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Møller P, Azqueta A, Boutet-Robinet E, Koppen G, Bonassi S, Milié M, Gajski G, Costa S, Teixeira JP, Costa Pereira C, Dusinska M, Godschalk R, Brunborg G, Gutzkow KB, Giovannelli L, Cooke MS, Richling E, Laffon B, Valdiglesias V, Langie SAS (2020) Minimum Information for Reporting on the Comet Assay (MIRCA): recommendations for describing comet assay procedures and results. Nat Protoc. https://doi.org/10.1038/s41596-020-0398-1

Nübler S, Esteban López M, Castaño A, Mol HGJ, Müller J, Schäfer M, Haji-Abbas-Zarrabi K, Hajslova J, Pulkrabova J, Dvorakova D, Urbancova K, Koch HM, Antignac JP, Sakhi AK, Vorkamp K, Burkhardt T, Scherer M, Göen T (2023) External Quality Assurance Schemes (EQUASs) and Inter-laboratory Comparison Investigations (ICIs) for human biomonitoring of polycyclic aromatic hydrocarbon (PAH) biomarkers in urine as part of the quality assurance programme under HBM4EU. Int J Hyg Environment Health 250:114169. https://doi.org/10.1016/j.ijheh.2023.114169

OECD (2020) Test No. 471: Bacterial Reverse Mutation Test, OECD Guidelines for the Testing of Chemicals, Section 4. OECD Publishing, Paris. https://doi.org/10.1787/9789264071247-en

Olsen M, Visconti A (1988) Metabolism of alternariol monomethyl ether by porcine liver and intestinal mucosa *in vitro*. Toxicol in Vitro 2(1):27-29. https://doi.org/10.1016/0887-2333(88)90033-1

Osborne LC, Jones VI, Peeler JT, Larkin EP (1988) Transformation of C3H/10T12 cells and induction of EBV-early antigen in Raji cells by altertoxins I and III. Toxicol in Vitro 2(2):97-102. https://doi.org/10.1016/0887-2333(88)90019-7

Pahlke G, Tiessen C, Domnanich K, Kahle N, Groh IAM, Schreck I, Weiss C, Marko D (2016) Impact of *Alternaria* toxins on CYP1A1 expression in different human tumor cells and relevance for genotoxicity. Toxicol Lett 240(1):93-104. https://doi.org/10.1016/j.toxlet.2015.10.003

Perrin L, Loiseau N, André F, Delaforge M (2011) Metabolism of N-methyl-amide by cytochrome P450s. FEBS J 278(12):2167-2178. https://doi.org/10.1111/j.1742-4658.2011.08133.x

Perrone G, Ferrara M, Medina A, Pascale M, Magan N (2020) Toxigenic fungi and mycotoxins in a climate change scenario: ecology, genomics, distribution, prediction and prevention of the risk. Microorganisms 8(10):1496. https://doi.org/10.3390/MICROORGANISMS8101496

Pfeiffer E, Eschbach S, Metzler M (2007) *Alternaria* toxins: DNA strand-breaking activity in mammalian cells *in vitro*. Mycotoxin Res 23(3):152-157. https://doi.org/10.1007/BF02951512

Pfeiffer E, Burkhardt B, Altemöller M, Podlech J, Metzler M (2008) Activities of human recombinant cytochrome P450 isoforms and human hepatic microsomes for the hydroxylation of *Alternaria* toxins. Mycotoxin Res 24(3):117-123. https://doi.org/10.1007/BF03032337

Pfeiffer E, Herrmann C, Altemöller M, Podlech J, Metzler M (2009a) Oxidative in vitro metabolism of the Alternaria toxins altenuene and isoaltenuene. Mol Nutr Food Res 53:452-459. https://doi.org/10.1002/mnfr.200700501

Pfeiffer E, Schmit C, Burkhardt B, Altemöller M, Podlech J, Metzler M (2009b) Glucuronidation of the mycotoxins alternariol

and alternariol-9-methyl ether *in vitro*: Chemical structures of glucuronides and activities of human UDP-glucuronosyltransferase isoforms. Mycotoxin Res 25(1):3-10. https://doi.org/10.1007/s12550-008-0001-z

Pollock GA, DiSabatino CE, Heimsch RC, Coulombe RA (1982) The distribution, elimination, and metabolism of ¹⁴C-alternariol monomethyl ether. J Environ Sci Health B 17(2):109-124. https://doi.org/10.1080/03601238209372306

Puntscher H, Kütt ML, Skrinjar P, Mikula H, Podlech J, Fröhlich J, Marko D, Warth B (2018) Tracking emerging mycotoxins in food: development of an LC-MS/MS method for free and modified *Alternaria* toxins. Anal Bioanal Chem 410(18):4481-4494. https://doi.org/10.1007/S00216-018-1105-8

Puntscher H, Aichinger G, Grabher S, Attakpah E, Krüger F, Tillmann K, Motschnig T, Hohenbichler J, Braun D, Plasenzotti R, Pahlke G, Höger H, Marko D, Warth B (2019a) Bioavailability, metabolism, and excretion of a complex Alternaria culture extract versus altertoxin II: a comparative study in rats. Arch Toxicol 93(11):3153-3167. https://doi.org/10.1007/s00204-019-02575-7

Puntscher H, Hankele S, Tillmann K, Attakpah E, Braun D, Kütt ML, Del Favero G, Aichinger G, Pahlke G, Höger H, Marko D, Warth B (2019b) First insights into Alternaria multi-toxin *in vivo* metabolism. Toxicol Lett 301:168-178. https://doi.org/10.1016/J.TOXLET.2018.10.006

