Browse > Article
http://dx.doi.org/10.5941/MYCO.2017.45.4.240

Influence of Temperature and Water Activity on Deleterious Fungi and Mycotoxin Production during Grain Storage  

Mannaa, Mohamed (Laboratory of Plant Disease and Biocontrol, Department of Biosystems and Biotechnology, Korea University)
Kim, Ki Deok (Laboratory of Plant Disease and Biocontrol, Department of Biosystems and Biotechnology, Korea University)
Publication Information
Mycobiology / v.45, no.4, 2017 , pp. 240-254 More about this Journal
Abstract
Cereal grains are the most important food source for humans. As the global population continues to grow exponentially, the need for the enhanced yield and minimal loss of agricultural crops, mainly cereal grains, is increasing. In general, harvested grains are stored for specific time periods to guarantee their continuous supply throughout the year. During storage, economic losses due to reduction in quality and quantity of grains can become very significant. Grain loss is usually the result of its deterioration due to fungal contamination that can occur from preharvest to postharvest stages. The deleterious fungi can be classified based on predominance at different stages of crop growth and harvest that are affected by environmental factors such as water activity ($a_w$) and eco-physiological requirements. These fungi include species such as those belonging to the genera Aspergillus and Penicillium that can produce mycotoxins harmful to animals and humans. The grain type and condition, environment, and biological factors can also influence the occurrence and predominance of mycotoxigenic fungi in stored grains. The main environmental factors influencing grain fungi and mycotoxins are temperature and $a_w$. This review discusses the effects of temperature and $a_w$ on fungal growth and mycotoxin production in stored grains. The focus is on the occurrence and optimum and minimum growth requirements for grain fungi and mycotoxin production. The environmental influence on aflatoxin production and hypothesized mechanisms of its molecular suppression in response to environmental changes are also discussed. In addition, the use of controlled or modified atmosphere as an environmentally safe alternative to harmful agricultural chemicals is discussed and recommended future research issues are highlighted.
Keywords
Aflatoxin; Aflatoxin biosynthesis gene cluster; Mycotoxin; Storage fungi; Temperature; Water activity;
Citations & Related Records
Times Cited By KSCI : 7  (Citation Analysis)
연도 인용수 순위
1 Magan N, Lacey J. Ecological determinants of mould growth in stored grain. Int J Food Microbiol 1988;7:245-56.   DOI
2 Squire RA. Ranking animal carcinogens: a proposed regulatory approach. Science 1981;214:877-80.   DOI
3 World Health Organization. Some naturally occurring substances: food items and constituents, heterocyclic aromatic amines and mycotoxins. IARC Monographs on the Evaluation of Carcinogenic Risks to Humans. Vol. 56. Lyon: IARC Press; 1993. p. 397-444.
4 Groopman JD, Kensler TW. The light at the end of the tunnel for chemical-specific biomarkers: daylight or headlight? Carcinogenesis 1999;20:1-11.   DOI
5 Gong Y, Hounsa A, Egal S, Turner PC, Sutcliffe AE, Hall AJ, Cardwell K, Wild CP. Postweaning exposure to aflatoxin results in impaired child growth: a longitudinal study in Benin, West Africa. Environ Health Perspect 2004;112:1334-8.   DOI
6 Ehrlich KC, Kobbeman K, Montalbano BG, Cotty PJ. Aflatoxin-producing Aspergillus species from Thailand. Int J Food Microbiol 2007;114:153-9.   DOI
7 Pildain MB, Frisvad JC, Vaamonde G, Cabral D, Varga J, Samson RA. Two novel aflatoxin-producing Aspergillus species from Argentinean peanuts. Int J Syst Evol Microbiol 2008; 58(Pt 3):725-35.   DOI
8 Scheidegger KA, Payne GA. Unlocking the secrets behind secondary metabolism: a review of Aspergillus flavus from pathogenicity to functional genomics. J Toxicol 2003;22:423-59.
