{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,28]],"date-time":"2026-08-28T00:18:00Z","timestamp":1787876280240,"version":"build-2784847793"},"reference-count":49,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,6,13]],"date-time":"2026-06-13T00:00:00Z","timestamp":1781308800000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"},{"start":{"date-parts":[[2026,8,28]],"date-time":"2026-08-28T00:00:00Z","timestamp":1787875200000},"content-version":"vor","delay-in-days":76,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-nd\/4.0"}],"funder":[{"DOI":"10.13039\/501100003500","name":"Universit\u00e0 degli Studi di Padova","doi-asserted-by":"publisher","award":["2025DAFNAE1SIDPROGETTI-0001, 2025DAFNAE1DOR-00533"],"award-info":[{"award-number":["2025DAFNAE1SIDPROGETTI-0001, 2025DAFNAE1DOR-00533"]}],"id":[{"id":"10.13039\/501100003500","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Sci Rep"],"DOI":"10.1038\/s41598-026-56263-5","type":"journal-article","created":{"date-parts":[[2026,6,13]],"date-time":"2026-06-13T15:24:20Z","timestamp":1781364260000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Sporulation generates stress-dependent phenotypic variability in the heterothallic industrial Saccharomyces cerevisiae Ethanol Red"],"prefix":"10.1038","volume":"16","author":[{"given":"Viviana","family":"Paulon","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ameya Pankaj","family":"Gupte","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Katarina","family":"Stojkov","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chiara","family":"Ruspi","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Debora","family":"Casagrande Pierantoni","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Marina","family":"Basaglia","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Gianluigi","family":"Cardinali","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5650-4422","authenticated-orcid":false,"given":"Lorenzo","family":"Favaro","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Laura","family":"Corte","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,6,13]]},"reference":[{"key":"56263_CR1","doi-asserted-by":"publisher","first-page":"920","DOI":"10.1080\/07388551.2022.2072268","volume":"43","author":"PV Attfield","year":"2023","unstructured":"Attfield, P. V. Crucial aspects of metabolism and cell biology relating to industrial production and processing of Saccharomyces biomass. Crit. Rev. Biotechnol. 43, 920\u2013937. https:\/\/doi.org\/10.1080\/07388551.2022.2072268 (2023).","journal-title":"Crit. Rev. Biotechnol."},{"key":"56263_CR2","doi-asserted-by":"publisher","first-page":"1","DOI":"10.3934\/microbiol.2020001","volume":"6","author":"M Parapouli","year":"2020","unstructured":"Parapouli, M., Vasileiadis, A., Afendra, A-S. & Hatziloukas, E. Saccharomyces cerevisiae and its industrial applications. AIMS Microbiol. 6, 1\u201331. https:\/\/doi.org\/10.3934\/microbiol.2020001 (2020).","journal-title":"AIMS Microbiol."},{"key":"56263_CR3","doi-asserted-by":"publisher","first-page":"61","DOI":"10.1186\/s13068-024-02510-8","volume":"17","author":"C Liu","year":"2024","unstructured":"Liu, C., Choi, B., Efimova, E., Nyg\u00e5rd, Y. & Santala, S. Enhanced upgrading of lignocellulosic substrates by coculture of Saccharomyces cerevisiae and Acinetobacter baylyi ADP1. Biotechnol. Biofuels Bioprod. 17, 61. https:\/\/doi.org\/10.1186\/s13068-024-02510-8 (2024).","journal-title":"Biotechnol. Biofuels Bioprod."