{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,2]],"date-time":"2026-10-02T12:58:42Z","timestamp":1790945922924,"version":"4.1.0"},"reference-count":74,"publisher":"American Chemical Society (ACS)","issue":"10","license":[{"start":{"date-parts":[[2026,8,31]],"date-time":"2026-08-31T00:00:00Z","timestamp":1788134400000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-029"},{"start":{"date-parts":[[2026,8,31]],"date-time":"2026-08-31T00:00:00Z","timestamp":1788134400000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-037"},{"start":{"date-parts":[[2026,8,31]],"date-time":"2026-08-31T00:00:00Z","timestamp":1788134400000},"content-version":"stm-asf","delay-in-days":0,"URL":"https:\/\/doi.org\/10.15223\/policy-045"}],"funder":[{"DOI":"10.13039\/501100002347","name":"Bundesministerium f?r Forschung, Technologie und Raumfahrt","doi-asserted-by":"publisher","award":["01KI2024"],"award-info":[{"award-number":["01KI2024"]}],"id":[{"id":"10.13039\/501100002347","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001659","name":"Deutsche Forschungsgemeinschaft","doi-asserted-by":"publisher","award":["505696476"],"award-info":[{"award-number":["505696476"]}],"id":[{"id":"10.13039\/501100001659","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001659","name":"Deutsche Forschungsgemeinschaft","doi-asserted-by":"publisher","award":["540833593"],"award-info":[{"award-number":["540833593"]}],"id":[{"id":"10.13039\/501100001659","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001659","name":"Deutsche Forschungsgemeinschaft","doi-asserted-by":"publisher","award":["545040837"],"award-info":[{"award-number":["545040837"]}],"id":[{"id":"10.13039\/501100001659","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2026,10,2]]},"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Ontogenetic shifts in diet are well documented in snakes and are increasingly linked to age-related venom variation. The common adder, Vipera berus, exhibits a dietary transition from predominantly ectothermic prey in its early life to increasingly incorporating endothermic prey as an adult. Here, we investigate whether this dietary shift is reflected in age-related changes in the venom composition and bioactivity of V. berus. Venoms from captive-bred V. berus from Germany were obtained and pooled across five age groups, from neonates to adults. Venom profiles were assessed by SDS-PAGE and genome-guided shotgun proteomics, with quantification based on normalized spectral abundance factors (NSAF) using a toxin-gene catalogue generated from a novel V. berus genome assembly. In parallel, we assayed general protease and PLA2 activities, as well as FXa-, thrombin-, and plasmin-like activities, and cytotoxicity toward mammalian cell lines. We identified two distinct age-related venom phenotypes (ontotypes): an svMP\/CTL-rich ontotype A (\u22641 year) and an svSP\/PLA2-rich ontotype B (\u22652 years). Functionally, protease activity decreased with age, whereas thrombin-like, plasmin-like and PLA2 activities, and cytotoxicity, increased. Our findings indicate an ontogenetic shift in composition and activities of V. berus venom that parallels dietary transitions and potentially reflect adaptation to differing prey physiologies.<\/jats:p>","DOI":"10.1021\/acs.jproteome.6c00042","type":"journal-article","created":{"date-parts":[[2026,9,3]],"date-time":"2026-09-03T04:27:50Z","timestamp":1788409670000},"page":"5103-5113","source":"Crossref","is-referenced-by-count":0,"title":["Ontogenetic\nVariation in Composition and Bioactivity\nof Common Adder (\n                    <i>Vipera berus<\/i>\n                    ) Venom Revealed by\nGenome-Guided Proteomics and In Vitro Functional Assays"],"prefix":"10.1021","volume":"25","author":[{"ORCID":"https:\/\/orcid.org\/0009-0008-5260-9797","authenticated-orcid":true,"given":"Lennart","family":"Schulte","sequence":"first","affiliation":[{"name":"Fraunhofer Institute for Molecular Biology and Applied Ecology (IME) , , Ohlebergsweg 12 , ,","place":["Giessen, Germany, 