9 Price MS, Yu J, Nierman WC, Kim HS, Pritchard B, Jacobus CA, Bhatnagar D, Cleveland TE, Payne GA. The aflatoxin pathway regulator AflR induces gene transcription inside and outside of the aflatoxin biosynthetic cluster. FEMS Microbiol Lett 2006;255:275-9.   DOI
10 Li F, Yoshizawa T. Alternaria mycotoxins in weathered wheat from China. J Agric Food Chem 2000;48:2920-4.   DOI
11 Meronuck RA, Steele JA, Mirocha CJ, Christensen CM. Tenuazonic acid, a toxin produced by Alternaria alternata. Appl Microbiol 1972;23:613-7.
12 Marsh SF, Payne GA. Scaning EM studies on the colonization of dent corn by Aspergillus flavus. Phytopathology 1984;74: 557-61.   DOI
13 Hill RA, Wilson DM, McMillian WW, Widstrom NW, Cole RJ, Sanders TH, Blankenship PD. Ecology of the Aspergillus flavus group and aflatoxin formation in maize and groundnut. In: Lacey J, editor. Trichothecenes and other mycotoxins. Chichester: Wiley & Sons; 1985. p. 79-95.
14 Magan N, Cayley GR, Lacey J. Effect of water activity and temperature on mycotoxin production by Alternaria alternata in culture and on wheat grain. Appl Environ Microbiol 1984; 47:1113-7.
15 Kosiak B, Torp M, Skjerve E, Andersen B. Alternaria and Fusarium in Norwegian grains of reduced quality: a matched pair sample study. Int J Food Microbiol 2004;93:51-62.   DOI
16 Azcarate MP, Patriarca A, Terminiello L, Fernandez Pinto V. Alternaria toxins in wheat during the 2004 to 2005 Argentinean harvest. J Food Prot 2008;71:1262-5.   DOI
17 Müller ME, Urban K, Köppen R, Siegel D, Korn U, Koch M. Mycotoxins as antagonistic or supporting agents in the interaction between phytopathogenic Fusarium and Alternaria fungi. World Mycotoxin J 2014;8:311-21.
18 Bezuidenhout SC, Gelderblom WC, Gorst-Allman CP, Horak RM, Marasas WF, Spiteller G, Vleggaar R. Structure elucidation of the fumonisins, mycotoxins from Fusarium moniliforme. J Chem Soc Chem Commun 1988;11:743-5.
19 Marasas WF, Riley RT, Hendricks KA, Stevens VL, Sadler TW, Gelineau-van Waes J, Missmer SA, Cabrera J, Torres O, Gelderblom WC, et al. Fumonisins disrupt sphingolipid metabolism, folate transport, and neural tube development in embryo culture and in vivo: a potential risk factor for human neural tube defects among populations consuming fumonisin-contaminated maize. J Nutr 2004;134:711-6.   DOI
20 Fleurat-Lessard F. Integrated management of the risks of stored grain spoilage by seedborne fungi and contamination by storage mould mycotoxins: an update. J Stored Prod Res 2017;71:22-40.   DOI
21 Lacey J. Pre-and post-harvest ecology of fungi causing spoilage of foods and other stored products. J Appl Microbiol 1989;67:11S-25S.
22 Flannigan B. Primary contamination of barley and wheat grain storage fungi. Trans Br Mycol Soc 1978;71:37-42.   DOI
23 Miller JD. Fungi and mycotoxins in grain: implications for stored product research. J Stored Prod Res 1995;31:1-16.   DOI
24 Oh JY, Sang MK, Oh JE, Lee HJ, Ryoo MI, Kim KD. Microbial population, aflatoxin contamination and predominant Aspergillus species in Korean stored rice. Plant Pathol J 2010;26:121-9.   DOI
25 Oh JY, Jee SN, Nam Y, Lee H, Ryoo MI, Kim KD. Populations of fungi and bacteria associated with samples of stored rice in Korea. Mycobiology 2007;35:36-8.   DOI
26 Oh JY, Kim EN, Ryoo MI, Kim KD. Morphological and molecular identification of Penicillium islandicum isolate KU101 from stored rice. Plant Pathol J 2008;24:469-73.   DOI
27 Oh JY, Sang MK, Lee H, Ryoo MI, Kim KD. First detection of Penicillium fellutanum from stored rice in Korea. Res Plant Dis 2011;17:216-21.   DOI
28 Wilson DM, Mubatanhema W, Jurjevic Z. Biology and ecology of mycotoxigenic Aspergillus species as related to economic and health concerns. Adv Exp Med Biol 2002; 504:3-17.
