{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,30]],"date-time":"2026-09-30T04:59:08Z","timestamp":1790744348260,"version":"4.1.0"},"reference-count":58,"publisher":"MDPI AG","issue":"19","license":[{"start":{"date-parts":[[2026,9,26]],"date-time":"2026-09-26T00:00:00Z","timestamp":1790380800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"name":"Ministry of Science and Higher Education of the Russian Federation","award":["075-15-2025-466"],"award-info":[{"award-number":["075-15-2025-466"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Int. J. Mol. Sci.","IJMS"],"abstract":"<jats:p>Autophagy is a conserved lysosome-dependent process essential for cellular homeostasis, and pharmacological activation of autophagy has emerged as a promising therapeutic strategy for age-related, neurodegenerative, metabolic, and malignant diseases. Despite the growing interest in natural autophagy modulators, lichens remain largely unexplored as a source of autophagy-inducing compounds. Here, we tested ethanolic extracts from four commercially available lichens using LC3-II accumulation as a primary readout. Usnea sp. was identified as the most active species. Activity-guided fractionation followed by HRMS and NMR analyses identified salazinic acid as the active metabolite. Salazinic acid increased LC3-II accumulation, enhanced autophagic flux in the presence of bafilomycin A1, and promoted autophagosome formation as confirmed by transmission electron microscopy. The compound exhibited low cytotoxicity in human cell lines at concentrations exceeding those required for autophagy induction. Mechanistically, salazinic acid activated the phosphorylation of AMPK, ULK1, and AKT; however, it retained its ability to stimulate autophagic flux in AMPK- and ULK1-deficient cells, indicating that its activity is not strictly dependent on the canonical AMPK\u2013ULK1 signaling pathway. Collectively, our findings identify salazinic acid as a previously unrecognized inducer of productive autophagic flux and demonstrate that lichen secondary metabolites represent a promising and largely untapped source of novel autophagy-modulating compounds.<\/jats:p>","DOI":"10.3390\/ijms27198621","type":"journal-article","created":{"date-parts":[[2026,9,28]],"date-time":"2026-09-28T16:50:54Z","timestamp":1790614254000},"page":"8621","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["The Reindeer\u2019s Secret: Salazinic Acid from Lichens as a Candidate Modulator of Autophagy"],"prefix":"10.3390","volume":"27","author":[{"ORCID":"https:\/\/orcid.org\/0009-0004-4722-8309","authenticated-orcid":false,"given":"Anastasiia K.","family":"Veretina","sequence":"first","affiliation":[{"name":"Center for Bio- and Medical Technologies, 121205 Moscow, Russia"},{"name":"Belozersky Institute of Physico-Chemical Biology, Lomonosov Moscow State University, 119991 Moscow, Russia"},{"name":"Department of Chemistry, Lomonosov Moscow State University, 119991 Moscow, Russia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8961-5890","authenticated-orcid":false,"given":"Vera A.","family":"Alferova","sequence":"additional","affiliation":[{"name":"Belozersky Institute of Physico-Chemical Biology, Lomonosov Moscow State University, 119991 Moscow, Russia"},{"name":"Shemyakin-Ovchinnikov Institute of Bioorganic Chemistry, Russian Academy of Science, 117997 Moscow, Russia"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"methodology","vocabulary":"credit"},{"role":"investigation","vocabulary":"credit"},{"role":"visualization","vocabulary":"credit"},{"role":"writing-original-draft","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1818-6590","authenticated-orcid":false,"given":"Ekaterina A.","family":"Guseva","sequence":"additional","affiliation":[{"name":"Belozersky Institute of Physico-Chemical Biology, Lomonosov Moscow State University, 119991 Moscow, Russia"},{"name":"Department of Chemistry, Lomonosov Moscow State University, 119991 Moscow, Russia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Anton R.","family":"Izzi","sequence":"additional","affiliation":[{"name":"Belozersky Institute of Physico-Chemical Biology, Lomonosov Moscow State University, 119991 Moscow, Russia"},{"name":"Faculty of Bioengineering and Bioinformatics, Lomonosov Moscow State University, 