Alternaria Mikotoksinleri ve Önemi

Yıl 2023, , 195 - 219, 01.06.2023

Berna Tunalı , Yeter Küçüktopcu , Nazlı Tunalı , Songül Erken Meral , Seçil Eker , Bayram Kansu

https://doi.org/10.20479/bursauludagziraat.1111062

Cited By: 1

Öz

Alternaria, dünyada yaygın olarak görülen önemli bir fungus cinsi olup Ascomycota bölümü,
Dothideomycetes sınıfı, Pleosporales takımı ve Pleosporaceae familyasında yer almaktadır. Alternaria cinsi
içerisinde, saprofitik, endofitik ve patojenik türler yer almaktadır. Patojen türler arasında ise bitki patojenleri,
hasat sonrası patojenler veya insan patojenleri de bulunmaktadır. Alternaria spp. alternariol (AOH), alternariol
monometil eter (AME), tenuazonik asit (TEA), altenuen (ALT) ve altertoksin (AT) gibi önemli bazı
mikotoksinleri üretmektedir. Mikotoksinler, insan besin zincirine çeşitli şekillerde girebilmekte, birçok farklı
gıda ve hayvan yemi ürününde bulunabilmektedir. Bu mikotoksinler, insanlar, memeliler ve diğer hayvanlar
tarafından ağız yoluyla alınırsa, mikotoksikoz adı verilen toksik bir tepkiye neden olabilmektedir. Birçoğunun
kanserojen olduğu bilinmektedir. Diğerlerinin de cilt hassasiyetinden immün yetmezliğe kadar değişen
nörotoksikolojik etkilerle birlikte karaciğer veya böbrek fonksiyonunun bozulması gibi insanlarda çeşitli farklı
tepkiler ortaya çıkardığı gösterilmiştir. Alternaria spp., özellikle su aktivitesi (aw), sıcaklık ve pH gibi abiyotik
faktörlerden etkilenmektedir. Literatürdeki çalışmalara göre bazı tahıl taneleri dahil sorgum, pamuk tohumu, domates ve soya fasulyesi gibi farklı substratlar fungusun çoğalması ve toksin üretimi ile ilişkilendirilmiştir.
Alternaria toksinlerinin incelenmesinde ELISA, sıvı kromatografi ve PCR temelli analizler en kullanışlı
yöntemler olarak görülmektedir. Bu derleme, Alternaria türlerinin önemini, ekolojilerini, mikotoksin üretimi ve
sıcakkanlılardaki etkileri ile mikotoksin analiz metotlarını içermektedir. Derleme özellikle, Alternaria türlerinin
oluşturdukları mikotoksinler hakkında genel bir bilgi sunmak ve önemine dikkat çekmek amacıyla
hazırlanmıştır.

Anahtar Kelimeler

Alternaria türleri, mikotoksin, insan sağlığı, kromatografi, inceleme

Alternaria Mycotoxins and Their Importance

Öz

Alternaria, which are an important genus of fungi that are ubiquitous in the environment, belongs to
the Ascomycota division, Dothideomycetes class, Pleosporales order, and Pleosporaceae family. The Alternaria
genus includes saprophytic, endophytic, and pathogenic species, whereas the pathogenic species include plant,
post-harvest, and human pathogens. Alternaria spp. produce some important mycotoxins such as alternariol
(AOH), alternariol monomethyl ether (AME), tenuazonic acid (TEA), altenuene (ALT) ve altertoxin (AT). These
mycotoxins can gain access to the human food chain and exist in many different food products and forage. When
ingested orally by humans, mammalians, or other animals, these mycotoxins may cause mycotoxicosis, which is
a toxic reaction. Many of these mycotoxins are known to be carcinogenic, while some others are shown to
disrupt liver and kidney functions as well as have neurotoxicological effects ranging from skin hypersensitivity
reactions to immunologic deficiency syndromes. Alternaria spp. can get affected by abiotic factors such as water
activity (aw), temperature, and pH. In literature, many crops including sorghum, cotton, tomato, and soya beans,
have been related the conditions that result in fungi to reproducing and producing toxins. The most effective
methods established for analyzing the Alternaria toxins are ELISA, liquid chromatography, and PCR-based
methods. This review includes the importance of Alternaria spp., their ecologies, mycotoxin production and their
effect on the homeotherms, and the methods for analyzing mycotoxins. It is specifically aimed to highlighting
the importance and providing a general knowledge of producing mycotoxins that Alternaria spp.