Qiao X, Li G, Zhang J, Du J, Yang Y, Yin J, Li H, Xie J, Jiang Y, Fang X, Dai X, Shao B (2022) Urinary analysis reveals high *Alternaria* mycotoxins exposure in the general population from Beijing, China. J Environ Sci 118:122-129. https://doi.org/10.1016/J.JES.2021.08.025

Samak MA, Elshatory A, Mohamed EM (2019) Outcomes of gallic acid on alternariol induced cyto-morphic and genotoxic *In Vivo* changes in parotid gland: 4-HNE incorporated. Biomedicines 7(4):84. https://doi.org/10.3390/BIOMEDICINES7040084

Schmutz C, Cenk E, Marko D (2019) The *Alternaria* mycotoxin alternariol triggers the immune response of IL-1β-stimulated, differentiated Caco-2 Cells. Mol Nutr Food Res 63(20):1-8. https://doi.org/10.1002/mnfr.201900341

Schrader TJ, Cherry W, Soper K, Langlois I, Vijay HM (2001) Examination of *Alternaria alternata* mutagenicity and effects of nitrosylation using the ames *Salmonella* test. Teratog Carcinog Mutagen 21(4):261-274. https://doi.org/10.1002/tcm.1014

Schrader TJ, Cherry W, Soper K, Langlois I (2006) Further examination of the effects of nitrosylation on *Alternaria alternata* mycotoxin mutagenicity in *vitro*. Mutat Res 606(1-2):61-71. https://doi.org/10.1016/j.MRGENTOX.2006.02.008

Schuchardt S, Ziemann C, Hansen T (2014) Combined toxicokinetic and *in vivo* genotoxicity study on *Alternaria* toxins. EFSA Support Publ 11(11):679E. https://doi.org/10.2903/SP.EFSA.2014.EN-679

Schwarz C, Tiessen C, Kreutzer M, Stark T, Hofmann T, Marko D (2012) Characterization of a genotoxic impact compound in *Alternaria alternata* infested rice as altertoxin II. Arch Toxicol 86(12):1911-1925. https://doi.org/10.1007/s00204-012-0958-4

Solhaug A, Vines LL, Ivanova L, Spilsberg B, Holme JA, Pestkab J, Collins A, Eriksen GS (2012) Mechanisms involved in alternariol-induced cell cycle arrest. Mutat Res Fund Mol Mech Mutagen 738-739(1):1-11. https://doi.org/10.1016/j.mrfmmm.2012.09.001

Solhaug A, Holme JA, Haglund K, Dendele B, Sergent O, Pestka J, Lagadic-Gossmann D, Eriksen GS (2013) Alternariol induces abnormal nuclear morphology and cell cycle arrest in murine RAW 264.7 macrophages. Toxicol Lett 219(1):8-17. https://doi.org/10.1016/j.toxlet.2013.02.012

Solhaug A, Torgersen ML, Holme JA, Lagadic-Gossmann D, Eriksen GS (2014) Autophagy and senescence, stress responses induced
Structure determination and cytotoxic activities. Biotechnol Rep 33:e00628. https://doi.org/10.1016/j.btre.2021.e00628

Mayer S, Twaruzek M, Blajet-Kosicka A, Grajewski J (2016) Occupational exposure to mould and microbial metabolites during onion sorting – insights into oan overlooked workplace. Environ Monit Assess 188:154. https://doi.org/10.1007/s10661-016-5150-5

Miao Y, Wang D, Chen Y, Zhu X, Tang X, Zhang J, Zhang L, Chen J (2022) General toxicity and genotoxicity of alternariol: a novel 28-day multi-endpoint assessment in male Sprague-Dawley rats. Mycotoxin Res 38(4):231–241. https://doi.org/10.1007/s12550-022-00466-y

Møller P, Azqueta A, Boutet-Robinet E, Koppen G, Bonassi S, Milić M, Gajski G, Costa S, Teixeira JP, Costa Pereira C, Dusinska M, Godschalk R, Brunborg G, Gutzkow KB, Giovannelli L, Cooke MS, Richling E, Laffon B, Valdiglesias V, Langie SAS (2020) Minimum Information for Reporting on the Comet Assay (MIRCA): recommendations for describing comet assay procedures and results. Nat Protoc. https://doi.org/10.1038/s41596-020-0398-1

Nübler S, Esteban López M, Castaño A, Mol HGJ, Müller J, Schäfer M, Haji-Abbas-Zarrabi K, Hajslova J, Pulkrabova J, Dvorakova D, Urbancova K, Koch HM, Antignac JP, Sakhi AK, Vorkamp K, Burkhardt T, Scherer M, Göen T (2023) External Quality Assurance Schemes (EQUASs) and Inter-laboratory Comparison Investigations (ICIs) for human biomonitoring of polycyclic aromatic hydrocarbon (PAH) biomarkers in urine as part of the quality assurance programme under HBM4EU. Int J Hyg Environment Health 250:114169. https://doi.org/10.1016/j.ijheh.2023.114169

OECD (2020) Test No. 471: Bacterial Reverse Mutation Test, OECD Guidelines for the Testing of Chemicals, Section 4. OECD Publishing, Paris. https://doi.org/10.1787/9789264071247-en

Olsen M, Visconti A (1988) Metabolism of alternariol monomethyl ether by porcine liver and intestinal mucosa *in vitro*. Toxicol in Vitro 2(1):27–29. https://doi.org/10.1016/0887-2333(88)90033-1