119991 Moscow, Russia"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"investigation","vocabulary":"credit"}]},{"given":"Arina A.","family":"Nikandrova","sequence":"additional","affiliation":[{"name":"Center for Bio- and Medical Technologies, 121205 Moscow, Russia"},{"name":"Belozersky Institute of Physico-Chemical Biology, Lomonosov Moscow State University, 119991 Moscow, Russia"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"investigation","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6405-6367","authenticated-orcid":false,"given":"Gulnara K.","family":"Kudryakova","sequence":"additional","affiliation":[{"name":"Shemyakin-Ovchinnikov Institute of Bioorganic Chemistry, Russian Academy of Science, 117997 Moscow, Russia"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"investigation","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0009-0007-8527-3540","authenticated-orcid":false,"given":"Mikhail Y.","family":"Zhitlov","sequence":"additional","affiliation":[{"name":"Shemyakin-Ovchinnikov Institute of Bioorganic Chemistry, Russian Academy of Science, 117997 Moscow, Russia"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"investigation","vocabulary":"credit"}]},{"given":"Julia A.","family":"Golubeva","sequence":"additional","affiliation":[{"name":"Center for Bio- and Medical Technologies, 121205 Moscow, Russia"},{"name":"Belozersky Institute of Physico-Chemical Biology, Lomonosov Moscow State University, 119991 Moscow, Russia"},{"name":"Department of Chemistry, Lomonosov Moscow State University, 119991 Moscow, Russia"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"investigation","vocabulary":"credit"}]},{"given":"Valeriya B.","family":"Vays","sequence":"additional","affiliation":[{"name":"Belozersky Institute of Physico-Chemical Biology, Lomonosov Moscow State University, 119991 Moscow, Russia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Irina M.","family":"Vangeli","sequence":"additional","affiliation":[{"name":"Belozersky Institute of Physico-Chemical Biology, Lomonosov Moscow State University, 119991 Moscow, Russia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yulia M.","family":"Khabibulina","sequence":"additional","affiliation":[{"name":"Faculty of Bioengineering and Bioinformatics, Lomonosov Moscow State University, 119991 Moscow, Russia"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"investigation","vocabulary":"credit"}]},{"given":"Lora E.","family":"Bakeeva","sequence":"additional","affiliation":[{"name":"Belozersky Institute of Physico-Chemical Biology, Lomonosov Moscow State University, 119991 Moscow, Russia"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"methodology","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8069-7087","authenticated-orcid":false,"given":"Anna A.","family":"Lukianova","sequence":"additional","affiliation":[{"name":"Shemyakin-Ovchinnikov Institute of Bioorganic Chemistry, Russian Academy of Science, 117997 Moscow, Russia"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"investigation","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9921-8031","authenticated-orcid":false,"given":"Lyamzaev G.","family":"Konstantin","sequence":"additional","affiliation":[{"name":"Belozersky Institute of Physico-Chemical Biology, Lomonosov Moscow State University, 119991 Moscow, Russia"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5203-0787","authenticated-orcid":false,"given":"Dmitry S.","family":"Karpov","sequence":"additional","affiliation":[{"name":"Engelhardt Institute of Molecular Biology, Russian Academy of Sciences, 119991 Moscow, Russia"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"project-administration","vocabulary":"credit"},{"role":"funding-acquisition","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1413-9559","authenticated-orcid":false,"given":"Dmitrii A.","family":"Lukianov","sequence":"additional","affiliation":[{"name":"Center for Bio- and Medical Technologies, 121205 Moscow, Russia"},{"name":"Department of Chemistry, Lomonosov Moscow State University, 119991 Moscow, Russia"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"conceptualization","vocabulary":"credit"},{"role":"methodology","vocabulary":"credit"},{"role":"writing-original-draft","vocabulary":"credit"}]},{"given":"Olga