Anahtar Kelimeler

Alternaria spp., mycotoxin, human health, chromatography, detection

Kaynakça

• Abdel-Hadi, A., Schmidt-Heydt, M., Parra, R., Geisen, R. and Magan, N. 2012. A systems approach to model the relationship between aflatoxin gene cluster expression, environmental factors, growth and toxin production by Aspergillus flavus. Journal of the Royal Society Interface, 9: 757-767.
• Aichinger, G., Puntscher, H., Beisl, J., Kütt, M-L., Warth, B. and Marko, D. 2018. Delphinidin protects colon carcinoma cells against the genotoxic effects of the mycotoxin altertoxin II. Toxicology Letters, 284: 136- 142.
• Akamatsu, H., Itoh, Y., Kodama, M., Otani, H. and Kohmoto, K. 1997. AAL-toxin-deficient mutanst of Alternaria alternata tomato pathotype by restriction enzyme-mediated integration. Phytopathology, 87: 967-972.
• Akamatsu, H., Taga, M., Kodama, M., Johnson, R., Otani, H., and Kohmoto, K. 1999. Molecular karyotypes for Alternaria plant pathogens known to produce host-specific toxins. Current Genetic, 35: 647–656. doi: 10.1007/s002940050464.
• Akinmitsu, K., Tsuge, T. and Kodama, M. 2013. Alternaria host-selective toxins: determinant factors of plant disease. Journal of Gen. Plant Pathology, 80: 109-122.
• AlMatar, M., Var, I., and Koksal, F. 2016. How Does Alternaria alternata-Derived Alternariol Affect Our Health? Mini-Reviews in Organic Chemistry, 13: 465-472.
• Andersen, B., Nielsen, K.F., Pinto, V.F. and Patriarca, A. 2015. Characterization of Alternaria strains from Argentinean blueberry, tomato, walnut and wheat. International Journal of Food Microbiology, 196: 1-10.
• Andrew, M., Peever, T. L., and Pryor, B. M. 2009. An expanded multilocus phylogeny does not resolve morphological species within the small-spored Alternaria species complex. Mycologia, 101: 95–109. doi: 10.3852/08-135.
• Arcella, D., Eskola, M., Ruiz, J.A.G. 2016. Dietary exposure assessment to Alternaria toxins in the European population. EFSA Journal 14:4654.
• Azcarate, M.P., Patriarca, A., Terminiello, L. and Pinto, V.F. 2008. Alternaria toxins in wheat during the 2004 to 2005 Argentinean harvest. Journal of Food Protection, 71: 1262-1265.
• Barkai-Golan, R. 2008. Alternaria Mycotoxins: Mycotoxins in Fruits and Vegetables. Ed: Barkai-Golan, R., Paster, N., Elsevier Inc., San Diego, CA, USA, pp: 185-203.
• Barkai-Golan, R. and Paster, N. 2008. Mouldy fruits and vegetables as a source of mycotoxins: part 1. World Mycotoxin Journal, 1:147-159.
• Bensassi, F., Gallerne, C., Sharaf El Dein, O., Hajlaoui, M.R., Bacha, H., Lemaire, C. 2012. Cell death induced by the Alternaria mycotoxin Alternariol. Toxicology in Vitro, 26:915-923.
• Berestetskly, A.O. 2008. A review of fungal phytotoxins: from basic studies to practical use. Applied Biochemical and Microbiology, 44: 453-465.
• Bottalico, A. and Logrieco, A. 1998. Alternaria species of economic importance: Mycotoxins in agriculture and food safety. Ed: Sinha, K.K., Bhatnagar, D., Marcel Dekker. New York, USA, p 65-108.
• Boutin, B., Peeler, J., Twedt, R. 1989. Effects of purified altertoxins I, II, and III in the metabolic communication V79 system. Journal of Toxicology and Environmental Health, Part A, 26:75-81.
• Brugger, E-M., Wagner, J., Schumacher, D.M., Koch, K., Podlech, J., Metzler, M. and Lehmann, L. 2006. Mutagenicity of the mycotoxin alternariol in cultured mammalian cells. Toxicology Letters, 164: 221-230.
• Charmley, L.L., Trenholm, H.L., Prelusky, D.B. and Rosenberg, A. 1995. Economic losses and decontamination. Natural Toxins, 3: 199-203.
• Chen, A., Mao, X., Sun, Q., Wei, Z., Li, J., You, Y., Zhao, J., Jiang, G., Wu, Y., Wang, L., Li, Y. 2021. Alternaria Mycotoxins: An Overview of Toxicity, Metabolism, and Analysis in Food. Journal of Agricultural and Food Chemistry, 69:7817-7830.