Osborne LC, Jones VI, Peeler JT, Larkin EP (1988) Transformation of C3H/10T12 cells and induction of EBV-early antigen in Raji cells by altertoxins I and III. Toxicol in Vitro 2(2):97–102. https://doi.org/10.1016/0887-2333(88)90019-7

Pahlke G, Tiessen C, Domnanich K, Kahle N, Groh IAM, Schreck I, Weiss C, Marko D (2016) Impact of *Alternaria* toxins on CYP1A1 expression in different human tumor cells and relevance for genotoxicity. Toxicol Lett 240(1):93–104. https://doi.org/10.1016/j.toxlet.2015.10.003

Perrin L, Loiseau N, André F, Delaforge M (2011) Metabolism of N-methyl-amide by cytochrome P450s. FEBS J 278(12):2167–2178. https://doi.org/10.1111/j.1742-4658.2011.08133.x

Perrone G, Ferrara M, Medina A, Pascale M, Magan N (2020) Toxigenic fungi and mycotoxins in a climate change scenario: ecology, genomics, distribution, prediction and prevention of the risk. Microorganisms 8(10):1496. https://doi.org/10.3390/MICROORGANISMS8101496

Pfeiffer E, Eschbach S, Metzler M (2007) *Alternaria* toxins: DNA strand-breaking activity in mammalian cells *in vitro*. Mycotoxin Res 23(3):152–157. https://doi.org/10.1007/BF02951512

Pfeiffer E, Burkhardt B, Altemöller M, Podlech J, Metzler M (2008) Activities of human recombinant cytochrome P450 isoforms and human hepatic microsomes for the hydroxylation of *Alternaria* toxins. Mycotoxin Res 24(3):117–123. https://doi.org/10.1007/BF03032337

Pfeiffer E, Herrmann C, Altemöller M, Podlech J, Metzler M (2009a) Oxidative in vitro metabolism of the Alternaria toxins altenuene and isoaltenuene. Mol Nutr Food Res 53:452–459. https://doi.org/10.1002/mnfr.200700501

Pfeiffer E, Schmit C, Burkhardt B, Altemöller M, Podlech J, Metzler M (2009b) Glucuronidation of the mycotoxins alternariol

and alternariol-9-methyl ether *in vitro*: Chemical structures of glucuronides and activities of human UDP-glucuronosyltransferase isoforms. Mycotoxin Res 25(1):3–10. https://doi.org/10.1007/s12550-008-0001-z

Pollock GA, DiSabatino CE, Heimsch RC, Coulombe RA (1982) The distribution, elimination, and metabolism of ¹⁴C-alternariol monomethyl ether. J Environ Sci Health B 17(2):109–124. https://doi.org/10.1080/03601238209372306

Puntscher H, Kütt ML, Skrinjar P, Mikula H, Podlech J, Fröhlich J, Marko D, Warth B (2018) Tracking emerging mycotoxins in food: development of an LC-MS/MS method for free and modified *Alternaria* toxins. Anal Bioanal Chem 410(18):4481–4494. https://doi.org/10.1007/S00216-018-1105-8

Puntscher H, Aichinger G, Grabher S, Attakpah E, Krüger F, Tillmann K, Motschnig T, Hohenbichler J, Braun D, Plasenzotti R, Pahlke G, Höger H, Marko D, Warth B (2019a) Bioavailability, metabolism, and excretion of a complex Alternaria culture extract versus altertoxin II: a comparative study in rats. Arch Toxicol 93(11):3153–3167. https://doi.org/10.1007/s00204-019-02575-7

Puntscher H, Hankele S, Tillmann K, Attakpah E, Braun D, Kütt ML, Del Favero G, Aichinger G, Pahlke G, Höger H, Marko D, Warth B (2019b) First insights into Alternaria multi-toxin *in vivo* metabolism. Toxicol Lett 301:168–178. https://doi.org/10.1016/J.TOXLET.2018.10.006

Qiao X, Li G, Zhang J, Du J, Yang Y, Yin J, Li H, Xie J, Jiang Y, Fang X, Dai X, Shao B (2022) Urinary analysis reveals high *Alternaria* mycotoxins exposure in the general population from Beijing, China. J Environ Sci 118:122–129. https://doi.org/10.1016/J.JES.2021.08.025

Samak MA, Elshatory A, Mohamed EM (2019) Outcomes of gallic acid on alternariol induced cyto-morphic and genotoxic *In Vivo* changes in parotid gland: 4-HNE incorporated. Biomedicines 7(4):84. https://doi.org/10.3390/BIOMEDICINES7040084

Schmutz C, Cenk E, Marko D (2019) The *Alternaria* mycotoxin alternariol triggers the immune response of IL-1β-stimulated, differentiated Caco-2 Cells. Mol Nutr Food Res 63(20):1–8. https://doi.org/10.1002/mnfr.201900341

Schrader TJ, Cherry W, Soper K, Langlois I, Vijay HM (2001) Examination of *Alternaria alternata* mutagenicity and effects of nitrosylation using the Ames *Salmonella* test. Teratog Carcinog Mutagen 21(4):261–274. https://doi.org/10.1002/tcm.1014

Schrader TJ, Cherry W, Soper K, Langlois I (2006) Further examination of the effects of nitrosylation on *Alternaria alternata* mycotoxin mutagenicity in *vitro*. Mutat Res 606(1-2):61–71. https://doi.org/10.1016/j.MRGENTOX.2006.02.008

