium Species. Plants, 8.","DOI":"10.3390\/plants8110506"},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"139","DOI":"10.1016\/j.envexpbot.2005.05.007","article-title":"Clustering of Halophytes from an Inland Salt Marsh in Turkey According to Their Ability to Accumulate Sodium and Nitrogenous Osmolytes","volume":"57","author":"Tipirdamaz","year":"2006","journal-title":"Environ. Exp. Bot."},{"key":"ref_59","doi-asserted-by":"crossref","unstructured":"Gonz\u00e1lez-Orenga, S., Grigore, M.-N., Boscaiu, M., and Vicente, O. (2021). Constitutive and Induced Salt Tolerance Mechanisms and Potential Uses of Limonium Mill. Species. Agronomy, 11.","DOI":"10.3390\/agronomy11030413"},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"231","DOI":"10.1016\/j.envexpbot.2005.02.007","article-title":"Effects of Water Deficit on Growth and Proline Metabolism in Sesuvium portulacastrum","volume":"56","author":"Slama","year":"2006","journal-title":"Environ. Exp. Bot."},{"key":"ref_61","doi-asserted-by":"crossref","unstructured":"Hnilickova, H., Kraus, K., Vachova, P., and Hnilicka, F. (2021). Salinity Stress Affects Photosynthesis, Malondialdehyde Formation, and Proline Content in Portulaca oleracea L.. Plants, 10.","DOI":"10.3390\/plants10050845"},{"key":"ref_62","doi-asserted-by":"crossref","unstructured":"Kazerooni, E.A., Maharachchikumbura, S.S.N., Adhikari, A., Al-Sadi, A.M., Kang, S.-M., Kim, L.-R., and Lee, I.-J. (2021). Rhizospheric Bacillus amyloliquefaciens Protects Capsicum annuum cv. Geumsugangsan from Multiple Abiotic Stresses via Multifarious Plant Growth-Promoting Attributes. Front. Plant Sci., 12.","DOI":"10.3389\/fpls.2021.669693"},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"174","DOI":"10.1016\/j.scienta.2011.01.037","article-title":"Involvement of Proline in Response of Chickpea (Cicer arietinum L.) to Chilling Stress at Reproductive Stage","volume":"128","author":"Kaur","year":"2011","journal-title":"Sci. Hortic."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"835","DOI":"10.1080\/11263504.2019.1701122","article-title":"Seed Germination, Antioxidant Enzymes Activity and Proline Content in Medicinal Plant Tagetes minuta under Salinity Stress","volume":"154","author":"Moghaddam","year":"2020","journal-title":"Plant Biosyst."},{"key":"ref_65","doi-asserted-by":"crossref","unstructured":"Liu, G., Zhang, Y., Yun, Z., Hu, M., Liu, J., Jiang, Y., and Zhang, Z. (2020). Melatonin Enhances Cold Tolerance by Regulating Energy and Proline Metabolism in Litchi Fruit. Foods, 9.","DOI":"10.3390\/foods9040454"},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"70","DOI":"10.1007\/s00425-005-1518-4","article-title":"Responsive Modes of Medicago sativa Proline Dehydrogenase Genes during Salt Stress and Recovery Dictate Free Proline Accumulation","volume":"222","author":"Miller","year":"2005","journal-title":"Planta"},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"121","DOI":"10.1007\/s11103-018-0767-0","article-title":"Synthesis versus Degradation: Directions of Amino Acid Metabolism during Arabidopsis Abiotic Stress Response","volume":"98","author":"Hildebrandt","year":"2018","journal-title":"Plant Mol. Biol."},{"key":"ref_68","doi-asserted-by":"crossref","unstructured":"Kov\u00e1cs, H., Aleksza, D., Baba, A.I., Hajdu, A., Kir\u00e1ly, A.M., Zsigmond, L., T\u00f3th, S.Z., Kozma-Bogn\u00e1r, L., and Szabados, L. (2019). Light Control of Salt-Induced Proline Accumulation Is Mediated by ELONGATED HYPOCOTYL 5 in Arabidopsis. Front. Plant Sci., 10.","DOI":"10.3389\/fpls.2019.01584"},{"key":"ref_69","doi-asserted-by":"crossref","unstructured":"Yang, D., Ni, R., Yang, S., Pu, Y., Qian, M., Yang, Y., and Yang, Y. (2021). Functional Characterization of the Stipa purpurea P5CS Gene under Drought Stress Conditions. Int. J. Mol. Sci., 22.","DOI":"10.3390\/ijms22179599"},{"key":"ref_70","doi-asserted-by":"crossref","unstructured":"Luan, Y., An, H., Chen, Z., Zhao, D., and Tao, J. (2024). Functional Characterization of the Paeonia ostii P5CS Gene under Drought Stress. Plants, 13.","DOI":"10.3390\/plants13152145"},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"109444","DOI":"10.1016\/j.scienta.2020.109444","article-title":"SlWRKY81 Reduces Drought Tolerance by Attenuating Proline Biosynthesis in Tomato","volume":"270","author":"Ahammed","year":"2020","journal-title":"Sci. Hortic."