• Chowdappa, P., Lakshmi, M. J., Madhura, S. 2013. Matrix assisted laser desorption ionization–time of flight (MALDI-TOF) mass spectrometry for identification of plant pathogenic Alternaria species. Phytoparasitica, 41: 169-179.
• Chulze, S.N., Torres, A.M., Dalcero, A.M., Etcheverry, M.G, Ramírez, M.L. and Farnochi, M.C. 1995. Alternaria mycotoxins in sunflower seeds: incidence and distribution of the toxins in oil and meal. Journal of Food Protection, 58: 1133-1134.
• Crespo-Sempere, A., Estiarte, N., Marín, S., Sanchis, V. and Ramos, A.J. 2013. Propidium monoazide combined with real-time quantitative PCR to quantify viable Alternaria spp. contamination in tomato products. International Journal of Food Microbiology, 165: 214-220.
• Dall’Asta, C., Cirlini, M., Falavigna, C. 2014. Chapter Three - Mycotoxins from Alternaria: Toxicological Implications. Advances in Molecular Toxicology, 8:107-121.
• Dang, H.X., Pryor, B., Peever, T. and Lawerence, C.B. 2015. The Alternaria genomes database: a comprehensive resource for a fungal genus comparised of saprophytes, plant pathogens, and allergic species. BMC Genomics, 16: 239.
• Davis, N.D., Diener, U.L., Morgan-Jones, G. 1977. Tenuazonic acid production by Alternaria alternata and Alternaria tenuissima isolated from cotton. Applied and Environmental Microbiology, 34:155-157.
• Dearborn, D.G., Infeld, M.D., Smith, P., Judge, C., Horgan, T.E., Allan, T., Zimomra, J.A., Mortensen, B.K., Burkett, S.A. and Winpisingerslay, K. 1995. Acute Pulmonary Hemorrhage Hemosiderosis Among InfantsCleveland, January-1993 Novenber-1994. Journal of the American Medical Association, 273: 281-282.
• Dellafiora, L., Dall’Asta, C., Cruciani, G., Galaverna, G. and Cozzini, P. 2015. Molecular modelling approach to evaluate poisoning of topoisomerase I by alternariol derivatives. Food Chemistry, 189: 93-101.
• Dellafiora, L., Warth, B., Schmidt, V., Del Favero, G., Mikula, H., Fröhlich, J. and Marko, D. 2018. An integrated in silico/in vitro approach to assess the xenoestrogenic potential of Alternaria mycotoxins and metabolites. Food Chemistry, 248: 253-261.
• EFSA Panel on Contaminants in the Food Chain (CONTAM) 2011. Scientific opinion on the risks for animal and public health related to the presence of Alternaria toxins in feed and food. EFSA Journal, 9: 2407.
• Escriva, L., Font, G., Manyes, L., Berrada, H. 2017. Studies on the Presence of Mycotoxins in Biological Samples: An Overview. Toxins, 9:251.
• Fabrega, A., Agut, M. and Calvo, M. 2002. Optimization of the method of detection of metabolites produced by the Alternaria genus: Alternariol, alternariol monomethyl ether, altenuene, altertoxin I and tentoxin. Journal of Food Science, 67: 802-806.
• Fehr, M., Pahlke, G., Fritz, J., Christensen, M.O., Boege, F., Altemöller, M. and Podlech J, Marko D. 2009. Alternariol acts as a topoisomerase poison, preferentially affecting the IIα isoform. Molecular nutrition & Food Research, 53: 441-451.
• Fleck, S.C., Sauter, F., Pfeiffer, E., Metzler, M., Hartwig, A., Koberle, B. 2016. DNA damage and repair kinetics of the Alternaria mycotoxins alternariol, altertoxin II and stemphyltoxin III in cultured cells. Mutation Research/Genetic Toxicology and Environmental Mutagenesis, 798-799:27-34.
• Freire, F.D.C.O. and da Rocha, M.E.B. 2017. Impact of Mycotoxins on Human Health: Fungal Metabolites. Ed.: Merillon, J.M., Ramawat, R.G., Springer International Publishing, Switzerland, pp: 239-255.
• Friesen, T.L., Faris, J.D., Solomon, P.S. and Oliver, R. P. 2008. Host-specific toxins: effectors of necrotrophic pathogenicity. Cell Microbiology, 10: 1421-1428.