Schuchardt S, Ziemann C, Hansen T (2014) Combined toxicokinetic and *in vivo* genotoxicity study on *Alternaria* toxins. EFSA Support Publ 11(11):679E. https://doi.org/10.2903/SP.EFSA.2014.EN-679

Schwarz C, Tiessen C, Kreutzer M, Stark T, Hofmann T, Marko D (2012) Characterization of a genotoxic impact compound in *Alternaria alternata* infested rice as altertoxin II. Arch Toxicol 86(12):1911–1925. https://doi.org/10.1007/s00204-012-0958-4

Solhaug A, Vines LL, Ivanova L, Spilsberg B, Holme JA, Pestkab J, Collins A, Eriksen GS (2012) Mechanisms involved in alternariol-induced cell cycle arrest. Mutat Res Fund Mol Mech Mutagen 738–739(1):1–11. https://doi.org/10.1016/j.mrfmmm.2012.09.001

Solhaug A, Holme JA, Haglund K, Dendele B, Sergent O, Pestka J, Lagadic-Gossmann D, Eriksen GS (2013) Alternariol induces abnormal nuclear morphology and cell cycle arrest in murine RAW 264.7 macrophages. Toxicol Lett 219(1):8–17. https://doi.org/10.1016/j.toxlet.2013.02.012

Solhaug A, Torgersen ML, Holme JA, Lagadic-Gossmann D, Eriksen GS (2014) Autophagy and senescence, stress responses induced
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Soukup ST, Fleck SC, Pfeiffer E, Podlech J, Kulling SE, Metzler M (2020) DNA reactivity of altertoxin II: Identification of two cova- lent guanine adducts formed under cell-free conditions. *Toxicol Lett* 331:75–81. https://doi.org/10.1016/j.TOXLET.2020.05.018

Stack ME, Prival MJ (1986) Mutagenicity of the *Alternaria* metabo- lites altertoxins I, II, and III. *Appl Environ Microbiol* 52(4):718. https://doi.org/10.1128/AEM.52.4.718-722.1986

Straumfors A, Uhlig S, Eriksen GS, Heldal KK, Eduard W, Kriska R, Sulyok M (2015) Mycotoxins and other fungal metabolites in grain dust from Norwegian grain elevators and compound feed mills. *World Mycotoxin J* 8(3):361–373. https://doi.org/10.3920/ WMJ2014.1799

Stypuła-Trębas S, Minta M, Radko L, Jedziniak P, Posyniak A (2017) Nonsteroidal mycotoxin alternariol is a full androgen agonist in the yeast reporter androgen bioassay. *Environ Toxicol Pharmacol* 55:208–211. https://doi.org/10.1016/J.ETAP.2017.08.036

Tang X, Chen Y, Zhu X, Miao Y, Wang D, Zhang J, Li R, Zhang L, Chen J (2022) Alternariol monomethyl ether toxicity and geno- toxicity in male Sprague-Dawley rats: 28-Day *in vivo* multi-end- point assessment. *Mutat Res Genetic Toxicol Environ Mutagen* 873:10. https://doi.org/10.1016/J.MRGENTOX.2021.503435

Tiemann U, Tomek W, Schneider F, Müller M, Pöhland R, Vanselow J (2009) The mycotoxins alternariol and alternariol methyl ether negatively affect progesterone synthesis in porcine granulosa cells *in vitro*. *Toxicol Lett* 186(2):139–145. https://doi.org/10. 1016/j.toxlet.2009.01.014

Tiessen C, Fehr M, Schwarz C, Baechler S, Domnanich K, Böttler U, Pahlke G, Marko D (2013a) Modulation of the cellular redox sta- tus by the *Alternaria* toxins alternariol and alternariol monome- thyl ether. *Toxicol Lett* 216(1):23–30. https://doi.org/10.1016/J. TOXLET.2012.11.005

Tiessen C, Gehrke H, Kropat C, Schwarz C, Bächler S, Fehr M, Pahlke G, Marko D (2013b) Role of topoisomerase inhibition and DNA repair mechanisms in the genotoxicity of alternariol and alter- toxin-II. *World Mycotoxin J* 6(3):233–244. https://doi.org/10. 3920/wmj2013.1592

Tiessen C, Ellmer D, Mikula H, Pahlke G, Warth B, Gehrke H, Zim- mermann K, Heiss E, Fröhlich J, Marko D (2017) Impact of phase I metabolism on uptake, oxidative stress and genotoxicity of the emerging mycotoxin alternariol and its monomethyl ether in esophageal cells. *Arch Toxicol* 91(3):1213–1226. https://doi. org/10.1007/s00204-016-1801-0

Tran VN, Viktorova J, Augustynkova K, Jelenova N, Dobiasova S, Rehorova K, Fenclova M, Stranska-Zachariasova M, Vitek L, Hajslova J, Ruml T (2020) *In silico* and *in vitro* studies of myco- toxins and their cocktails; Their toxicity and its mitigation by silibinin pre-treatment. Toxins. https://doi.org/10.3390/toxin s12030148

Vejdovszky K, Sack M, Jarolim K, Aichinger G, Somoza MM, Marko D (2017) *In vitro* combinatory effects of the *Alternaria* mycotox- ins alternariol and altertoxin II and potentially involved miRNAs. *Toxicol Lett* 267:45–52. https://doi.org/10.1016/j.toxlet.2016.12. 011