},{"key":"56263_CR4","doi-asserted-by":"publisher","first-page":"R381","DOI":"10.1016\/j.cub.2019.04.025","volume":"29","author":"J Steensels","year":"2019","unstructured":"Steensels, J., Gallone, B., Voordeckers, K. & Verstrepen, K. J. Domestication of Industrial Microbes. Curr. Biol. 29, R381\u2013R393. https:\/\/doi.org\/10.1016\/j.cub.2019.04.025 (2019).","journal-title":"Curr. Biol."},{"key":"56263_CR5","doi-asserted-by":"publisher","first-page":"1397","DOI":"10.1016\/j.cell.2016.08.020","volume":"166","author":"B Gallone","year":"2016","unstructured":"Gallone, B. et al. Domestication and Divergence of Saccharomyces cerevisiae Beer Yeasts. Cell 166, 1397\u20131410e16. https:\/\/doi.org\/10.1016\/j.cell.2016.08.020 (2016).","journal-title":"Cell"},{"key":"56263_CR6","doi-asserted-by":"publisher","first-page":"800","DOI":"10.1080\/07388551.2019.1619157","volume":"39","author":"L Favaro","year":"2019","unstructured":"Favaro, L., Jansen, T. & van Zyl, W. H. Exploring industrial and natural Saccharomyces cerevisiae strains for the bio-based economy from biomass: the case of bioethanol. Crit. Rev. Biotechnol. 39, 800\u2013816. https:\/\/doi.org\/10.1080\/07388551.2019.1619157 (2019).","journal-title":"Crit. Rev. Biotechnol."},{"key":"56263_CR7","doi-asserted-by":"publisher","first-page":"264","DOI":"10.3390\/jof9020264","volume":"9","author":"AP Gupte","year":"2023","unstructured":"Gupte, A. P. et al. Renewing Lost Genetic Variability with a Classical Yeast Genetics Approach. J. Fungi. 9, 264. https:\/\/doi.org\/10.3390\/jof9020264 (2023).","journal-title":"J. Fungi"},{"key":"56263_CR8","doi-asserted-by":"publisher","first-page":"947","DOI":"10.1111\/1574-6976.12073","volume":"38","author":"J Steensels","year":"2014","unstructured":"Steensels, J. et al. Improving industrial yeast strains: exploiting natural and artificial diversity. FEMS Microbiol. Rev. 38, 947\u2013995. https:\/\/doi.org\/10.1111\/1574-6976.12073 (2014).","journal-title":"FEMS Microbiol. Rev."},{"key":"56263_CR9","doi-asserted-by":"publisher","first-page":"102","DOI":"10.1016\/j.ijfoodmicro.2012.03.008","volume":"156","author":"L P\u00e9rez-Trav\u00e9s","year":"2012","unstructured":"P\u00e9rez-Trav\u00e9s, L., Lopes, C. A., Barrio, E. & Querol, A. Evaluation of different genetic procedures for the generation of artificial hybrids in Saccharomyces genus for winemaking. Int. J. Food Microbiol. 156, 102\u2013111. https:\/\/doi.org\/10.1016\/j.ijfoodmicro.2012.03.008 (2012).","journal-title":"Int. J. Food Microbiol."},{"key":"56263_CR10","doi-asserted-by":"publisher","first-page":"85","DOI":"10.1007\/s13213-010-0086-4","volume":"61","author":"M Sipiczki","year":"2011","unstructured":"Sipiczki, M. Diversity, variability and fast adaptive evolution of the wine yeast (Saccharomyces cerevisiae) genome\u2014a review. Ann. Microbiol. 61, 85\u201393. https:\/\/doi.org\/10.1007\/s13213-010-0086-4 (2011).","journal-title":"Ann. Microbiol."},{"key":"56263_CR11","doi-asserted-by":"publisher","first-page":"121","DOI":"10.1111\/j.1365-294X.2009.04436.x","volume":"19","author":"S-L Ezov Tk, Chang","year":"2010","unstructured":"Ezov Tk, Chang, S-L. et al. Heterothallism in Saccharomyces cerevisiae isolates from nature: effect of HO locus on the mode of reproduction. Mol. Ecol. 19, 121\u2013131. https:\/\/doi.org\/10.1111\/j.1365-294X.2009.04436.x (2010).","journal-title":"Mol. Ecol."