• Frizzell, C., Ndossi, D., Kalayou, S., Eriksen, G.S., Verhaegen, S., Sørlie, M., Elliott, C.T, Ropstad, E. and Connolly L. 2013. An in vitro investigation of endocrine disrupting effects of the mycotoxin alternariol. Toxicology and Applied Pharmacology, 271: 64-71.
• Goryacheva, I. Y., De Saeger, S., Eremin, S. A., Van Peteghem, C. 2007. Immunochemical methods for rapid mycotoxin detection: Evolution from single to multiple analyte screening: A review. Food Additives and Contaminants, 24(10): 1169-1183.
• Griffin, G.F. and Chu, F. 1983. Toxicity of the Alternaria metabolites alternariol, alternariol methyl ether, altenuene, and tenuazonic acid in the chicken embryo assay. Applied and Environmental Microbiology, 46: 1420-1422.
• Gross, M., Curtui, V., Ackermann, Y, Latif H, and Usleber E. 2011. Enzyme immunoassay for tenuazonic acid in apple and tomato products. Journal of Agricultural and Food Chemistry, 59: 12317-12322.
• Gürcan, Ş., Pişkin, S., Kılıç, H., Temelli, B.A. ve Yalçın, Ö. 2009. İmmün Sistemi Sağlam Bir Konakta Alternaria alternata ile Oluşan Deri Enfeksiyonu. Mikrobiyoloji Bülteni, 43: 163-167.
• Harvan, D. and Pero, R. 1974. Gas chromatographic analysis of the Alternaria metabolite, tenuazonic acid. Journal of Chromatography A, 101: 222-224.
• Hasan, H. 1995. Alternaria mycotoxins in black rot lesion of tomato fruit: conditions and regulation of their production. Mycopathologia, 130: 171-177.
• Hatta, R., Ito, K., Hosaki, Y., Tanaka, T., Tanaka, A. and Yamamoto, M. 2002. A conditionally dispensable chromosome controls host-specific pathogenicity in the fungal plant pathogen Alternaria alternata. Genetics, 161: 59–70.
• Haubruge, E., Chasseur, C., Debouck, C., Begaux, F., Suetens, C., Mathieu, F., Michel, V., Gaspar, C., Rooze, M. and Hinsenkamp M. 2001. The prevalence of mycotoxins in Kashin-Beck disease. International Orthopaedics, 25: 159-161.
• Hou, Y., Ma, X., Wan, W., Long, N., Zhang, J., Tan, Y., Duan, S., Zeng, Y. and Dong, Y. 2016. Comparative Genomics of Pathogens Causing Brown Spot Disease of Tobacco: Alternaria longipes and Alternaria alternata. PLoS ONE, 11: e0155258.
• Howlett, B. J. 2006. Secondary metabolite toxins and nutrition of plant pathogenic fungi. Current Opinion Plant Biology, 9: 371-375.
• Hu, J., Chen, C., Peever, T., Dang, H., Lawrence, C.B. and Mitchell, T.K. 2012. Genomic characterization of the conditionally dispensable chromosome in Alternaria arborescens provides evidence for horizontal gene transfer. BMC Genomics, 13: 171.
• Izmailov, N.A. and Shraiber, M.S. 1938. A drop-chromatographic method of analysis and its applications to pharmacy. Farmatsiya, 3:1-3.
• Karabulut, Ö.A. ve Değirmencioğlu, T. 2002. Hayvan Yemi Olarak Kullanılan Buğday Danelerinde Toksin Oluşumuna Neden Olan Fungusların Sodyum Hidroksit Uygulamasıyla Engellenmesi. Uludağ Üniversitesi, Ziraat Fakültesi Dergisi, 16: 129-138.
• Kaya, B. ve Zorba, N.N. 2021. Alternaria Genusu Üyelerinin Meyve ve Sebzeler Üzerine Etkileri. Mantar Dergisi, 12:223-239.
• Kocher, U. 2006. Multimethode zur Bestimmung von Alternaria-toxinen mittels LC-MS/MS. In: Gesellschaft für Mykotoxinforschung. Proceedings of the 28th Mycotoxin Workshop. Bydgoszcz, Poland: Gesellschaftfür Mykotoxin Forschung, p92.
• Köppen, R., Koch, M., Siegel, D., Merkel, S., Maul, R., Nehls, I. 2010. Determination of mycotoxins in foods: current state of analytical methods and limitations. Applied Microbiology and Biotechnology, 86: 1595-1612.
• Lee, H.B., Patriarca, A. and Magan, H. 2015. Alternaria in food: ecophysiology, mycotoxin production and toxicology. Microbiology, 43: 93-106.
• Lehmann, L., Esch, H.L., Wagner, J., Rohnstock, L., Metzler, M. 2005. Estrogenic and genotoxic potential of equol and two hydroxylated metabolites of Daidzein in cultured human Ishikawa cells. Toxicology Letters, 158:72-86.