A.","family":"Dontsova","sequence":"additional","affiliation":[{"name":"Center for Bio- and Medical Technologies, 121205 Moscow, Russia"},{"name":"Department of Chemistry, Lomonosov Moscow State University, 119991 Moscow, Russia"},{"name":"Shemyakin-Ovchinnikov Institute of Bioorganic Chemistry, Russian Academy of Science, 117997 Moscow, Russia"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"supervision","vocabulary":"credit"},{"role":"project-administration","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8866-1863","authenticated-orcid":false,"given":"Petr V.","family":"Sergiev","sequence":"additional","affiliation":[{"name":"Center for Bio- and Medical Technologies, 121205 Moscow, Russia"},{"name":"Belozersky Institute of Physico-Chemical Biology, Lomonosov Moscow State University, 119991 Moscow, Russia"},{"name":"Department of Chemistry, Lomonosov Moscow State University, 119991 Moscow, Russia"}],"role":[{"vocabulary":"crossref","role":"author"},{"role":"conceptualization","vocabulary":"credit"},{"role":"supervision","vocabulary":"credit"},{"role":"project-administration","vocabulary":"credit"}]}],"member":"1968","published-online":{"date-parts":[[2026,9,26]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"1159","DOI":"10.4161\/auto.20527","article-title":"Autophagy across the Eukaryotes: Is S. Cerevisiae the Odd One Out?","volume":"8","author":"King","year":"2012","journal-title":"Autophagy"},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"2570","DOI":"10.1128\/MCB.00166-09","article-title":"Hypoxia-Induced Autophagy Is Mediated through Hypoxia-Inducible Factor Induction of BNIP3 and BNIP3L via Their BH3 Domains","volume":"29","author":"Bellot","year":"2009","journal-title":"Mol. Cell. Biol."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"377","DOI":"10.1038\/cdd.2014.150","article-title":"Oxidative Stress and Autophagy: The Clash between Damage and Metabolic Needs","volume":"22","author":"Filomeni","year":"2015","journal-title":"Cell Death Differ."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"461","DOI":"10.1038\/nrm4024","article-title":"Autophagy at the Crossroads of Catabolism and Anabolism","volume":"16","author":"Kaur","year":"2015","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"9","DOI":"10.1038\/nrm3028","article-title":"Mechanisms of Mitophagy","volume":"12","author":"Youle","year":"2011","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"176","DOI":"10.4161\/auto.5269","article-title":"Promoting Basal Levels of Autophagy in the Nervous System Enhances Longevity and Oxidant Resistance in Adult Drosophila","volume":"4","author":"Simonsen","year":"2008","journal-title":"Autophagy"},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"858","DOI":"10.1038\/s41418-019-0480-9","article-title":"Autophagy and Disease: Unanswered Questions","volume":"27","author":"Yang","year":"2020","journal-title":"Cell Death Differ."},{"key":"ref_8","doi-asserted-by":"crossref","unstructured":"Ichimiya, T., Yamakawa, T., Hirano, T., Yokoyama, Y., Hayashi, Y., Hirayama, D., Wagatsuma, K., Itoi, T., and Nakase, H. (2020). Autophagy and Autophagy-Related Diseases: A Review. Int. J. Mol. Sci., 21.","DOI":"10.3390\/ijms21238974"},{"key":"ref_9","first-page":"1","article-title":"Natural Activators of Autophagy","volume":"89","author":"Pavlova","year":"2024","journal-title":"Biochemistry"},{"key":"ref_10","first-page":"27","article-title":"Synthetic Activators of Autophagy","volume":"89","author":"Guseva","year":"2024","journal-title":"Biochemistry"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"487","DOI":"10.1038\/nrd.2017.22","article-title":"Pharmacological Modulation of Autophagy: Therapeutic Potential and Persisting Obstacles","volume":"16","author":"Galluzzi","year":"2017","journal-title":"Nat. Rev. Drug Discov."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"14","DOI":"10.1172\/JCI73938","article-title":"Development of Autophagy Inducers in Clinical Medicine","volume":"125","author":"Levine","year":"2015","journal-title":"J. Clin. Investig."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"879","DOI":"10.1038\/nm.4132","article-title":"Urolithin A Induces Mitophagy and Prolongs Lifespan in C. Elegans and Increases Muscle Function in Rodents","volume":"22","author":"Ryu","year":"2016","journal-title":"Nat. Med."