35392"]},{"name":"Justus Liebig University Giessen , , Heinrich-Buff-Ring 58 , ,","place":["Giessen, Germany, 35392"]},{"name":"LOEWE-Centre for Translational Biodiversity Genomics , Senckenberganlage 25 , ,","place":["Frankfurt, Germany, 60325"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Johanna","family":"Eichberg","sequence":"additional","affiliation":[{"name":"Fraunhofer Institute for Molecular Biology and Applied Ecology (IME) , , Ohlebergsweg 12 , ,","place":["Giessen, Germany, 35392"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alfredo","family":"Cabrera-Orefice","sequence":"additional","affiliation":[{"name":"Justus Liebig University Giessen , , Friedrichstrasse 24 , ,","place":["Giessen, Germany, 35392"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Harry","family":"Wo\u0308lfel","sequence":"additional","affiliation":[{"name":"Venture for Interconnection, Protection, Education and Research in Adders e.V. (VIPERA) , Brunnenplatz 9 , ,","place":["Velburg, Germany, 92355"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Paul","family":"Hien","sequence":"additional","affiliation":[{"name":"Venture for Interconnection, Protection, Education and Research in Adders e.V. (VIPERA) , Brunnenplatz 9 , ,","place":["Velburg, Germany, 92355"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0810-3699","authenticated-orcid":true,"given":"Maik","family":"Damm","sequence":"additional","affiliation":[{"name":"Fraunhofer Institute for Molecular Biology and Applied Ecology (IME) , , Ohlebergsweg 12 , ,","place":["Giessen, Germany, 35392"]},{"name":"Justus Liebig University Giessen , , Heinrich-Buff-Ring 58 , ,","place":["Giessen, Germany, 35392"]},{"name":"LOEWE-Centre for Translational Biodiversity Genomics , Senckenberganlage 25 , ,","place":["Frankfurt, Germany, 60325"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6431-6018","authenticated-orcid":true,"given":"Tilman","family":"Schell","sequence":"additional","affiliation":[{"name":"Fraunhofer Institute for Molecular Biology and Applied Ecology (IME) , , Ohlebergsweg 12 , ,","place":["Giessen, Germany, 35392"]},{"name":"Senckenberg Research Institute and Natural History Museum Frankfurt , Senckenberganlage 25 , ,","place":["Frankfurt, Germany, 60325"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Charlotte","family":"Gerheim","sequence":"additional","affiliation":[{"name":"Fraunhofer Institute for Molecular Biology and Applied Ecology (IME) , , Ohlebergsweg 12 , ,","place":["Giessen, Germany, 35392"]},{"name":"Senckenberg Research Institute and Natural History Museum Frankfurt , Senckenberganlage 25 , ,","place":["Frankfurt, Germany, 60325"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Alexander Ben","family":"Hamadou","sequence":"additional","affiliation":[{"name":"Fraunhofer Institute for Molecular Biology and Applied Ecology (IME) , , Ohlebergsweg 12 , ,","place":["Giessen, Germany, 35392"]},{"name":"Senckenberg Research Institute and Natural History Museum Frankfurt , Senckenberganlage 25 , ,","place":["Frankfurt, Germany, 60325"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sabine","family":"Hurka","sequence":"additional","affiliation":[{"name":"Fraunhofer Institute for Molecular Biology and Applied Ecology (IME) , , Ohlebergsweg 12 , ,","place":["Giessen, Germany, 35392"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Thomas","family":"Lindner","sequence":"additional","affiliation":[{"name":"University of Regensburg , , Universita\u0308tsstrasse 31 , ,","place":["Regensburg, Germany, 93053"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Gregor","family":"Geisler","sequence":"additional","affiliation":[{"name":"Laboklin Laboratory for Clinical Diagnostics GmbH & Co , Steubenstrasse 4 , ,","place":["Bad Kissingen, Germany, 97688"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ignazio","family":"Avella","sequence":"additional","affiliation":[{"name":"Fraunhofer Institute for Molecular Biology and Applied Ecology (IME) , , Ohlebergsweg 12 , ,","place":["Giessen, Germany, 