},{"key":"56263_CR12","doi-asserted-by":"publisher","first-page":"230","DOI":"10.3390\/genes13020230","volume":"13","author":"F-Y Bai","year":"2022","unstructured":"Bai, F-Y., Han, D-Y., Duan, S-F. & Wang, Q-M. The Ecology and Evolution of the Baker\u2019s Yeast Saccharomyces cerevisiae. Genes 13, 230. https:\/\/doi.org\/10.3390\/genes13020230 (2022).","journal-title":"Genes"},{"key":"56263_CR13","doi-asserted-by":"publisher","first-page":"337","DOI":"10.1038\/nature07743","volume":"458","author":"G Liti","year":"2009","unstructured":"Liti, G. et al. Population genomics of domestic and wild yeasts. Nature 458, 337\u2013341. https:\/\/doi.org\/10.1038\/nature07743 (2009).","journal-title":"Nature"},{"key":"56263_CR14","doi-asserted-by":"publisher","first-page":"148","DOI":"10.1016\/j.copbio.2017.08.005","volume":"49","author":"B Gallone","year":"2018","unstructured":"Gallone, B. et al. Origins, evolution, domestication and diversity of Saccharomyces beer yeasts. Curr. Opin. Biotechnol. 49, 148\u2013155. https:\/\/doi.org\/10.1016\/j.copbio.2017.08.005 (2018).","journal-title":"Curr. Opin. Biotechnol."},{"key":"56263_CR15","doi-asserted-by":"publisher","first-page":"117","DOI":"10.1186\/s13068-024-02562-w","volume":"17","author":"J Ren","year":"2024","unstructured":"Ren, J. et al. Furfural tolerance of mutant Saccharomyces cerevisiae selected via ionizing radiation combined with adaptive laboratory evolution. Biotechnol. Biofuels Bioprod. 17, 117. https:\/\/doi.org\/10.1186\/s13068-024-02562-w (2024).","journal-title":"Biotechnol. Biofuels Bioprod."},{"key":"56263_CR16","doi-asserted-by":"publisher","first-page":"22875","DOI":"10.1038\/s41598-024-74174-1","volume":"14","author":"F Zuo","year":"2024","unstructured":"Zuo, F., Wu, Y., Sun, Y., Xie, C. & Tang, Y. Mechanism of enhanced salt tolerance in Saccharomyces cerevisiae by CRZ1 overexpression. Sci. Rep. 14, 22875. https:\/\/doi.org\/10.1038\/s41598-024-74174-1 (2024).","journal-title":"Sci. Rep."},{"key":"56263_CR17","doi-asserted-by":"publisher","first-page":"133528","DOI":"10.1016\/j.biortech.2025.133528","volume":"441","author":"R My","year":"2026","unstructured":"My, R. et al. Formic acid resistance in Saccharomyces cerevisiae strains: the role of SAT4 in a proposed molecular model. Bioresour Technol. 441, 133528. https:\/\/doi.org\/10.1016\/j.biortech.2025.133528 (2026).","journal-title":"Bioresour Technol."},{"key":"56263_CR18","doi-asserted-by":"publisher","first-page":"25","DOI":"10.1016\/S0960-8524(99)00161-3","volume":"74","author":"E Palmqvist","year":"2000","unstructured":"Palmqvist, E. & Hahn-H\u00e4gerdal, B. Fermentation of lignocellulosic hydrolysates. II: inhibitors and mechanisms of inhibition. Bioresour Technol. 74, 25\u201333. https:\/\/doi.org\/10.1016\/S0960-8524(99)00161-3 (2000).","journal-title":"Bioresour Technol."},{"key":"56263_CR19","doi-asserted-by":"publisher","first-page":"1545","DOI":"10.1039\/C9SE00029A","volume":"3","author":"D Greetham","year":"2019","unstructured":"Greetham, D., Zaky, A. S. & Du, C. Exploring the tolerance of marine yeast to inhibitory compounds for improving bioethanol production. Sustain. Energy Fuels. 3, 1545\u20131553. https:\/\/doi.org\/10.1039\/C9SE00029A (2019).","journal-title":"Sustain. Energy Fuels"},{"key":"56263_CR20","doi-asserted-by":"publisher","first-page":"1333777","DOI":"10.3389\/fmicb.2023.1333777","volume":"14","author":"L Yao","year":"2024","unstructured":"Yao, L. et al. Adaptive laboratory evolution to obtain furfural tolerant Saccharomyces cerevisiae for bioethanol production and the underlying mechanism. Front. Microbiol. 