Viegas S, Assunção R, Nunes C, Osteresch B, Twaružek M, Kosicki R, Grajewski J, Martins C, Alvito P, Almeida A, Viegas C (2018) Exposure assessment to mycotoxins in a Portuguese fresh bread dough company by using a multi-biomarker approach. Toxins 10(9):342. https://doi.org/10.3390/toxins10090342

Vila-Donat P, Fernández-Blanco C, Sagratini G, Font G, Ruiz MJ (2015) Effects of soyasaponin I and soyasaponins-rich extract on the Alternariol-induced cytotoxicity on Caco-2 cells. Food Chem Toxicol 77:44–49. https://doi.org/10.1016/j.fct.2014.12.016

VKM, Steffensen I, Fæste CK, Husøy T, Knutsen HK, Mathisen GH, Ørnsrud R, Agdestein A, Bodin J, Elvevoll E, Hessen DO, Hof- shagen M, Krogdahl A, Nilsen AM, Rafoss T, Skjerdal T, Velle G, Wasteson Y, Hemre G, Vandvik V, Alexander J (2019) Rank- ing of substances for monitoring in foods, drinks and dietary supplements - based on risk and knowledge gaps. Scientific Opinion of the Scientific Steering Committee of the Norwegian Scientific Committee for Food and Environment. VKM report 2019:13, ISBN: 978-82-8259-329-8. https://fhi.brage.unit.no/ fhi-xmlui/bitstream/handle/11250/2627587/Steffensen_2019_ Ran.pdf?sequence=2. Accessed 20 Aug 2021

Wang Y, Liu HX, Chen YC, Sun ZH, Li HH, Li SN, Yan ML, Zhang WM (2017) Two new metabolites from the endophytic fungus *Alternaria* sp. A744 derived from *Morinda officinalis*. *Molecules*. https://doi.org/10.3390/molecules22050765

Wollenhaupt K, Schneider F, Tiemann U (2008) Influence of alternariol (AOH) on regulator proteins of cap-dependent translation in porcine endometrial cells. *Toxicol Lett* 182(1–3):57–62. https://doi. org/10.1016/j.TOXLET.2008.08.005

World Health Organization (WHO) (2009) WHO Guideline for indoor air quality: dampness and mould. WHO Regional Office for Europe, Copenhagen, Denmark. https://www.who.int/publicatio ns/i/item/9789289041683. Accessed 18 Jan 2023

Xiao J, Zhang Q, Gao YQ, Tang JJ, Zhang AL, Gao JM (2014) Second- ary metabolites from the endophytic botryosphaeria dothidea of melia azedarach and their antifungal, antibacterial, antioxidant, and cytotoxic activities. J Agric Food Chem 62(16):3584–3590. https://doi.org/10.1021/jf500054f

Yekeler H, Bitmiş K, Özçelik N, Doymaz MZ, Calta M (2001) Analysis of toxic effects of *Alternaria* toxins on esophagus of mice by light and electron microscopy. *Toxicol Pathol* 29(4):492–497. https://doi.org/10.1080/01926230152499980

Zhao DL, Cao F, Wang CY, Yang LJ, Shi T, Wang KL, Shao CL, Wang CY (2019) Alternatone A, an unusual perylenequinone-related compound from a soft-coral-derived strain of the fungus *Alternaria alternata*. *J Nat Prod* 82(11):3201–3204. https://doi.org/ 10.1021/acs.jnatprod.9b00905

Zhu X, Chen Y, Tang X, Wang D, Miao Y, Zhang J, Li R, Zhang L, Chen J (2022) General toxicity and genotoxicity of altertoxin I: a novel 28-day multiendpoint assessment in male Sprague-Dawley rats. *J Appl Toxicol JAT* 42(8):1310–1322. https://doi.org/10. 1002/JAT.4297

**Publisher's Note** Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
by the DNA-damaging mycotoxin alternariol. Toxicology
326:119-129. https://doi.org/10.1016/j.tox.2014.10.009

Solhaug A, Wisbech C, Christoffersen TE, Hult LO, Lea T, Eriksen GS, Holme JA (2015) The mycotoxin alternariol induces DNA damage and modify macrophage phenotype and inflammatory responses. *Toxicol Lett* 239(1):9–21. https://doi.org/10.1016/j.toxlet.2015.08.1107

Solhaug A, Karlsøen LM, Holme JA, Kristoffersen AB, Eriksen GS (2016b) Immunomodulatory effects of individual and combined mycotoxins in the THP-1 cell line. *Toxicol in Vitro* 36:120–132. https://doi.org/10.1016/j.tiv.2016.07.012

Solhaug A, Eriksen GS, Holme JA (2016) Mechanisms of action and toxicity of the mycotoxin alternariol: a review. https://doi.org/10.1111/bcpt.12635

Soukup ST, Fleck SC, Pfeiffer E, Podlech J, Kulling SE, Metzler M (2020) DNA reactivity of altertoxin II: Identification of two cova- lent guanine adducts formed under cell-free conditions. *Toxicol Lett* 331:75–81. https://doi.org/10.1016/j.TOXLET.2020.05.018

Stack ME, Prival MJ (1986) Mutagenicity of the *Alternaria* metabo- lites altertoxins I, II, and III. *Appl Environ Microbiol* 52(4):718. https://doi.org/10.1128/AEM.52.4.718-722.1986

Straumfors A, Uhlig S, Eriksen GS, Heldal KK, Eduard W, Kriska R, Sulyok M (2015) Mycotoxins and other fungal metabolites in grain dust from Norwegian grain elevators and compound feed mills. *World Mycotoxin J* 8(3):361–373. https://doi.org/10.3920/ WMJ2014.1799