29 Sanchis V, Magan N. Environmental conditions affecting mycotoxins. In: Magan N, Olsen M, editors. Mycotoxins in food: detection and control. Cambridge: Woodhead Publishing Ltd.; 2004. p. 174-89.
30 Hocking AD, Pitt JI. Water relations of some Penicillium species at $25{\circ}C$. Trans Br Mycol Soc 1979;73:141-5.   DOI
31 Northolt MD, Van Egmond HP, Paulsch WE. Differences between Aspergillus flavus strains in growth and aflatoxin $B_{1}$ production in relation to water activity and temperature. J Food Prot 1977;40:778-81.   DOI
32 Klich MA. Environmental and developmental factors influencing aflatoxin production by Aspergillus flavus and Aspergillus parasiticus. Mycoscience 2007;48:71-80.   DOI
33 Abdel-Hadi A, Schmidt-Heydt M, Parra R, Geisen R, Magan N. A systems approach to model the relationship between aflatoxin gene cluster expression, environmental factors, growth and toxin production by Aspergillus flavus. J R Soc Interface 2012;9:757-67.   DOI
34 Mousa W, Ghazali FM, Jinap S, Ghazali HM, Radu S. Modeling growth rate and assessing aflatoxins production by Aspergillus flavus as a function of water activity and temperature on polished and brown rice. J Food Sci 2013;78:M56-63.   DOI
35 Ogundero VW. Temperature and aflatoxin production by Aspergillus flavus and A. parasiticus strains from Nigerian groundnuts. J Basic Microbiol 1987;27:511-4.   DOI
36 Sedmikova M, Reisnerova H, Dufkova Z, Barta I, Jilek F. Potential hazard of simultaneous occurrence of aflatoxins $B_{1}$ and ochratoxin A. Vet Med 2001;46:169-74.   DOI
37 Kheiralla ZH, Hassanin NI, Amra H. Effect of incubation time, temperature and substrate on growth and aflatoxin production. Int Biodeterior Biodegradation 1992;30:17-27.   DOI
38 Medina A, Gilbert MK, Mack BM, OBrian GR, Rodriguez A, Bhatnagar D, Payne G, Magan N. Interactions between water activity and temperature on the Aspergillus flavus transcriptome and aflatoxin $B_{1}$ production. Int J Food Microbiol 2017; 256:36-44.   DOI
39 Yu J, Fedorova ND, Montalbano BG, Bhatnagar D, Cleveland TE, Bennett JW, Nierman WC. Tight control of mycotoxin biosynthesis gene expression in Aspergillus flavus by temperature as revealed by RNA-Seq. FEMS Microbiol Lett 2011;322:145-9.   DOI
40 Sanchis V, Abadias M, Oncins L, Sala N, Viñas I, Canela R. Occurrence of fumonisins $B_{1}$ and $B_{2}$ in corn-based products from the Spanish market. Appl Environ Microbiol 1994;60: 2147-8.
41 Frisvad JC, Frank JM, Houbraken JA, Kuijpers AF, Samson RA. New ochratoxin A producing species of Aspergillus section Circumdati. Stud Mycol 2004;50:23-43.