• Licorish, K., Novey, H.S., Kozak, P., Fairshter, R.D. and Wilson, A.F. 1985. Role of Altemaria and Penicillium spores in the pathogenesis of asthma. Journal of Allergy and Clinical Immunology, 76: 819-825.
• Liu, G., Qian, Y., Zhang, P., Dong, Z., Shi, Z., Zhen, Y., Miao, J. and Xu, Y. 1991. Relationships between Alternaria alternata and oesophageal cancer. IARC Scientific Publications, 105: 258-262.
• Liu, G., Qian, Y., Zhang, P., Dong, W., Qi, Y. and Guo, H. 1992. Etiological role of Alternaria alternata in human esophageal cancer. Chinese Medical Journal, 105: 394-400.
• Logrieco, A., Bottalico, A., Mule, G., Moretti, A., Perrone, G. 2003. Epidemiology of toxigenic fungi and their associated mycotoxins for some Mediterranean crops. European Journal of Plant Pathology, 109:645-667.
• Logrieco, A., Moretti, A., Solfrizzo, M. 2009. Alternaria toxins and plant diseases: an overview of origin, occurence and risks. World Mycotoxin Journal, 2:129-140.
• López, P., Venema, D., de Rijk, T., de Kok, A., Scholten, J.M., Mol, H.G., de Nijs, M. 2016. Occurrence of Alternaria toxins in food products in The Netherlands. Food Control, 60: 196-204.
• Magan, N., Cayley, G.R. and Lacey, J. 1984. Effect of water activity and temperature on mycotoxin production by Alternaria alternata in culture and on wheat grain. Applied and Environmental Microbiology, 47: 1113- 1117.
• Magan, N. and Baxter, E. 1994. Relationship between environmental factors and tenuazonic acid production by Alternaria isolates from sorghum. In: Scudamore, K.A., editor. Occurrence and significance of mycotoxins. Slough: Central Science Laboratories. p 309-313.
• Magan, N. and Aldred, D. 2007. Why do fungi produce mycotoxins? In: Dijksterhuis, J., Samson, R.A., editors. Food mycology: a multifaceted approach to fungi and food. Boca Raton: Taylor and Francis. p 121-133.
• Man, Y., Liang, G., Li, A., Pan, L. 2017. Analytical Methods for the Determination of Alternaria Mycotoxins. Chromatographia, 80:9-22.
• Mannon, J. and Johnson, E. 1985. Fungi down on the farm. New Scientist, 105: 12-16.
• Markham, J.E. 2001. Host-selective toxins as agents of cell death in plant-fungus interactions. Molecular Plant Pathology, 2(4): 229-239.
• Matysik, G. and Giryn, H. 1996. Gradient thin-layer chromatography and densitometry determination of Alternaria mycotoxins. Chromatographia, 42: 555-558.
• Medina, A., Schmidt-Heydt, M., Cárdenas-Chávez, D.L., Parra, R., Geisen, R. and Magan, N. 2013. Integrating toxin gene expression, growth and fumonisin B1 and B2 production by a strain of Fusarium verticillioides under different environmental factors. Journal of the Royal Society Interface, 10: 20130320.
• Meena, M., Zehra, A., Dubey, M.K., Aamir, M., Gupta, V.K. and Upadhyay, R.S. 2016. Comparative evaluation of biochemical changes in tomato (Lycopersicon esculentum Mill.) infected by Alternaria alternata and its toxic metabolites (TeA, AOH, and AME). Frontier Plant Science, 7: 1408.
• Meena, M. and Samal, S. 2019. Alternaria host-specific (HSTs) toxins: An overview of chemical characterization, target sites, regulation and their toxic effects. Toxicology Reports, 6: 745-758.
• Menzies, D., Comtois, P., Pasztor, J., Nunes, F. and Hanley, J.A. 1998. Aeroallergens and work-related respiratory symptoms among office workers. Journal of Allergy and Clinical Immunology, 101: 38-44.
• Moressi, M., Zon, A., Fernandez, H., Rivas, G. and Solis, V. 1999. Amperometric quantification of Alternaria mycotoxins with a mushroom tyrosinase modified carbon paste electrode. Electrochemistry Communications, 1:472-476.
• Morgan, M. R. A. 1989. Mycotoxin immunoassays (with special reference to elisas). Tetrahedron, 45(8): 2237- 2249.
• Mujahid, C., Marie-Claude, S., Basle, Q., Woo, P.M., Yean Ee, E.C., Mottier, P., Bessaire, T. 2020. Levels of Alternaria Toxins in Selected Food Commodities Including Green Coffee. Toxins, 12:595.