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"1305","DOI":"10.1038\/ncb1975","article-title":"Induction of Autophagy by Spermidine Promotes Longevity","volume":"11","author":"Eisenberg","year":"2009","journal-title":"Nat. Cell Biol."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"e10","DOI":"10.1038\/cddis.2009.8","article-title":"Caloric Restriction and Resveratrol Promote Longevity through the Sirtuin-1-Dependent Induction of Autophagy","volume":"1","author":"Morselli","year":"2010","journal-title":"Cell Death Dis."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"1377","DOI":"10.1124\/dmd.104.000885","article-title":"High absorption but very low bioavailability of oral resveratrol in humans","volume":"32","author":"Walle","year":"2004","journal-title":"Drug Metab. Dispos."},{"key":"ref_17","doi-asserted-by":"crossref","unstructured":"Gasulla, F., Del Campo, E.M., Casano, L.M., and Gu\u00e9ra, A. (2021). Advances in Understanding of Desiccation Tolerance of Lichens and Lichen-Forming Algae. Plants, 10.","DOI":"10.3390\/plants10040807"},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"71","DOI":"10.1007\/s10886-008-9584-2","article-title":"Lichen Substances Prevent Lichens from Nutrient Deficiency","volume":"35","author":"Hauck","year":"2009","journal-title":"J. Chem. Ecol."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"1490","DOI":"10.1039\/c3np70064j","article-title":"UV-Protectant Metabolites from Lichens and Their Symbiotic Partners","volume":"30","author":"Nguyen","year":"2013","journal-title":"Nat. Prod. Rep."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"1730","DOI":"10.1039\/C7CS00431A","article-title":"Secondary Metabolism in the Lichen Symbiosis","volume":"47","author":"Calcott","year":"2018","journal-title":"Chem. Soc. Rev."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"801","DOI":"10.1039\/C5NP00134J","article-title":"Dibenzofurans and Derivatives from Lichens and Ascomycetes","volume":"33","author":"Millot","year":"2016","journal-title":"Nat. Prod. Rep."},{"key":"ref_22","doi-asserted-by":"crossref","unstructured":"Ure\u00f1a-Vacas, I., Gonz\u00e1lez-Burgos, E., Divakar, P.K., and G\u00f3mez-Serranillos, M.P. (2023). Lichen Depsides and Tridepsides: Progress in Pharmacological Approaches. JoF, 9.","DOI":"10.3390\/jof9010116"},{"key":"ref_23","doi-asserted-by":"crossref","unstructured":"Le Pogam, P., and Boustie, J. (2016). Xanthones of Lichen Source: A 2016 Update. Molecules, 21.","DOI":"10.3390\/molecules21030294"},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"294","DOI":"10.1038\/s41597-019-0305-1","article-title":"A Database of High-Resolution MS\/MS Spectra for Lichen Metabolites","volume":"6","author":"Rondeau","year":"2019","journal-title":"Sci. Data"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"157","DOI":"10.1515\/znc-2010-3-401","article-title":"Current Results on Biological Activities of Lichen Secondary Metabolites: A Review","volume":"65","author":"Farkas","year":"2010","journal-title":"Z. Naturforschung C"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"137","DOI":"10.1007\/s00114-002-0305-3","article-title":"A Review on Usnic Acid, an Interesting Natural Compound","volume":"89","author":"Cocchietto","year":"2002","journal-title":"Naturwissenschaften"},{"key":"ref_27","doi-asserted-by":"crossref","unstructured":"Kumari, M., Kamat, S., Singh, S.K., Kumar, A., and Jayabaskaran, C. (2023). Inhibition of Autophagy Increases Cell Death in HeLa Cells through Usnic Acid Isolated from Lichens. Plants, 12.","DOI":"10.3390\/plants12030519"},{"key":"ref_28","doi-asserted-by":"crossref","unstructured":"Bolikhova, A.K., Alferova, V.A., Izzi, A.R., Nikandrova, A.A., Kudryakova, G.K., Zhitlov, M.Y., Guseva, E.A., Karpov, D.S., Lukianov, D.A., and Dontsova, O.A. (2026). HR-LCMS\/MS-Based Dereplication of Plant-Derived Autophagy Inducers Revealed Astragalus Dasyanthus as a New Glabrol Producer. Metabolites, 16.","DOI":"10.3390\/metabo16050311"},{"key":"ref_29","unstructured":"Muchnik, E.E., Insarova, I.D., and Kazakova, M.V. (2011). Educational Key to Lichens of Central Russia: A Textbook, Ryazan State University named for S.A. Yesenin."