14, 1333777. https:\/\/doi.org\/10.3389\/fmicb.2023.1333777 (2024).","journal-title":"Front. Microbiol."},{"key":"56263_CR21","doi-asserted-by":"publisher","first-page":"630","DOI":"10.1128\/aem.61.2.630-638.1995","volume":"61","author":"AC Cod\u00f3n","year":"1995","unstructured":"Cod\u00f3n, A. C., Gasent-Ram\u00edrez, J. M. & Ben\u00edtez, T. Factors which affect the frequency of sporulation and tetrad formation in Saccharomyces cerevisiae baker\u2019s yeasts. Appl. Environ. Microbiol. 61, 630\u2013638. https:\/\/doi.org\/10.1128\/aem.61.2.630-638.1995 (1995).","journal-title":"Appl. Environ. Microbiol."},{"key":"56263_CR22","doi-asserted-by":"publisher","first-page":"657","DOI":"10.1046\/j.1365-2672.2000.01163.x","volume":"89","author":"M Budroni","year":"2000","unstructured":"Budroni, M., Giordano, G., Pinna, G. & Farris, G. A. A genetic study of natural flor strains of Saccharomyces cerevisiae isolated during biological ageing from Sardinian wines. J. Appl. Microbiol. 89, 657\u2013662. https:\/\/doi.org\/10.1046\/j.1365-2672.2000.01163.x (2000).","journal-title":"J. Appl. Microbiol."},{"key":"56263_CR23","doi-asserted-by":"publisher","first-page":"999","DOI":"10.1002\/yea.1808","volume":"27","author":"M Bahalul","year":"2010","unstructured":"Bahalul, M., Kaneti, G. & Kashi, Y. Ether-zymolyase ascospore isolation procedure: an efficient protocol for ascospores isolation in Saccharomyces cerevisiae yeast. Yeast 27, 999\u20131003. https:\/\/doi.org\/10.1002\/yea.1808 (2010).","journal-title":"Yeast"},{"key":"56263_CR24","doi-asserted-by":"publisher","first-page":"768562","DOI":"10.3389\/fmicb.2021.768562","volume":"12","author":"N Gronchi","year":"2021","unstructured":"Gronchi, N. et al. Natural Saccharomyces cerevisiae strain reveals peculiar genomic traits for starch-to-bioethanol production: the design of an amylolytic consolidated bioprocessing yeast. Front. Microbiol. 12, 768562. https:\/\/doi.org\/10.3389\/fmicb.2021.768562 (2021).","journal-title":"Front. Microbiol."},{"key":"56263_CR25","doi-asserted-by":"publisher","first-page":"63","DOI":"10.1016\/j.nbt.2024.12.004","volume":"85","author":"R My","year":"2025","unstructured":"My, R. et al. Unveiling the fitness of Saccharomyces cerevisiae strains for lignocellulosic bioethanol: a genomic exploration through fermentation stress tests. New. Biotechnol. 85, 63\u201374. https:\/\/doi.org\/10.1016\/j.nbt.2024.12.004 (2025).","journal-title":"New. Biotechnol."},{"key":"56263_CR26","doi-asserted-by":"publisher","first-page":"3406","DOI":"10.3390\/ijms23063406","volume":"23","author":"C Du","year":"2022","unstructured":"Du, C., Li, Y., Xiang, R. & Yuan, W. Formate Dehydrogenase Improves the Resistance to Formic Acid and Acetic Acid Simultaneously in Saccharomyces cerevisiae. Int. J. Mol. Sci. 23, 3406. https:\/\/doi.org\/10.3390\/ijms23063406 (2022).","journal-title":"Int. J. Mol. Sci."},{"key":"56263_CR27","doi-asserted-by":"publisher","first-page":"34","DOI":"10.1007\/s11274-021-03222-z","volume":"38","author":"L Zeng","year":"2022","unstructured":"Zeng, L. et al. Transcriptomic analysis of formic acid stress response in Saccharomyces cerevisiae. World J. Microbiol. Biotechnol. 38, 34. https:\/\/doi.org\/10.1007\/s11274-021-03222-z (2022).","journal-title":"World J. Microbiol. Biotechnol."},{"key":"56263_CR28","doi-asserted-by":"publisher","first-page":"e0121623","DOI":"10.1128\/spectrum.01216-23","volume":"11","author":"L Qi","year":"2023","unstructured":"Qi, L. et al. Nonlethal Furfural Exposure Causes Genomic Alterations and Adaptability Evolution in Saccharomyces cerevisiae. Microbiol. Spectr. 