42 Lee HB, Magan N. Impact of environment and interspecific interactions between spoilage fungi and Aspergillus ochraceus on growth and ochratoxin production in maize grain. Int J Food Microbiol 2000;61:11-6.   DOI
43 Pardo E, Marin S, Sanchis V, Ramos AJ. Prediction of fungal growth and ochratoxin A production by Aspergillus ochraceus on irradiated barley grain as influenced by temperature and water activity. Int J Food Microbiol 2004;95:79-88.   DOI
44 Wearing AH, Burgess LW. Water potential and the saprophytic growth of Fusarium roseum "Graminearum." Soil Biol Biochem 1979;11:661-7.   DOI
45 Madhyastha S, Marquardt RR, Abramson D. Effect of ochratoxin producing fungi on the chemical composition of wheat and barley. J Food Qual 1993;16:287-99.   DOI
46 Häggblom P. Production of ochratoxin A in barley by Aspergillus ochraceus and Penicillium viridicatum: effect of fungal growth, time, temperature, and inoculum size. Appl Environ Microbiol 1982;43:1205-7.
47 Cairns-Fuller V, Aldred D, Magan N. Water, temperature and gas composition interactions affect growth and ochratoxin A production by isolates of Penicillium verrucosum on wheat grain. J Appl Microbiol 2005;99:1215-21.   DOI
48 Peraica M, Radic B, Lucic A, Pavlovic M. Toxic effects of mycotoxins in humans. Bull World Health Organ 1999;77: 754-66.
49 World Health Organization. Basic food safety for health workers [Internet]. Geneva: World Health Organization; 1999 [cited 2017 Jul 25]. Available from: http://apps.who.int/iris/bitstream/10665/65992/1/WHO_SDE_PHE_FOS_99.1.
50 Pitt JI. Penicillium toxins. In: Champ BR, Highley E, Hocking AD, Pitt JI, editors. Fungi and mycotoxins in stored products: Proceedings of an International Conference; 1991 Apr 23-26;Bangkok, Thailand. Canberra: Australian Centre for International Agricultural Research; 1991. p. 99-103.
51 Agrios G. Plant pathology. 5th ed. Boston (MA): Elsevier Academic Press; 2005.
52 Kushiro M. Historical review of researches on yellow rice and mycotoxigenic fungi adherent to rice in Japan. JSM Mycotoxins 2015;65:19-23.   DOI
53 Lutsky II, Mor N. Alimentary toxic aleukia (septic angina, endemic panmyelotoxicosis, alimentary hemorrhagic aleukia): t-2 toxin-induced intoxication of cats. Am J Pathol 1981;104: 189-91.
54 Azziz-Baumgartner E, Lindblade K, Gieseker K, Rogers HS, Kieszak S, Njapau H, Schleicher R, McCoy LF, Misore A, DeCock K, et al. Case-control study of an acute aflatoxicosis outbreak, Kenya, 2004. Environ Health Perspect 2005;113: 1779-83.   DOI
55 Krishnamachari KA, Bhat RV, Nagarajan V, Tilak TB. Hepatitis due to aflatoxicosis: an outbreak in western India. Lancet 1975;305:1061-3.   DOI
56 Wagacha JM, Muthomi JW. Mycotoxin problem in Africa: current status, implications to food safety and health and possible management strategies. Int J Food Microbiol 2008; 124:1-12.   DOI
57 Liu GT, Qian YZ, Zhang P, Dong ZM, Shi ZY, Zhen YZ, Miao J, Xu YM. Relationships between Alternaria alternata and oesophageal cancer. IARC Sci Publ 1991;(105):258-62.