},{"key":"ref_30","unstructured":"Golubkova, N.S., Dombrovskaja, A.V., Zhurbenko, M.P., Kotlov, Y.V., and Krusanova, Z.G. (1996). The Handbook of the Lichens of Russia, Russian Academy of Sciences."},{"key":"ref_31","unstructured":"Abramov, I.I. (1978). Handbook of the Lichens of the USSR, Nauka."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"163","DOI":"10.1159\/000328516","article-title":"Resveratrol-Activated AMPK\/SIRT1\/Autophagy in Cellular Models of Parkinson\u2019s Disease","volume":"19","author":"Wu","year":"2011","journal-title":"Neurosignals"},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"1625","DOI":"10.3390\/ijms15011625","article-title":"Resveratrol Partially Prevents Rotenone-Induced Neurotoxicity in Dopaminergic SH-SY5Y Cells through Induction of Heme Oxygenase-1 Dependent Autophagy","volume":"15","author":"Lin","year":"2014","journal-title":"Int. J. Mol. Sci."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"5720","DOI":"10.1093\/emboj\/19.21.5720","article-title":"LC3, a Mammalian Homologue of Yeast Apg8p, Is Localized in Autophagosome Membranes after Processing","volume":"19","author":"Kabeya","year":"2000","journal-title":"EMBO J."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1080\/15548627.2020.1797280","article-title":"Guidelines for the Use and Interpretation of Assays for Monitoring Autophagy (4th Edition)1","volume":"17","author":"Klionsky","year":"2021","journal-title":"Autophagy"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"265","DOI":"10.1134\/S1062359015030036","article-title":"Distribution of Mycotoxins and Usnic Acid in the Thalli of Epigenous Lichens","volume":"42","author":"Kononenko","year":"2015","journal-title":"Biol. Bull."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"556","DOI":"10.12659\/MSM.908568","article-title":"Usnic Acid Induces Cycle Arrest, Apoptosis, and Autophagy in Gastric Cancer Cells In Vitro and In Vivo","volume":"24","author":"Geng","year":"2018","journal-title":"Med. Sci. Monit."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"472","DOI":"10.1002\/1097-458X(200006)38:6<472::AID-MRC658>3.0.CO;2-P","article-title":"NMR Spectral Data of Salazinic Acid Isolated from Some Species of Parmotrema","volume":"38","author":"Sperry","year":"2000","journal-title":"Magn. Reson. Chem."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"8023","DOI":"10.1074\/jbc.M900301200","article-title":"An ATP-Competitive Mammalian Target of Rapamycin Inhibitor Reveals Rapamycin-Resistant Functions of mTORC1","volume":"284","author":"Thoreen","year":"2009","journal-title":"J. Biol. Chem."},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"5","DOI":"10.1172\/JCI73937","article-title":"Pharmacologic Agents Targeting Autophagy","volume":"125","author":"Xia","year":"2015","journal-title":"J. Clin. Investig."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"283","DOI":"10.1042\/BJ20101894","article-title":"The Serine\/Threonine Kinase ULK1 Is a Target of Multiple Phosphorylation Events","volume":"440","author":"Bach","year":"2011","journal-title":"Biochem. J."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"577","DOI":"10.1016\/S0092-8674(03)00929-2","article-title":"TSC2 Mediates Cellular Energy Response to Control Cell Growth and Survival","volume":"115","author":"Inoki","year":"2003","journal-title":"Cell"},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1016\/j.tem.2019.09.005","article-title":"Growth Factor-Dependent and -Independent Activation of mTORC2","volume":"31","author":"Knudsen","year":"2020","journal-title":"Trends Endocrinol. Metab."},{"key":"ref_44","doi-asserted-by":"crossref","unstructured":"Bessadottir, M., Egilsson, M., Einarsdottir, E., Magnusdottir, I.H., Ogmundsdottir, M.H., Omarsdottir, S., and Ogmundsdottir, H.M. (2012). Proton-Shuttling Lichen Compound Usnic Acid Affects Mitochondrial and Lysosomal Function in Cancer Cells. PLoS ONE, 7.","DOI":"10.1371\/journal.pone.0051296"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"855","DOI":"10.1055\/a-1482-6381","article-title":"Lichen Depsidones with Biological Interest","volume":"88","author":"Divakar","year":"2022","journal-title":"Planta Med."