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"1227","DOI":"10.1016\/S0031-9422(00)90389-3","article-title":"Effects of Water-Stress on Proline Content and Ornithine Aminotransferase Activity of Radish Cotyledons","volume":"37","author":"Hervieu","year":"1994","journal-title":"Phytochemistry"},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"433","DOI":"10.1007\/s13580-020-00231-z","article-title":"Evaluation of Chlorophyll Fluorescence Parameters and Proline Content in Tomato Seedlings Grown under Different Salt Stress Conditions","volume":"61","author":"Shin","year":"2020","journal-title":"Hortic. Environ. Biotechnol."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"4245","DOI":"10.1093\/jxb\/erm284","article-title":"Compatible Solute Accumulation and Stress-Mitigating Effects in Barley Genotypes Contrasting in Their Salt Tolerance","volume":"58","author":"Chen","year":"2007","journal-title":"J. Exp. Bot."},{"key":"ref_75","first-page":"619","article-title":"Detection of Temperature Stress Using Chlorophyll Fluorescence Parameters and Stress-Related Chlorophyll and Proline Content in Paprika (Capsicum annuum L.) Seedlings","volume":"36","author":"Bhandari","year":"2018","journal-title":"Hortic. Sci. Technol."},{"key":"ref_76","doi-asserted-by":"crossref","unstructured":"Mircea, D.M., Estrelles, E., Al Hassan, M., Soriano, P., Sestras, R.E., Boscaiu, M., Sestras, A.F., and Vicente, O. (2023). Effect of Water Deficit on Germination, Growth and Biochemical Responses of Four Potentially Invasive Ornamental Grass Species. Plants, 12.","DOI":"10.3390\/plants12061260"},{"key":"ref_77","doi-asserted-by":"crossref","unstructured":"Todea (Morar), I.M., Gonz\u00e1lez-Orenga, S., Boscaiu, M., Plazas, M., Sestras, A.F., Prohens, J., Vicente, O., and Sestras, R.E. (2020). Responses to Water Deficit and Salt Stress in Silver Fir (Abies alba Mill.) Seedlings. Forests, 11.","DOI":"10.3390\/f11040395"},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1111\/j.1469-8137.1985.tb02816.x","article-title":"Tansley Review No. 2: Regulation of pH and Generation of Osmolarity in Vascular Plants: A Cost-Benefit Analysis in Relation to Efficiency of Use of Energy, Nitrogen and Water","volume":"101","author":"Raven","year":"1985","journal-title":"New Phytol."},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"plu049","DOI":"10.1093\/aobpla\/plu049","article-title":"Responses of Five Mediterranean Halophytes to Seasonal Changes in Environmental Conditions","volume":"6","author":"Gil","year":"2014","journal-title":"AoB Plants"},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"9","DOI":"10.15835\/nbha3927176","article-title":"Soluble Carbohydrates as Osmolytes in Several Halophytes from a Mediterranean Salt Marsh","volume":"39","author":"Gil","year":"2011","journal-title":"Not. Bot. Horti Agrobot. Cluj."},{"key":"ref_81","doi-asserted-by":"crossref","unstructured":"Homayouni, H., Razi, H., Izadi, M., Alemzadeh, A., Kazemeini, S.A., Niazi, A., and Vicente, O. (2024). Temporal Changes in Biochemical Responses to Salt Stress in Three Salicornia Species. Plants, 13.","DOI":"10.3390\/plants13070979"},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"772","DOI":"10.3832\/ifor1233-007","article-title":"Effect of Salt and Drought on Growth, Physiological and Biochemical Responses of Two Tamarix Species","volume":"8","author":"Dawalibi","year":"2015","journal-title":"iForest"},{"key":"ref_83","doi-asserted-by":"crossref","unstructured":"Patel, J., Khandwal, D., Choudhary, B., Ardeshana, D., Jha, R.K., Tanna, B., Yadav, S., Mishra, A., Varshney, R.K., and