Stypuła-Trębas S, Minta M, Radko L, Jedziniak P, Posyniak A (2017) Nonsteroidal mycotoxin alternariol is a full androgen agonist in the yeast reporter androgen bioassay. *Environ Toxicol Pharmacol* 55:208–211. https://doi.org/10.1016/J.ETAP.2017.08.036

Tang X, Chen Y, Zhu X, Miao Y, Wang D, Zhang J, Li R, Zhang L, Chen J (2022) Alternariol monomethyl ether toxicity and geno- toxicity in male Sprague-Dawley rats: 28-Day *in vivo* multi-end- point assessment. *Mutat Res Genetic Toxicol Environ Mutagen* 873:10. https://doi.org/10.1016/J.MRGENTOX.2021.503435

Tiemann U, Tomek W, Schneider F, Müller M, Pöhland R, Vanselow J (2009) The mycotoxins alternariol and alternariol methyl ether negatively affect progesterone synthesis in porcine granulosa cells *in vitro*. *Toxicol Lett* 186(2):139–145. https://doi.org/10. 1016/j.toxlet.2009.01.014

Tiessen C, Fehr M, Schwarz C, Baechler S, Domnanich K, Böttler U, Pahlke G, Marko D (2013a) Modulation of the cellular redox sta- tus by the *Alternaria* toxins alternariol and alternariol monome- thyl ether. *Toxicol Lett* 216(1):23–30. https://doi.org/10.1016/J. TOXLET.2012.11.005

Tiessen C, Gehrke H, Kropat C, Schwarz C, Bächler S, Fehr M, Pahlke G, Marko D (2013b) Role of topoisomerase inhibition and DNA repair mechanisms in the genotoxicity of alternariol and alter- toxin-II. *World Mycotoxin J* 6(3):233–244. https://doi.org/10. 3920/wmj2013.1592

Tiessen C, Ellmer D, Mikula H, Pahlke G, Warth B, Gehrke H, Zim- mermann K, Heiss E, Fröhlich J, Marko D (2017) Impact of phase I metabolism on uptake, oxidative stress and genotoxicity of the emerging mycotoxin alternariol and its monomethyl ether in esophageal cells. *Arch Toxicol* 91(3):1213–1226. https://doi. org/10.1007/s00204-016-1801-0

Tran VN, Viktorova J, Augustynkova K, Jelenova N, Dobiasova S, Rehorova K, Fenclova M, Stranska-Zachariasova M, Vitek L, Hajslova J, Ruml T (2020) *In silico* and *in vitro* studies of myco- toxins and their cocktails; Their toxicity and its mitigation by silibinin pre-treatment. Toxins. https://doi.org/10.3390/toxin s12030148

Vejdovszky K, Sack M, Jarolim K, Aichinger G, Somoza MM, Marko D (2017) *In vitro* combinatory effects of the *Alternaria* mycotox- ins alternariol and altertoxin II and potentially involved miRNAs. *Toxicol Lett* 267:45–52. https://doi.org/10.1016/j.toxlet.2016.12. 011

Viegas S, Assunção R, Nunes C, Osteresch B, Twaružek M, Kosicki R, Grajewski J, Martins C, Alvito P, Almeida A, Viegas C (2018) Exposure assessment to mycotoxins in a Portuguese fresh bread dough company by using a multi-biomarker approach. Toxins 10(9):342. https://doi.org/10.3390/toxins10090342

Vila-Donat P, Fernández-Blanco C, Sagratini G, Font G, Ruiz MJ (2015) Effects of soyasaponin I and soyasaponins-rich extract on the Alternariol-induced cytotoxicity on Caco-2 cells. Food Chem Toxicol 77:44–49. https://doi.org/10.1016/j.fct.2014.12.016

VKM, Steffensen I, Fæste CK, Husøy T, Knutsen HK, Mathisen GH, Ørnsrud R, Agdestein A, Bodin J, Elvevoll E, Hessen DO, Hof- shagen M, Krogdahl A, Nilsen AM, Rafoss T, Skjerdal T, Velle G, Wasteson Y, Hemre G, Vandvik V, Alexander J (2019) Rank- ing of substances for monitoring in foods, drinks and dietary supplements - based on risk and knowledge gaps. Scientific Opinion of the Scientific Steering Committee of the Norwegian Scientific Committee for Food and Environment. VKM report 2019:13, ISBN: 978-82-8259-329-8. https://fhi.brage.unit.no/ fhi-xmlui/bitstream/handle/11250/2627587/Steffensen_2019_ Ran.pdf?sequence=2. Accessed 20 Aug 2021

Wang Y, Liu HX, Chen YC, Sun ZH, Li HH, Li SN, Yan ML, Zhang WM (2017) Two new metabolites from the endophytic fungus *Alternaria* sp. A744 derived from *Morinda officinalis*. *Molecules*. https://doi.org/10.3390/molecules22050765

Wollenhaupt K, Schneider F, Tiemann U (2008) Influence of alternariol (AOH) on regulator proteins of cap-dependent translation in porcine endometrial cells. *Toxicol Lett* 182(1–3):57–62. https://doi. org/10.1016/j.TOXLET.2008.08.005

World Health Organization (WHO) (2009) WHO Guideline for indoor air quality: dampness and mould. WHO Regional Office for Europe, Copenhagen, Denmark. https://www.who.int/publicatio ns/i/item/9789289041683. Accessed 18 Jan 2023