35392"]},{"name":"Justus Liebig University Giessen , , Heinrich-Buff-Ring 58 , ,","place":["Giessen, Germany, 35392"]},{"name":"LOEWE-Centre for Translational Biodiversity Genomics , Senckenberganlage 25 , ,","place":["Frankfurt, Germany, 60325"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Kornelia","family":"Hardes","sequence":"additional","affiliation":[{"name":"Fraunhofer Institute for Molecular Biology and Applied Ecology (IME) , , Ohlebergsweg 12 , ,","place":["Giessen, Germany, 35392"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Carola","family":"Greve","sequence":"additional","affiliation":[{"name":"Fraunhofer Institute for Molecular Biology and Applied Ecology (IME) , , Ohlebergsweg 12 , ,","place":["Giessen, Germany, 35392"]},{"name":"Senckenberg Research Institute and Natural History Museum Frankfurt , Senckenberganlage 25 , ,","place":["Frankfurt, Germany, 60325"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tim","family":"Lu\u0308ddecke","sequence":"additional","affiliation":[{"name":"Fraunhofer Institute for Molecular Biology and Applied Ecology (IME) , , Ohlebergsweg 12 , ,","place":["Giessen, Germany, 35392"]},{"name":"LOEWE-Centre for Translational Biodiversity Genomics , Senckenberganlage 25 , ,","place":["Frankfurt, Germany, 60325"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"316","published-online":{"date-parts":[[2026,8,31]]},"reference":[{"key":"2026100208412718700_cit1","volume-title":"Venom: The Secrets of\nNature\u2019s Deadliest Weapon","author":"Jenner","year":"2017"},{"issue":"11","key":"2026100208412718700_cit2","doi-asserted-by":"publisher","first-page":"666","DOI":"10.3390\/toxins11110666","article-title":"The\nDiversity\nof Venom: The Importance of Behavior and Venom System Morphology in\nUnderstanding Its Ecology and 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Govern Venom Variation\nin a Widespread Rattlesnake Species","volume":"286","author":"Zancolli","year":"2019","journal-title":"Proceedings\nof the Royal Society B: Biological Sciences"},{"issue":"8","key":"2026100208412718700_cit9","doi-asserted-by":"publisher","first-page":"570","DOI":"10.1016\/j.tips.2020.05.006","article-title":"Causes and Consequences\nof Snake Venom Variation","volume":"41","author":"Casewell","year":"2020","journal-title":"Trends Pharmacol.\nSci."},{"issue":"1","key":"2026100208412718700_cit10","doi-asserted-by":"publisher","first-page":"67","DOI":"10.1016\/0041-0101(95)00122-0","article-title":"Electrophoretic Profiles and Biological\nActivities:\nIntraspecific Variation in the Venom of the Malayan Pit Viper (Calloselasma Rhodostoma)","volume":"34","author":"Daltry","year":"1996","journal-title":"Toxicon"},{"key":"2026100208412718700_cit11","doi-asserted-by":"publisher","first-page":"74","DOI":"10.1016\/j.toxicon.2015.01.010","article-title":"Early 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Ammodytes","volume":"310","author":"Lakus\u030cic\u0301","year":"2025","journal-title":"Journal of Proteomics"},{"key":"2026100208412718700_cit15","doi-asserted-by":"publisher","DOI":"10.1016\/j.toxicon.2025.108291","article-title":"Beyond Sexual Maturity:\nImportance of Dietary Changes\nin Venom Variation in Vipera Ammodytes","volume":"257","author":"Lakus\u030cic\u0301","year":"2025","journal-title":"Toxicon"},{"key":"2026100208412718700_cit16","doi-asserted-by":"publisher","DOI":"10.1016\/j.jprot.2022.104613","article-title":"Interpopulational Variation and Ontogenetic\nShift in the Venom Composition of Lataste\u2019s Viper (Vipera Latastei, Bosca\u0301 1878) from Northern Portugal","volume":"263","author":"Avella","year":"2022","journal-title":"Journal of Proteomics"},{"key":"2026100208412718700_cit17","volume-title":"True Vipers: Natural\nHistory and Toxinology of Old World Vipers","author":"Mallow","year":"2003"},{"key":"2026100208412718700_cit18","volume-title":"Venomous Snakes of Europe, Northern,\nCentral and Western Asia","author":"David","year":"2006"},{"key":"2026100208412718700_cit19","volume-title":"Snakes of Europe, North\nAfrica and the Middle East - A Photographic Guide: A Photographic\nGuide","author":"Geniez","year":"2018"},{"key":"2026100208412718700_cit20","doi-asserted-by":"crossref","unstructured":"Luiselli, L.; Anibaldi, C.; Capula, M.