• Muller, M. and Lepom, P. 1992. The detection of Alternaria mycotoxins in laboratory culture. Zentralblatt fur Mikrobiologie, 147:197-206.
• Müller, M.E. and Korn, U. 2013. Alternaria mycotoxins in wheat–a 10 years survey in the Northeast of Germany. Food Control, 34: 191-197.
• Newberne, P.M. and Butler, W.H. 1969. Acute and Chronic Effects of Aflatoxin on the Liver of Domestic and Laboratory Animals : A Review. Cancer Research, 29:236-250.
• Nishimura, S. and Kohmoto, K. 1983. Host-specific toxins and chemical structures form Alternaria species. Annual Review Phytopathology, 21: 87-116.
• Noser, J., Schneider, P., Rother, M., Schmutz, H. 2011. Determination of six Alternaria toxins with UPLCMS/MS and their occurrence in tomatoes and tomato products from the Swiss market. Mycological Research, 27:265-271.
• Ostry, V., Skarkova, J., Prochazkova, I., Kubatova, A. and Ruprich, J. 2007. Mycobiota of Czech wine grapes and occurrence of ochratoxin A and Alternaria mycotoxins in fresh grape juice, must and wine. Czech Mycology, 59: 241.
• Ostry, V. 2008. Alternaria mycotoxins: an overview of chemical characterization, producers, toxicity, analysis and occurrence in foodstuffs. World Mycotoxin Journal, 1: 175-188.
• Oviedo, M., Ramirez, M., Barros, G. and Chulze, S. 2009. Effect of environmental factors on tenuazonic acid production by Alternaria alternata on soybean‐based media. Journal of Applied Microbiology, 107: 1186- 1192.
• Oviedo, M., Ramirez, M., Barros, G. and Chulze, S. 2010. Impact of water activity and temperature on growth and alternariol and alternariol monomethyl ether production of Alternaria alternata isolated from soybean. Journal of Food Protection, 73: 336-343.
• Oviedo, M.S., Ramirez, M.L., Barros, G.G. and Chulze, S.N. 2011. Influence of water activity and temperature on growth and mycotoxin production by Alternaria alternata on irradiated soya beans. International Journal of Food Microbiology, 149: 127-132.
• Oviedo, M.S., Sturm, M.E., Reynoso, M.M., Chulze, S.N. and Ramirez, M.L. 2013. Toxigenic profile and AFLP variability of Alternaria alternata and Alternaria infectoria occurring on wheat. Brazilian Journal of Microbiology, 44:447-455.
• Patriarca, A., Medina, A., Pinto, V.F. and Magan, N. 2014. Temperature and water stress impacts on growth and production of altertoxin-II by strains of Alternaria tenuissima from Argentinean wheat. World Mycotoxin Journal, 7: 329-334.
• Pero, R., Posner, H., Blois, M., Harvan, D. and Spalding, J. 1973. Toxicity of metabolites produced by the" Alternaria". Environmental Health Perspectives, 4: 87-94.
• Pinto, V.E.F. and Patriarca, A. 2017. Alternaria Species and Their Associated Mycotoxins: Mycotoxigenic Fungi: Methods and Protocols. Ed: Moretti, A., Susca, A., New York, USA: Springer Science+Business Media LLC., pp: 13-32.
• Pose, G., Patriarca, A., Kyanko, V., Pardo, A. and Pinto, V.F. 2010. Water activity and temperature effects on mycotoxin production by Alternaria alternata on a synthetic tomato medium. International Journal of Food Microbiology, 142: 348-353.
• Pryor, B.M. and Gilbertson, R.L. 2000. Molecular phylogenetic relationships amongst Alternaria species and related fungi based upon analysis of nuclear ITS and mt SSU rDNA sequences. Mycological Research 104:1312-1321.
• Rahman, H.U., Yue, X., Yu, Q., Xie, H., Zhang, W., Zhang, Q., Li, P. 2019. Specific antigen-based and emerging detection technologies of mycotoxins. Journal of the Science of Food and Agriculture, 99:4869- 4877.
• Rivoire, B., Attucci, S., Anthonioz, P., Carre, P., Lemarie, E. and Hazouard, E. 2001. Occupational acute lung injury due to Alternaria alternata: early stage of organic dust toxic syndrome requires no corticosteroids. Intensive Care Medicine, 27: 1236.