Xiao J, Zhang Q, Gao YQ, Tang JJ, Zhang AL, Gao JM (2014) Second- ary metabolites from the endophytic botryosphaeria dothidea of melia azedarach and their antifungal, antibacterial, antioxidant, and cytotoxic activities. J Agric Food Chem 62(16):3584–3590. https://doi.org/10.1021/jf500054f

Yekeler H, Bitmiş K, Özçelik N, Doymaz MZ, Calta M (2001) Analysis of toxic effects of *Alternaria* toxins on esophagus of mice by light and electron microscopy. *Toxicol Pathol* 29(4):492–497. https://doi.org/10.1080/01926230152499980

Zhao DL, Cao F, Wang CY, Yang LJ, Shi T, Wang KL, Shao CL, Wang CY (2019) Alternatone A, an unusual perylenequinone-related compound from a soft-coral-derived strain of the fungus *Alternaria alternata*. *J Nat Prod* 82(11):3201–3204. https://doi.org/ 10.1021/acs.jnatprod.9b00905

Zhu X, Chen Y, Tang X, Wang D, Miao Y, Zhang J, Li R, Zhang L, Chen J (2022) General toxicity and genotoxicity of altertoxin I: a novel 28-day multiendpoint assessment in male Sprague-Dawley rats. *J Appl Toxicol JAT* 42(8):1310–1322. https://doi.org/10. 1002/JAT.4297

**Publisher's Note** Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
Authors and Affiliations

Henriqueta Louro<fnref n="1" /> ID · Ariane Vettorazzi<fnref n="2" /> ID · Adela López de Cerain<fnref n="2" /> ID · Anastasia Spyropoulou<fnref n="3" /> ID ·

Anita Solhaug<fnref n="4" /> ID · Anne Straumfors<fnref n="4" /> ID · Anne-Cathrin Behr<fnref n="5" /> ID · Birgit Mertens<fnref n="6" /> ID · Bojana Žegura<fnref n="7" /> ID ·

Christiane Kruse Fæste<fnref n="4" /> ID · Dieynaba Ndiaye<fnref n="8" /> ID · Eliana Spilioti<fnref n="3" /> ID · Elisabeth Varga<fnref n="9" />,<fnref n="10" /> ID · Estelle Dubreil<fnref n="11" /> ID ·

Eszter Borsos<fnref n="10" /> ID · Francesco Crudo<fnref n="10" /> ID · Gunnar Sundstøl Eriksen<fnref n="4" /> ID · Igor Snapkow<fnref n="12" /> ID · Jérôme Henri<fnref n="11" /> ID ·

Julie Sanders<fnref n="6" /> ID · Kyriaki Machera<fnref n="3" /> ID · Laurent Gaté<fnref n="8" /> ID · Ludovic Le Hegarat<fnref n="11" /> ID · Matjaž Novak<fnref n="7" /> ID ·

Nicola M. Smith<fnref n="12" /> ID · Solveig Krapf<fnref n="4" /> ID · Sonja Hager<fnref n="9" /> ID · Valérie Fessard<fnref n="11" /> ID · Yvonne Kohl<fnref n="13" /> ID · Maria João Silva<fnref n="1" /> ID ·

Hubert Dirven<fnref n="12" /> ID · Jessica Dietrich<fnref n="14" /> ID · Doris Marko<fnref n="9" /> ID

<figure><img src="image_16.png" /></figure>

Doris Marko

doris.marko@univie.ac.at

<footnote n="1">¹ Department of Human Genetics, National Institute of Health Dr. Ricardo Jorge (INSA) and Centre for Toxicogenomics and Human Health (ToxOmics), NOVA Medical School, Universidade Nova de Lisboa, Av. Padre Cruz, 1649-016 Lisbon, Portugal</footnote>

<footnote n="2">² MITOX Research Group, Department of Pharmaceutical Sciences, Faculty of Pharmacy and Nutrition, UNAV-University of Navarra, Pamplona, Spain</footnote>

<footnote n="3">³ Laboratory of Toxicological Control of Pesticides, Scientific Directorate of Pesticides' Control and Phytopharmacy, Benaki Phytopathological Institute, 145 61 Attica, Greece</footnote>

<footnote n="4">⁴ Norwegian Veterinary Institute, PO Box 64, 1431 Ås, Norway</footnote>

<footnote n="5">⁵ Department Food Safety, BfR, German Federal Institute for Risk Assessment, Max-Dohrnstraße 8-10, 10589 Berlin, Germany</footnote>

<footnote n="6">⁶ Department of Chemical and Physical Health Risks, Sciensano, Brussels, Belgium</footnote>

<footnote n="7">⁷ Department of Genetic Toxicology and Cancer Biology, National Institute of Biology, Večna Pot 111, 1000 Ljubljana, Slovenia</footnote>

<footnote n="8">⁸ INRS, Institut National de Recherche et de Sécurité pour la Prévention des accidents du travail et des maladies professionnelles, Rue du Morvan, CS 60027, 54519 Vandœuvre Lès Nancy Cedex, France</footnote>

<footnote n="9">⁹ Department of Food Chemistry and Toxicology, Faculty of Chemistry, University of Vienna, Vienna, Austria</footnote>

<footnote n="10">¹⁰ Food Hygiene and Technology, University of Veterinary Medicine, Vienna, Veterinärplatz 1, 1210 Vienna, Austria</footnote>