\n          Diet of Juvenile Adders,\nVipera Berus, in an Alpine Habitat. 1995.10.1163\/156853895X00488.","DOI":"10.1163\/156853895X00488"},{"key":"2026100208412718700_cit21","volume-title":"Die Kreuzotter - Ein Leben in ziemlich\nfesten Bahnen","author":"Otte","year":"2020"},{"issue":"1","key":"2026100208412718700_cit22","doi-asserted-by":"publisher","first-page":"11","DOI":"10.1007\/s10344-022-01559-w","article-title":"A Hidden Threat\nin Abandoned Fields: Frequent Nest Predation by Common Adder on Ground-Nesting\nPasserines","volume":"68","author":"Samsonov","year":"2022","journal-title":"Eur. J. Wildl Res."},{"issue":"16","key":"2026100208412718700_cit23","doi-asserted-by":"publisher","first-page":"447","DOI":"10.3238\/arztebl.m2025.0091","article-title":"Kreuzotter-Bisse in Deutschland Zwischen\n2012 Und 2023","volume":"122","author":"Rae","year":"2025","journal-title":"Deutsches A\u0308rzteblatt"},{"issue":"5","key":"2026100208412718700_cit24","doi-asserted-by":"publisher","first-page":"362","DOI":"10.1097\/MEJ.0000000000000577","article-title":"Envenomation by the\nCommon European Adder (Vipera Berus): A Case Series\nof 219 Patients","volume":"26","author":"Hermansen","year":"2019","journal-title":"European Journal of Emergency\nMedicine"},{"issue":"10","key":"2026100208412718700_cit25","doi-asserted-by":"publisher","first-page":"2247","DOI":"10.1111\/jdv.16722","article-title":"Vipera Snakebite\nin Europe: A Systematic Review of a Neglected Disease","volume":"34","author":"Paolino","year":"2020","journal-title":"Journal of the European Academy of Dermatology and Venereology"},{"key":"2026100208412718700_cit26","doi-asserted-by":"publisher","DOI":"10.1016\/j.tox.2021.152724","article-title":"Vipers of Major Clinical Relevance in Europe: Taxonomy,\nVenom Composition, Toxicology and Clinical Management of Human Bites","volume":"453","author":"Di Nicola","year":"2021","journal-title":"Toxicology"},{"key":"2026100208412718700_cit27","doi-asserted-by":"publisher","first-page":"34","DOI":"10.1016\/j.jprot.2016.06.020","article-title":"Venomics of Vipera Berus Berus to Explain Differences in Pathology Elicited by Vipera Ammodytes Ammodytes Envenomation: Therapeutic Implications","volume":"146","author":"Latinovic\u0301","year":"2016","journal-title":"Journal of Proteomics"},{"issue":"10","key":"2026100208412718700_cit28","doi-asserted-by":"publisher","first-page":"1398","DOI":"10.3390\/molecules21101398","article-title":"Proteome and Peptidome of Vipera Berus Berus Venom","volume":"21","author":"Bocian","year":"2016","journal-title":"Molecules"},{"issue":"2","key":"2026100208412718700_cit29","doi-asserted-by":"publisher","first-page":"90","DOI":"10.3390\/toxins11020090","article-title":"Vipera Berus Berus Venom from Russia: Venomics, Bioactivities and\nPreclinical Assessment\nof Microgen Antivenom","volume":"11","author":"Al-Shekhadat","year":"2019","journal-title":"Toxins"},{"issue":"8","key":"2026100208412718700_cit30","doi-asserted-by":"publisher","first-page":"3524","DOI":"10.1021\/acs.jproteome.4c00171","article-title":"Venomics and Peptidomics of Palearctic\nVipers: A Clade-Wide Analysis of Seven Taxa of the Genera Vipera, Montivipera, Macrovipera, and Daboia across Tu\u0308rkiye","volume":"23","author":"Damm","year":"2024","journal-title":"J. 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Conda Package and Environment\nManager, 2017. https:\/\/docs.conda.io\/."},{"issue":"7","key":"2026100208412718700_cit40","doi-asserted-by":"publisher","first-page":"1023","DOI":"10.1038\/s41587-021-01156-3","article-title":"SignalP 6.0 Predicts All Five Types\nof Signal Peptides Using Protein Language Models","volume":"40","author":"Teufel","year":"2022","journal-title":"Nat. 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