Siddique, K.H.M. (2022). Differential Physio-Biochemical and Metabolic Responses of Peanut (Arachis hypogaea L.) under Multiple Abiotic Stress Conditions. Int. J. Mol. Sci., 23.","DOI":"10.3390\/ijms23020660"},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"10571","DOI":"10.1007\/s42729-025-02810-4","article-title":"Foliar Application of Seaweed-Based Biostimulants Enhances Drought Resilience, Physiological Performance, and Seed Oil Quality in Sesame (Sesamum indicum L.)","volume":"25","author":"Alayafi","year":"2025","journal-title":"J. Soil Sci. Plant Nutr."},{"key":"ref_85","first-page":"1413","article-title":"Morphophysiological Response of Rice Genotypes to Seed Priming in Various Environments","volume":"11","author":"Akmal","year":"2025","journal-title":"J. Penelit. Pendidik. IPA"},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"14328","DOI":"10.1038\/s41598-021-93697-5","article-title":"Heat-Tolerant Hot Pepper Exhibits Constant Photosynthesis via Increased Transpiration Rate, High Proline Content and Fast Recovery in Heat Stress Condition","volume":"11","author":"Rajametov","year":"2021","journal-title":"Sci. Rep."},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"193","DOI":"10.1016\/j.sajb.2018.05.022","article-title":"Comparative Analysis of Water Deficit and Salt Tolerance Mechanisms in Silene","volume":"117","author":"Kozminska","year":"2018","journal-title":"S. Afr. J. Bot."},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"1061","DOI":"10.1093\/treephys\/tpt074","article-title":"Physiological, Biochemical and Molecular Responses in Four Prunus Rootstocks Submitted to Drought Stress","volume":"33","author":"Jimenez","year":"2013","journal-title":"Tree Physiol."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"195","DOI":"10.1034\/j.1399-3054.1998.1040207.x","article-title":"Stress-induced Changes in Polyamine and Tyramine Levels Can Regulate Proline Accumulation in Tomato Leaf Discs Treated with Sodium Chloride","volume":"104","author":"Aziz","year":"1998","journal-title":"Physiol. Plant."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"1050","DOI":"10.1111\/plb.12178","article-title":"Genotype Differences in the Metabolism of Proline and Polyamines under Moderate Drought in Tomato Plants","volume":"16","author":"Romero","year":"2014","journal-title":"Plant Biol. J."},{"key":"ref_91","doi-asserted-by":"crossref","unstructured":"El-Beltagi, H.S., Mohamed, H.I., and Sofy, M.R. (2020). Role of Ascorbic Acid, Glutathione and Proline Applied as Singly or in Sequence Combination in Improving Chickpea Plant through Physiological Change and Antioxidant Defense under Different Levels of Irrigation Intervals. Molecules, 25.","DOI":"10.3390\/molecules25071702"},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"787","DOI":"10.1111\/j.1399-3054.1997.tb01064.x","article-title":"Water and Salt Stress-induced Alterations in Proline Metabolism of Triticum durum Seedlings","volume":"101","author":"Mattioni","year":"1997","journal-title":"Physiol. Plant."},{"key":"ref_93","doi-asserted-by":"crossref","unstructured":"Al Hassan, M., Pacurar, A., L\u00f3pez-Gresa, M.P., Donat-Torres, M.P., Llinares, J.V., Boscaiu, M., and Vicente, O. (2016). Effects of Salt Stress on Three Ecologically Distinct Plantago Species. PLoS ONE, 11.","DOI":"10.1371\/journal.pone.0160236"},{"key":"ref_94","doi-asserted-by":"crossref","unstructured":"Jaarsma, R., De Vries, R.S.M., and De Boer, A.H. (2013). Effect of Salt Stress on Growth, Na+ Accumulation and Proline Metabolism in Potato (Solanum tuberosum) Cultivars. PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0060183"},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"plw055","DOI":"10.1093\/aobpla\/plw055","article-title":"Effect of Salt Stress on Ion Concentration, Proline Content, Antioxidant Enzyme Activities and Gene Expression in Tomato Cultivars","volume":"8","author":"Gharsallah","year":"2016","journal-title":"AoB Plants"},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"188","DOI":"10.1038\/newbio236188a0","article-title":"Proline Accumulation and Varietal Adaptability to Drought in Barley: A Potential Metabolic Measure of Drought Resistance","volume":"236","author":"Singh","year":"1972","journal-title":"Nat. New Biol."