58 Zhang F, Guo Z, Zhong H, Wang S, Yang W, Liu Y, Wang S. RNA-Seq-based transcriptome analysis of aflatoxigenic Aspergillus flavus in response to water activity. Toxins (Basel) 2014;6: 3187-207.   DOI
59 Faraj MK, Smith JE, Harran G. Interaction of water activity and temperature on aflatoxin production by Aspergillus flavus and A. parasiticus in irradiated maize seeds. Food Addit Contam 1991;8:731-6.   DOI
60 Cuero RG, Smith JE, Lacey J. Interaction of water activity, temperature and substrate on mycotoxin production by Aspergillus flavus, Penicillium viridicatum and Fusarium graminearum in irradiated grains. Trans Br Mycol Soc 1987; 89:221-6.   DOI
61 Georgianna DR, Payne GA. Genetic regulation of aflatoxin biosynthesis: from gene to genome. Fungal Genet Biol 2009; 46:113-25.   DOI
62 Abdel-Hadi A, Carter D, Magan N. Temporal monitoring of the nor-1 (aflD) gene of Aspergillus flavus in relation to aflatoxin B1 production during storage of peanuts under different water activity levels. J Appl Microbiol 2010;109:1914-22.   DOI
63 Yu J, Bhatnagar D, Cleveland TE. Completed sequence of aflatoxin pathway gene cluster in Aspergillus parasiticus. FEBS Lett 2004;564:126-30.   DOI
64 Nierman WC, Yu J, Fedorova-Abrams ND, Losada L, Cleveland TE, Bhatnagar D, Bennett JW, Dean R, Payne GA. Genome sequence of Aspergillus flavus NRRL 3357, a strain that causes aflatoxin contamination of food and feed. Genome Announc 2015;3:e00168-15.
65 O'Brian GR, Georgianna DR, Wilkinson JR, Yu J, Abbas HK, Bhatnagar D, Cleveland TE, Nierman W, Payne GA. The effect of elevated temperature on gene transcription and aflatoxin biosynthesis. Mycologia 2007;99:232-9.   DOI
66 Sutton JC, Baliko W, Funnell HS. Relation of weather variables to incidence of zearalenone in corn in southern Ontario. Can J Plant Sci 1980;60:149-55.   DOI
67 Hagler WM Jr, Towers NR, Mirocha CJ, Eppley RM, Bryden WL. Zearalenone: mycotoxin or mycoestrogen? In: Summerell BA, Leslie JF, Backhouse D, Bryden WL, Burgess LW, editors. Fursarium: Paul E. Nelson Memorial Symposium. St. Paul (MN): APS Press; 2001. p. 321-31.
68 Magan N, Lacey J. Water relations of some Fusarium species from infected wheat ears and grain. Trans Br Mycol Soc 1984;83:281-5.   DOI
69 Versonder RF, Ellis JJ, Kwolek WF, DeMarini DJ. Production of vomitoxin on corn by Fusarium graminearum NRRL 5883 and Fusarium roseum NRRL 6101. Appl Environ Microbiol 1982;43:967-70.
70 Eugenio CP, Christensen CM, Mirocha CJ. Factors affecting production of the mycotoxin F-2 by Fusarium roseum. Phytopathology 1970;60:1055-7.   DOI
71 Ryu D, Bullerman LB. Effect of cycling temperatures on the production of deoxynivalenol and zearalenone by Fusarium graminearum NRRL 5883. J Food Prot 1999;62:1451-5.   DOI
72 Hope R, Magan N. Two‐dimensional environmental profiles of growth, deoxynivalenol and nivalenol production by Fusarium culmorum on a wheat-based substrate. Lett Appl Microbiol 2003;37:70-4.   DOI
73 Jayas DS, Jeyamkondan S. PH-postharvest technology: modified atmosphere storage of grains meats fruits and vegetables. Biosyst Eng 2002;82:235-51.   DOI
74 Landers KE, Davis ND, Diener UL. Influence of atmospheric gases on aflatoxin production by Aspergillus flavus in peanuts. Phytopathology 1967;57:1086-90.
75 Flieger M, Wurst M, Shelby R. Ergot alkaloids: sources, structures and analytical methods. Folia Microbiol 1997;42:3-30.   DOI
76 Serna-Saldivar SO. Cereal grains: properties, processing, and nutritional attributes. Boca Raton (FL): CRC Press; 2016.