• Rychlik, M., Lepper, H., Weidner, C., Asam, S. 2016. Risk evaluation of the Alternaria mycotoxin tenuazonic acid in foods for adults and infants and subsequent risk management. Food Control, 68:181-185.
• Sanchis, V. and Magan, N. 2004. Environmental profiles for growth and mycotoxin production: Mycotoxins in food: detection and control, Ed: Magan, N., Olsen, M.,Cambridge: Woodhead Publishing Ltd., pp: 174-189.
• Sauer, D.B., Seitz, L.M., Burroughs, R., Mohr, H.E., West, J.L., Milleret, R.J. and Anthony, H.D. 1978. Toxicity of Alternaria metabolites found in weathered sorghum grain at harvest. Journal of Agricultural and Food Chemistry, 26: 1380-1383.
• Schrader, T., Cherry, W., Soper, K. and Langlois, I. 2006. Further examination of the effects of nitrosylation on Alternaria alternata mycotoxin mutagenicity in vitro. Mutation Research/Genetic Toxicology and Environmental Mutagenesis, 606: 61-71.
• Scott, P. M. 1995. Mycotoxin methodology. Food Additives and Contaminants, 12(3): 395-403.
• Scott, P.M., Weber, D., Kanhere, S.R. 1997. Gas chromatographymass spectrometry of Alternaria mycotoxins. Journal of Chromatography A, 765:255-263.
• Simmons, E.G. 1992. Alternaria taxonomy: current status, viewpoint, challenge: Alternaria biology, plant diseases and metabolites, Ed: Chelkowski, J. and Visconti, A., Amsterdam, Elsevier Science Publisher, pp: 1-35.
• Sivagnanam, K., Komatsu, E., Rampitsch, C., Perreault, H., Grafenhan, T. 2017. Rapid screening of Alternaria mycotoxins using MALDI-TOF mass spectrometry. Journal of the Science of Food and Agriculture, 97:357- 361.
• Skarkova, J., Ostry, V. and Prochazkova, I. 2005. Planar chromatographic determination of Alternaria toxins in selected foodstuffs. In: Proceedings of the international symposium on planar separations, planar chromatography, milestones in instrumental TLC. Siofok, Hungary: RIMP. p 29-31.
• Smith, J.E., Solomons, G., Lewis, C. and Anderson, J.G. 1995. Role of Mycotoxins in Human and Animal Nutrition and Health. Natural Toxins, 3:187-192.
• Solfrizzo, M., De Girolamo, A., Vitti, C., Visconti, A. and van den Bulk, R. 2004. Liquid chromatographic determination of Alternaria toxins in carrots. Journal of AOAC International, 87: 101-106.
• Solhaug, A., Vines, L.L., Ivanova, L., Spilsberg, B., Holme, J.A., Pestka, J., Collins, A., Eriksen, G.S. 2012. Mechanisms involved in alternariol-induced cell cycle arrest. Mutation Research/Genetic Toxicology and Environmental Mutagenesis, 738-739:1-11.
• Somma, S., Pose, G., Pardo, A., Mulè, G., Pinto, V.F., Moretti, A. and Logrieco, A.F. 2011. AFLP variability, toxin production, and pathogenicity of Alternaria species from Argentinean tomato fruits and puree. International Journal of Food Microbiology, 145: 414-419.
• Stack, M.E., Mislivec, P.B., Roach, J.A. and Pohland, A.E. 1985. Liquid chromatographic determination of tenuazonic acid and alternariol methyl ether in tomatoes and tomato products. Journal of the Association of Official Analytical Chemists, 68: 640-642.
• Steyn, P. and Stander, M. 1999. Mycotoxins with special reference to the carcinogenic mycotoxins: aflatoxins, ochratoxins and fumonisins. General and Applied Toxicology. 2nd Edition United Kingdom: Macmillan Reference Ltd:2145-2176.
• Sydenham, E.W. and Shephard, G.S. 1996. Chromatographic and allied methods of analysis for selected mycotoxins; Progress in Food Contaminant Analysis, Ed: J. Gilbert, Norwich, UK, Blackie Academic&Professional, pp: 65-146.
• Tatlıdil, S., Bıçakçı, A., Akkaya, A. ve Malyer H. 2001. Burdur Atmosferindeki Allerjen Cladosporium sp. ve Alternaria sp. Sporları. Süleyman Demirel Üniversitesi Tıp Fakültesi Dergisi, 8: 1-3.