11, e0121623. https:\/\/doi.org\/10.1128\/spectrum.01216-23 (2023).","journal-title":"Microbiol. Spectr."},{"key":"56263_CR29","doi-asserted-by":"publisher","first-page":"376","DOI":"10.3389\/fbioe.2019.00376","volume":"7","author":"L Favaro","year":"2019","unstructured":"Favaro, L. et al. Metabolomic Alterations Do Not Induce Metabolic Burden in the Industrial Yeast M2n[pBKD2-Pccbgl1]-C1 Engineered by Multiple \u03b4-Integration of a Fungal \u03b2-Glucosidase Gene. Front. Bioeng. Biotechnol. 7, 376. https:\/\/doi.org\/10.3389\/fbioe.2019.00376 (2019).","journal-title":"Front. Bioeng. Biotechnol."},{"key":"56263_CR30","doi-asserted-by":"publisher","first-page":"293","DOI":"10.1007\/s00374-024-01885-y","volume":"61","author":"C Ruspi","year":"2025","unstructured":"Ruspi, C. et al. Plant Growth-Promoting Yeasts (PGPYs) as a sustainable solution to mitigate salt-induced stress on zucchini plant growth. Biol. Fertil. Soils. 61, 293\u2013309. https:\/\/doi.org\/10.1007\/s00374-024-01885-y (2025).","journal-title":"Biol. Fertil. Soils"},{"key":"56263_CR31","doi-asserted-by":"publisher","first-page":"2099","DOI":"10.1039\/C0AN00127A","volume":"135","author":"L Roscini","year":"2010","unstructured":"Roscini, L. et al. Influence of cell geometry and number of replicas in the reproducibility of whole cell FTIR analysis. Analyst 135, 2099\u20132105. https:\/\/doi.org\/10.1039\/C0AN00127A (2010).","journal-title":"Analyst"},{"key":"56263_CR32","doi-asserted-by":"publisher","first-page":"e55160","DOI":"10.7554\/eLife.55160","volume":"9","author":"S Kamrad","year":"2020","unstructured":"Kamrad, S. et al. Pyphe, a python toolbox for assessing microbial growth and cell viability in high-throughput colony screens. eLife 9, e55160. https:\/\/doi.org\/10.7554\/eLife.55160 (2020).","journal-title":"eLife"},{"key":"56263_CR33","unstructured":"R Core Team. R: A language and environment for statistical computing (R Foundation for Statistical Computing, 2024)."},{"key":"56263_CR34","doi-asserted-by":"publisher","first-page":"140","DOI":"10.3390\/metabo10040140","volume":"10","author":"L Corte","year":"2020","unstructured":"Corte, L. et al. Delta-Integration of Single Gene Shapes the Whole Metabolomic Short-Term Response to Ethanol of Recombinant Saccharomyces cerevisiae Strains. Metabolites 10, 140. https:\/\/doi.org\/10.3390\/metabo10040140 (2020).","journal-title":"Metabolites"},{"key":"56263_CR35","doi-asserted-by":"publisher","first-page":"fow072","DOI":"10.1093\/femsyr\/fow072","volume":"16","author":"Z Guo","year":"2016","unstructured":"Guo, Z. & Olsson, L. Physiological responses to acid stress by Saccharomyces cerevisiae when applying high initial cell density. FEMS Yeast Res. 16, fow072. https:\/\/doi.org\/10.1093\/femsyr\/fow072 (2016).","journal-title":"FEMS Yeast Res."},{"key":"56263_CR36","doi-asserted-by":"publisher","first-page":"106099","DOI":"10.1016\/j.ibiod.2025.106099","volume":"202","author":"Z Zhang","year":"2025","unstructured":"Zhang, Z. et al. Enhanced tolerance of Saccharomyces cerevisiae to industrial inhibitors through multi-transcription factor engineering for environmental and biotechnological applications. Int. Biodeterior. Biodegrad. 