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"e15394","DOI":"10.7717\/peerj.15394","article-title":"Recent Advances on Natural Depsidones: Sources, Biosynthesis, Structure-Activity Relationship, and Bioactivities","volume":"11","author":"Khayat","year":"2023","journal-title":"PeerJ"},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"1166","DOI":"10.1016\/j.phymed.2012.07.012","article-title":"Chemical Composition of Three Parmelia Lichens and Antioxidant, Antimicrobial and Cytotoxic Activities of Some Their Major Metabolites","volume":"19","year":"2012","journal-title":"Phytomedicine"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"619","DOI":"10.1515\/znc-2007-7-827","article-title":"Antimicrobial Activity of Extracts of the Lichen Parmelia Sulcata and Its Salazinic Acid Constituent","volume":"62","author":"Candan","year":"2007","journal-title":"Z. Naturforschung C"},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"12991","DOI":"10.1039\/D3RA01542D","article-title":"Salazinic Acid Attenuates Male Sexual Dysfunction and Testicular Oxidative Damage in Streptozotocin-Induced Diabetic Albino Rats","volume":"13","author":"Killari","year":"2023","journal-title":"RSC Adv."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"F1177","DOI":"10.1152\/ajprenal.00034.2011","article-title":"AMPK Protects Proximal Tubular Cells from Stress-Induced Apoptosis by an ATP-Independent Mechanism: Potential Role of Akt Activation","volume":"301","author":"Lieberthal","year":"2011","journal-title":"Am. J. Physiol.-Ren. Physiol."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"365","DOI":"10.2307\/1218225","article-title":"Chemistry of Lichen Substances","volume":"21","author":"Asahina","year":"1972","journal-title":"Taxon"},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"108588","DOI":"10.1016\/j.bioorg.2025.108588","article-title":"AMPK-Specific Autophagy Activator Based on 1,3-Diaza-2-Oxophenoxazine","volume":"162","author":"Guseva","year":"2025","journal-title":"Bioorganic Chem."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"129980","DOI":"10.1016\/j.bmcl.2024.129980","article-title":"Modified (2\u2032-Deoxy)Adenosines Activate Autophagy Primarily through AMPK\/ULK1-Dependent Pathway","volume":"113","author":"Guseva","year":"2024","journal-title":"Bioorganic Med. Chem. Lett."},{"key":"ref_54","doi-asserted-by":"crossref","unstructured":"Guseva, E.A., Kamzeeva, P.N., Sokolskaya, S.Y., Myasnikov, B.P., Golubeva, J.A., Alferova, V.A., Zaranaina, R.-L., Vays, V.B., Vangeli, I.M., and Belyaev, E.S. (2026). Cyanine Dye Conjugates of a 2\u2032-Deoxycytidine-Based Auto- and Mitophagy Activator Extend Caenorhabditis Elegan s Lifespan. bioRxiv.","DOI":"10.64898\/2026.01.23.701232"},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"74","DOI":"10.1016\/j.isci.2018.09.012","article-title":"A Potent and Selective ULK1 Inhibitor Suppresses Autophagy and Sensitizes Cancer Cells to Nutrient Stress","volume":"8","author":"Martin","year":"2018","journal-title":"iScience"},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"447","DOI":"10.1038\/s41587-023-01690-2","article-title":"Integrative Analysis of Multimodal Mass Spectrometry Data in MZmine 3","volume":"41","author":"Schmid","year":"2023","journal-title":"Nat. Biotechnol."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"3760","DOI":"10.1021\/pr900171m","article-title":"TOPPView: An Open-Source Viewer for Mass Spectrometry Data","volume":"8","author":"Sturm","year":"2009","journal-title":"J. Proteome Res."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"101154","DOI":"10.1016\/j.xpro.2022.101154","article-title":"Analyzing Autophagosomes and Mitophagosomes in the Mouse Brain Using Electron Microscopy","volume":"3","author":"Duan","year":"2022","journal-title":"STAR. Protoc."}],"container-title":["International Journal of Molecular Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1422-0067\/27\/19\/8621\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,30]],"date-time":"2026-09-30T04:31:15Z","timestamp":1790742675000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1422-0067\/27\/19\/8621"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,26]]},"references-count":58,"journal-issue":{"issue":"19","published-online":{"date-parts":[[2026,10]]}},"alternative-id":["ijms27198621"],"URL":"https:\/\/doi.org\/10.3390\/ijms27198621","relation":{},"ISSN":["1422-0067"],"issn-type":[{"value":"1422-0067","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,26]]}}}