<footnote n="11">¹¹ Toxicology of Contaminants Unit, Fougères Laboratory, French Agency for Food, Environmental and Occupational Health and Safety, 10 B rue Claude Bourgelat, 35306 Fougères, France</footnote>

<footnote n="12">¹² Department of Chemical Toxicology, Norwegian Institute of Public Health, Lovisenberggate 8, 0456 Oslo, Norway</footnote>

<footnote n="13">¹³ Fraunhofer Institute for Biomedical Engineering IBMT, Joseph-Von-Fraunhofer-Weg 1, 66280 Sulzbach, Germany</footnote>

<footnote n="14">¹⁴ Department Safety in the Food Chain, BfR, German Federal Institute for Risk Assessment, Max-Dohrn-Straße 8-10, 10589 Berlin, Germany</footnote>
Authors and Affiliations

Henriqueta Louro<fnref n="1" /> ID · Ariane Vettorazzi<fnref n="2" /> ID · Adela López de Cerain<fnref n="2" /> ID · Anastasia Spyropoulou<fnref n="3" /> ID ·

Anita Solhaug<fnref n="4" /> ID · Anne Straumfors<fnref n="4" /> ID · Anne-Cathrin Behr<fnref n="5" /> ID · Birgit Mertens<fnref n="6" /> ID · Bojana Žegura<fnref n="7" /> ID ·

Christiane Kruse Fæste<fnref n="4" /> ID · Dieynaba Ndiaye<fnref n="8" /> ID · Eliana Spilioti<fnref n="3" /> ID · Elisabeth Varga<fnref n="9" />,<fnref n="10" /> ID · Estelle Dubreil<fnref n="11" /> ID ·

Eszter Borsos<fnref n="10" /> ID · Francesco Crudo<fnref n="10" /> ID · Gunnar Sundstøl Eriksen<fnref n="4" /> ID · Igor Snapkow<fnref n="12" /> ID · Jérôme Henri<fnref n="11" /> ID ·

Julie Sanders<fnref n="6" /> ID · Kyriaki Machera<fnref n="3" /> ID · Laurent Gaté<fnref n="8" /> ID · Ludovic Le Hegarat<fnref n="11" /> ID · Matjaž Novak<fnref n="7" /> ID ·

Nicola M. Smith<fnref n="12" /> ID · Solveig Krapf<fnref n="4" /> ID · Sonja Hager<fnref n="9" /> ID · Valérie Fessard<fnref n="11" /> ID · Yvonne Kohl<fnref n="13" /> ID · Maria João Silva<fnref n="1" /> ID ·

Hubert Dirven<fnref n="12" /> ID · Jessica Dietrich<fnref n="14" /> ID · Doris Marko<fnref n="9" /> ID

<figure><img src="image_18.png" /></figure>

Doris Marko
doris.marko@univie.ac.at

<footnote n="1">¹ Department of Human Genetics, National Institute of Health Dr. Ricardo Jorge (INSA) and Centre for Toxicogenomics and Human Health (ToxOmics), NOVA Medical School, Universidade Nova de Lisboa, Av. Padre Cruz, 1649-016 Lisbon, Portugal</footnote>

<footnote n="2">² MITOX Research Group, Department of Pharmaceutical Sciences, Faculty of Pharmacy and Nutrition, UNAV-University of Navarra, Pamplona, Spain</footnote>

<footnote n="3">³ Laboratory of Toxicological Control of Pesticides, Scientific Directorate of Pesticides' Control and Phytopharmacy, Benaki Phytopathological Institute, 145 61 Attica, Greece</footnote>

<footnote n="4">⁴ Norwegian Veterinary Institute, PO Box 64, 1431 Ås, Norway</footnote>

<footnote n="5">⁵ Department Food Safety, BfR, German Federal Institute for Risk Assessment, Max-Dohrnstraße 8-10, 10589 Berlin, Germany</footnote>

<footnote n="6">⁶ Department of Chemical and Physical Health Risks, Sciensano, Brussels, Belgium</footnote>

<footnote n="7">⁷ Department of Genetic Toxicology and Cancer Biology, National Institute of Biology, Večna Pot 111, 1000 Ljubljana, Slovenia</footnote>

<footnote n="8">⁸ INRS, Institut National de Recherche et de Sécurité pour la Prévention des accidents du travail et des maladies professionnelles, Rue du Morvan, CS 60027, 54519 Vandœuvre Lès Nancy Cedex, France</footnote>

<footnote n="9">⁹ Department of Food Chemistry and Toxicology, Faculty of Chemistry, University of Vienna, Vienna, Austria</footnote>

<footnote n="10">¹⁰ Food Hygiene and Technology, University of Veterinary Medicine, Vienna, Veterinärplatz 1, 1210 Vienna, Austria</footnote>

<footnote n="11">¹¹ Toxicology of Contaminants Unit, Fougères Laboratory, French Agency for Food, Environmental and Occupational Health and Safety, 10 B rue Claude Bourgelat, 35306 Fougères, France</footnote>

<footnote n="12">¹² Department of Chemical Toxicology, Norwegian Institute of Public Health, Lovisenberggate 8, 0456 Oslo, Norway</footnote>

<footnote n="13">¹³ Fraunhofer Institute for Biomedical Engineering IBMT, Joseph-Von-Fraunhofer-Weg 1, 66280 Sulzbach, Germany</footnote>

<footnote n="14">¹⁴ Department Safety in the Food Chain, BfR, German Federal Institute for Risk Assessment, Max-Dohrn-Straße 8-10, 10589 Berlin, Germany</footnote>