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"720","DOI":"10.2135\/cropsci1977.0011183X001700050012x","article-title":"Evaluation of Free Proline Accumulation as an Index of Drought Resistance Using Two Contrasting Barley Cultivars","volume":"17","author":"Hanson","year":"1977","journal-title":"Crop Sci."},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"17","DOI":"10.1080\/02571862.1996.10634368","article-title":"Evaluation of Proline Accumulation as an Indicator of Drought Tolerance in Spring Wheat Cultivars","volume":"13","year":"1996","journal-title":"S. Afr. J. Plant Soil"},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"35","DOI":"10.1007\/s11120-020-00727-w","article-title":"Proline-Stimulated Signaling Primarily Targets the Chlorophyll Degradation Pathway and Photosynthesis Associated Processes to Cope with Short-Term Water Deficit in Maize","volume":"144","author":"Demiralay","year":"2020","journal-title":"Photosynth. Res."},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"98","DOI":"10.1016\/S0254-6299(16)30780-3","article-title":"Root Growth, Water Relations, Abscisic Acid and Proline Levels of Drought-Resistant and Drought-Sensitive Maize Cultivars in Response to Water Stress","volume":"59","author":"Cress","year":"1993","journal-title":"S. Afr. J. Bot."},{"key":"ref_101","doi-asserted-by":"crossref","unstructured":"Al Hassan, M., Morosan, M., L\u00f3pez-Gresa, M., Prohens, J., Vicente, O., and Boscaiu, M. (2016). Salinity-Induced Variation in Biochemical Markers Provides Insight into the Mechanisms of Salt Tolerance in Common (Phaseolus vulgaris) and Runner (P. coccineus) Beans. Int. J. Mol. Sci., 17.","DOI":"10.3390\/ijms17091582"},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"247","DOI":"10.24190\/ISSN2564-615X\/2017\/03.09","article-title":"Comparative Analysis of Drought Responses in Phaseolus vulgaris (Common Bean) and P. coccineus (Runner Bean) Cultivars","volume":"1","author":"Morosan","year":"2017","journal-title":"EuroBiotech J."},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"43","DOI":"10.1016\/j.molp.2022.09.007","article-title":"Metabolomics-Centered Mining of Plant Metabolic Diversity and Function: Past Decade and Future Perspectives","volume":"16","author":"Shen","year":"2023","journal-title":"Mol. Plant"},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"949","DOI":"10.1093\/plphys\/kiad357","article-title":"Advances in Single-Cell Metabolomics to Unravel Cellular Heterogeneity in Plant Biology","volume":"193","author":"Pandian","year":"2023","journal-title":"Plant Physiol."},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"1911","DOI":"10.1111\/nph.19734","article-title":"Isotopically Nonstationary Metabolic Flux Analysis of Plants: Recent Progress and Future Opportunities","volume":"242","author":"Koley","year":"2024","journal-title":"New Phytol."}],"container-title":["Plants"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2223-7747\/15\/16\/2478\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,17]],"date-time":"2026-08-17T08:57:59Z","timestamp":1786957079000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2223-7747\/15\/16\/2478"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,8,15]]},"references-count":105,"journal-issue":{"issue":"16","published-online":{"date-parts":[[2026,8]]}},"alternative-id":["plants15162478"],"URL":"https:\/\/doi.org\/10.3390\/plants15162478","relation":{},"ISSN":["2223-7747"],"issn-type":[{"value":"2223-7747","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,8,15]]}}}