• Thomma, B.P. 2003. Alternaria spp.: from general saprophyte to specific parasite. Molecular Plant Pathology, 4: 225-236.
• Tsuge, T., Harimoto, Y., Akimitsu, K., Ohtani, K., Kodama, M., Akagi, Y., Egusa, M., Yamamoto, M. and Otani, H. 2013. Host-selective toxins produced by the plant pathogenic fungus Alternaria alternata. FEMS Microbiology Reviews, 37: 44-66.
• Tunail, N. 2000. Funguslar ve mikotoksinler. Gıda Mikrobiyolojisi ve Uygulamaları, Genişletilmiş 2: 1-50.
• Turner, N.W., Subrahmanyam, S., Piletsky, S.A. 2009. Analytical methods for determination of mycotoxins: A review. Analytica Chimica Acta, 632(2): 168-180.
• Visconti, A., Sibilia, A., Palmisano, F. 1991. Selective determination of altertoxins by high-performance liquid chromatography with electrochemical detection with dual “in-series” electrodes. Journal of Chromatography A, 540: 376-382.
• Vlata, Z., Porichis, F., Tzanakakis, G., Tsatsakis, A. and Krambovitis, E. 2005. In vitro cytopathic effects of mycotoxin T-2 on human peripheral blood T lymphocytes. Toxicological Letters, 160(1): 60-68.
• Vlata, Z., Porichis, F., Tzanakakis, G., Tsatsakis, A. and Krambovitis, E. 2006. A study of zearalenone cytotoxicity on human peripheral blood mononuclear cells. Toxicological Letters, 165(3): 274-281.
• Walravens, J., Mikula, H., Rychlik, M., Asam, S., Ediage, E.N., Di Mavungu, J.D., Van Landschoot, A., Vanhaecke, L. and De Saeger, S. 2014. Development and validation of an ultra-high-performance liquid chromatography tandem mass spectrometric method for the simultaneous determination of free and conjugated Alternaria toxins in cereal-based foodstuffs. Journal of Chromatography A, 1372: 91-101.
• Wang, Y-J., Nie, J-Y., Zhen, Y., Li, Z-X., Cheng, Y. and Farooq, S. 2018. Multi-mycotoxin exposure and risk assessments for Chinese consumption of nuts and dried fruits. Journal of Integrative Agriculture, 17:1676- 1690.
• Warth, B., Parich, A., Atehnkeng, J., Bandyopadhyay, R., Schuhmacher, R., Sulyok, M., Krska, R. 2012. Quantitation of mycotoxins in food and feed from Burkina Faso and Mozambique using a modern LCMS/MS multitoxin method. Journal of Agricultural and Food Chemistry, 60:9352-9363.
• Wojciechowska, E., Weinert, C.H., Egert, B., Trierweiler, B., Schmidt-Heydt, M., Horneburg, B., GraeffHönninger, S., Kulling, S.E. and Geisen, R. 2014. Chlorogenic acid, a metabolite identified by untargeted metabolome analysis in resistant tomatoes, inhibits the colonization by Alternaria alternata by inhibiting alternariol biosynthesis. European Journal of Plant Pathology, 139: 735-747.
• Yamagishi, D., Akamatsu, H., Otani, H., and Kodama, M. 2006. Pathological evaluation of host-specific AALtoxins and fumonisin mycotoxins produced by Alternaria and Fusarium species. Journal of General Plant Pathology, 72: 323–326. doi: 10.1007/s10327-006-0291-y
• Yoder, O. 1980. Toxins in pathogenesis. Annual Review of Phytopathology, 18: 103-129.
• Yu, W., Yu, F-Y., Undersander, D.J. and Chu, F.S. 1999. Immunoassays of selected mycotoxins in hay, silage and mixed feed. Food and Agricultural Immunology, 11: 307-319.
• Zhang, Y., Li, H., Zhang, J., Shao, B. 2019. Determination of Alternaria toxins in drinking water by ultraperformance liquid chromatography tandem mass spectrometry. Environmental Science and Pollution Research, 26:22485-22493.
• Zhao, X., Liu, D.L., Yang, Y., Zhang, L., Yang, M. 2022. Detection of seven Alternaria toxins in edible and medicinal herbs using ultra-high performance liquid chromatography-tandem mass spectrometry. Food Chemistry: X, 13:100186.
• Zur, G., Hallerman, E.M., Sharf, R., and Kashi, Y. 1999. Development of a polymerase chain reaction-based assay for the detection of Alternaria fungal contamination in food products. Journal of Food Protection, 62: 1191-1197.