77 Walter GH, Chandrasekaran S, Collins PJ, Jagadeesan R, Mohankumar S, Alagusundaram K, Ebert PR, Daglish GJ, Nayak MK, Mohan S, et al. The grand challenge of food security: general lessons from a comprehensive approach to protecting stored grain from insect pests in Australia and India. Indian J Entomol 2016;78:7-16.   DOI
78 Food and Agriculture Organization of the United Nations. FAOSTAT statistics database [Internet]. Rome: FAO; 2017 [cited 2017 Jul 25]. Available from: http://www.fao.org/worldfoodsituation/csdb/en/.
79 Shih CN, Marth EH. Aflatoxin produced by Aspergillus parasiticus when incubated in the presence of different gases. J Milk Food Technol 1973;36:421-5.   DOI
80 Food and Agriculture Organization. Feeding the world, eradicating hunger: executive summary. World summit on food security [Internet]. Rome: FAO; 2009 [cited 2017 Jul 25]. Available from: http://www.fao.org/fileadmin/templates/wsfs/Summit/WSFS_Issues_papers/WSFS_Background_paper_Feeding_the_world.pdf.
81 Thompson C, Henke SE. Effects of climate and type of storage container on aflatoxin production in corn and its associated risks to wildlife species. J Wildl Dis 2000;36:172-9.   DOI
82 Magan N, Medina A, Aldred D. Possible climate-change effects on mycotoxin contamination of food crops pre- and postharvest. Plant Pathol 2011;60:150-63.   DOI
83 Paterson RR, Lima N. How will climate change affect mycotoxins in food? Food Res Int 2010;43:1902-14.   DOI
84 Paterson RR, Lima N. Further mycotoxin effects from climate change. Food Res Int 2011;44:2555-66.   DOI
85 Van Egmond HP. Worldwide regulations for mycotoxins. Adv Exp Med Biol 2002;504:257-69.
86 Henry SH, Bosch FX, Troxell TC, Bolger PM. Reducing liver cancer: global control of aflatoxin. Science 1999;286:2453-4.   DOI
87 Benford D, Boyle C, Dekant W, Fuchs R, Gaylor DW, Hard G, McGregory DB, Pitt JI, Plestina R, Shephard G, et al. Ochratoxin A. Safety evaluation of certain mycotoxins in food. WHO Food Additives Series 47. FAO Food and Nutrition Paper [Internet]. Geneva: World Health Organization IPCS; 2001 [cited 2017 Jul 25]. Available from: http://www.inchem.org/documents/jecfa/jecmono/v47je01.htm.
88 Shapira R, Paster N. Control of mycotoxins in storage and techniques for their decontamination. In: Magan N, Olsen M, editors. Mycotoxins in food: detection and control. Cambridge: Woodhead Publishing Ltd.; 2004. p. 190-223.
89 Schmidt-Heydt M, Abdel-Hadi A, Magan N, Geisen R. Complex regulation of the aflatoxin biosynthesis gene cluster of Aspergillus flavus in relation to various combinations of water activity and temperature. Int J Food Microbiol 2009; 135:231-7.   DOI
90 Pose G, Patriarca A, Kyanko V, Pardo A, Pinto VF. Effect of water activity and temperature on growth of Alternaria alternata on a synthetic tomato medium. Int J Food Mircobiol 2009; 135:60-3.   DOI
91 Schmidt-Heydt M, Rüfer CE, Abdel-Hadi A, Magan N, Geisen R. The production of aflatoxin $B_{1}$ or $G_{1}$ by Aspergillus parasiticus at various combinations of temperature and water activity is related to the ratio of aflS to aflR expression. Mycotoxin Res 2010;26:241-6.   DOI
92 Chang PK. The Aspergillus parasiticus protein AFLJ interacts with the aflatoxin pathway-specific regulator AFLR. Mol Genet Genomics 2003;268:711-9.