202, 106099. https:\/\/doi.org\/10.1016\/j.ibiod.2025.106099 (2025).","journal-title":"Int. Biodeterior. Biodegrad"},{"key":"56263_CR37","doi-asserted-by":"publisher","first-page":"300","DOI":"10.1128\/MMBR.66.2.300-372.2002","volume":"66","author":"S Hohmann","year":"2002","unstructured":"Hohmann, S. Osmotic stress signaling and osmoadaptation in yeasts. Microbiol. Mol. Biol. Rev. 66, 300\u2013372. https:\/\/doi.org\/10.1128\/MMBR.66.2.300-372.2002 (2002).","journal-title":"Microbiol. Mol. Biol. Rev."},{"key":"56263_CR38","doi-asserted-by":"publisher","first-page":"28","DOI":"10.1186\/s13068-022-02127-9","volume":"15","author":"B Li","year":"2022","unstructured":"Li, B., Liu, N. & Zhao, X. Response mechanisms of Saccharomyces cerevisiae to the stress factors present in lignocellulose hydrolysate and strategies for constructing robust strains. Biotechnol. Biofuels. 15, 28. https:\/\/doi.org\/10.1186\/s13068-022-02127-9 (2022).","journal-title":"Biotechnol. Biofuels"},{"key":"56263_CR39","doi-asserted-by":"publisher","first-page":"foac035","DOI":"10.1093\/femsyr\/foac035","volume":"22","author":"A Blomberg","year":"2022","unstructured":"Blomberg, A. Yeast osmoregulation \u2013 glycerol still in pole position. FEMS Yeast Res. 22, foac035. https:\/\/doi.org\/10.1093\/femsyr\/foac035 (2022).","journal-title":"FEMS Yeast Res."},{"key":"56263_CR40","doi-asserted-by":"publisher","first-page":"104451","DOI":"10.1016\/j.fbio.2024.104451","volume":"60","author":"A Chen","year":"2024","unstructured":"Chen, A. et al. Osmotic tolerance in Saccharomyces cerevisiae: Implications for food and bioethanol industries. Food Biosci. 60, 104451. https:\/\/doi.org\/10.1016\/j.fbio.2024.104451 (2024).","journal-title":"Food Biosci."},{"key":"56263_CR41","doi-asserted-by":"publisher","first-page":"1528","DOI":"10.3390\/horticulturae11121528","volume":"11","author":"K Stojkov","year":"2025","unstructured":"Stojkov, K. et al. Decoupling Microbial Activity from Metabolite Action: A Comparative Assessment of EM Technology and Its Cell-Free Extract as Nature-Based Solutions for Plant Biostimulation. Horticulturae 11, 1528. https:\/\/doi.org\/10.3390\/horticulturae11121528 (2025).","journal-title":"Horticulturae"},{"key":"56263_CR42","doi-asserted-by":"publisher","first-page":"953479","DOI":"10.3389\/fmicb.2022.953479","volume":"13","author":"RA Ribeiro","year":"2022","unstructured":"Ribeiro, R. A., Bourbon-Melo, N. & S\u00e1-Correia, I. The cell wall and the response and tolerance to stresses of biotechnological relevance in yeasts. Front. Microbiol. 13, 953479. https:\/\/doi.org\/10.3389\/fmicb.2022.953479 (2022).","journal-title":"Front. Microbiol."},{"key":"56263_CR43","doi-asserted-by":"publisher","first-page":"53","DOI":"10.1016\/s0009-3084(98)00080-2","volume":"96","author":"JL Arrondo","year":"1998","unstructured":"Arrondo, J. L. & Go\u00f1i, F. M. Infrared studies of protein-induced perturbation of lipids in lipoproteins and membranes. Chem. Phys. Lipids. 96, 53\u201368. https:\/\/doi.org\/10.1016\/s0009-3084(98)00080-2 (1998).","journal-title":"Chem. Phys. Lipids"},{"key":"56263_CR44","doi-asserted-by":"publisher","first-page":"23982","DOI":"10.1038\/s41598-024-75108-7","volume":"14","author":"A Shirvanyan","year":"2024","unstructured":"Shirvanyan, A. & Trchounian, K. Sodium transport and redox regulation in Saccharomyces cerevisiae under osmotic stress depending on oxygen availability. Sci. Rep. 14, 23982. https:\/\/doi.org\/10.1038\/s41598-024-75108-7 (2024).","journal-title":"Sci. Rep."},{"key":"56263_CR45","doi-asserted-by":"publisher","first-page":"189","DOI":"10.1007\/BF02786949","volume":"56","author":"SC Sharma","year":"1996","unstructured":"Sharma, S. C., Raj, D., Forouzandeh, M. & Bansal, M. P. Salt-induced changes in lipid composition and ethanol tolerance in Saccharomyces cerevisiae. Appl. Biochem. Biotechnol. 56, 189\u2013195. https:\/\/doi.org\/10.1007\/BF02786949 (1996).","journal-title":"Appl. Biochem. Biotechnol."},{"key":"56263_CR46","doi-asserted-by":"publisher","first-page":"110","DOI":"10.1128\/AEM.01360-06","volume":"73","author":"S Rodr\u00edguez-Vargas","year":"2007","unstructured":"Rodr\u00edguez-Vargas, S., S\u00e1nchez-Garc\u00eda, A., Mart\u00ednez-Rivas, J. M., Prieto, J. A. & Randez-Gil, F. Fluidization of membrane lipids enhances the tolerance of Saccharomyces cerevisiae to freezing and salt stress. Appl. Environ. Microbiol. 73, 110\u2013116. https:\/\/doi.org\/10.1128\/AEM.01360-06 (2007).","journal-title":"Appl. Environ. Microbiol."},{"key":"56263_CR47","doi-asserted-by":"publisher","first-page":"39","DOI":"10.1016\/S0009-3084(02)00005-1","volume":"115","author":"H Binder","year":"2002","unstructured":"Binder, H. & Zsch\u00f6rnig, O. The effect of metal cations on the phase behavior and hydration characteristics of phospholipid membranes. Chem. Phys. Lipids. 115, 39\u201361. https:\/\/doi.org\/10.1016\/S0009-3084(02)00005-1 (2002).","journal-title":"Chem. Phys. Lipids"},{"key":"56263_CR48","doi-asserted-by":"publisher","first-page":"2630","DOI":"10.3390\/ijms24032630","volume":"24","author":"B Gieroba","year":"2023","unstructured":"Gieroba, B. et al. Application of vibrational spectroscopic techniques in the study of the natural polysaccharides and their cross-linking process. Int. JMol Sci. 24, 2630. https:\/\/doi.org\/10.3390\/ijms24032630 (2023).","journal-title":"Int. JMol Sci."},{"key":"56263_CR49","doi-asserted-by":"publisher","first-page":"179","DOI":"10.1111\/j.1574-6968.2001.tb10586.x","volume":"197","author":"A Galichet","year":"2001","unstructured":"Galichet, A., Sockalingum, G. D., Belarbi, A. & Manfait, M. FTIR spectroscopic analysis of Saccharomyces cerevisiae cell walls: study of an anomalous strain exhibiting a pink-colored cell phenotype. FEMS Microbiol. Lett. 197, 179\u2013186. https:\/\/doi.org\/10.1111\/j.1574-6968.2001.tb10586.x (2001).","journal-title":"FEMS Microbiol. Lett."}],"container-title":["Scientific Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41598-026-56263-5","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-026-56263-5.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-026-56263-5.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,27]],"date-time":"2026-08-27T23:45:53Z","timestamp":1787874353000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41598-026-56263-5"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,6,13]]},"references-count":49,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["56263"],"URL":"https:\/\/doi.org\/10.1038\/s41598-026-56263-5","relation":{},"ISSN":["2045-2322"],"issn-type":[{"value":"2045-2322","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,6,13]]},"assertion":[{"value":"9 April 2026","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"29 May 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"13 June 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","label":"Competing interests","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"Not applicable. All the authors agree to the submission and publication of this manuscript.","order":2,"name":"Ethics","label":"Consent for publication","group":{"name":"EthicsHeading","label":"Declarations"}}],"article-number":"27046"}}