93 Gallo A, Solfrizzo M, Epifani F, Panzarini G, Perrone G. Effect of temperature and water activity on gene expression and aflatoxin biosynthesis in Aspergillus flavus on almond medium. Int J Food Microbiol 2016;217:162-9.   DOI
94 Bernaldez V, Cordoba JJ, Magan N, Peromingo B, Rodriguez A. The influence of ecophysiological factors on growth, aflR gene expression and aflatoxin $B_{1}$ production by a type strain of Aspergillus flavus. Food Sci Technol 2017;83:283-91.
95 Liu BH, Chu FS. Regulation of aflR and its product, AflR, associated with aflatoxin biosynthesis. Appl Environ Microbiol 1998;64:3718-23.
96 Chang PK. Lack of interaction between AFLR and AFLJ contributes to nonaflatoxigenicity of Aspergillus sojae. J Biotechnol 2004;107:245-53.   DOI
97 Yu J. Current understanding on aflatoxin biosynthesis and future perspective in reducing aflatoxin contamination. Toxins (Basel) 2012;4:1024-57.   DOI
98 Paster N, Barkai-Golan R, Calderon M. Control of T-2 toxin production using atmospheric gases. J Food Prot 1986;49:615-7.   DOI
99 Lillehoj EB, Milburn MS, Ciegler A. Control of Penicillium martensii development and penicillic acid production by atmospheric gases and temperatures. Appl Microbiol 1972; 24:198-201.
100 Paster N, Lisker N, Chet I. Ochratoxin A production by Aspergillus ochraceus Wilhelm grown under controlled atmospheres. Appl Environ Microbiol 1983;45:1136-9.
101 Paster N, Blumenthal-Yonassi J, Barkai-Golan R, Menasherov M. Production of zearalenone in vitro and in corn grains stored under modified atmospheres. Int J Food Microbiol 1991;12:157-65.   DOI
102 Samapundo S, De Meulenaer B, Atukwase A, Debevere J, Devlieghere F. The influence of modified atmospheres and their interaction with water activity on the radial growth and fumonisin $B_{1}$ production of Fusarium verticillioides and F. proliferatum on corn. Part I: The effect of initial headspace carbon dioxide concentration. Int J Food Microbiol 2007; 114:160-7.   DOI
103 Cole RJ, Cox RH. Handbook of toxic fungal metabolites. New York: Academic Press; 1981.
104 Magan N, Aldred D. Post-harvest control strategies: minimizing mycotoxins in the food chain. Int J Food Microbiol 2007;119:131-9.   DOI
105 Christensen CM, Kaufmann HH. Grain storage: the role of fungi in quality loss. Minneapolis (MN): University of Minnesota Press; 1969.
106 National Academy of Sciences. Post-harvest food losses in developing countries: a bibliography. Washington, D.C.: National Academy of Sciences; 1978.
107 Sinha RN. The stored-grain ecosystem. In: Jayas DS, White ND, Muir WE, editors. Stored-grain ecosystems. New York: Marcel Dekker; 1995. p. 1-32.
108 Levinson H, Levinson A. Control of stored food pests in the ancient orient and classical antiquity. J Appl Entomol 1998;122:137-44.   DOI
109 Mannaa M, Kim KD. Microbe-mediated control of mycotoxigenic grain fungi in stored rice with focus on aflatoxin biodegradation and biosynthesis inhibition. Mycobiology 2016;44:67-78.   DOI
110 Mannaa M, Oh JY, Kim KD. Microbe-mediated control of Aspergillus flavus in stored rice grains with a focus on aflatoxin inhibition and biodegradation. Ann Appl Biol 2017; 171:376-92.   DOI
111 Mannaa M, Oh JY, Kim KD. Biocontrol activity of volatileproducing Bacillus megaterium and Pseudomonas protegens against Aspergillus flavus and aflatoxin production on stored rice grains. Mycobiology 2017;45:213-9.   DOI
112 Magan N, Lacey J. Effect of temperature and pH on water relations of field and storage fungi. Trans Br Mycol Soc 1984;82:71-81.   DOI
113 Magan N, Lacey J. Effect of water activity, temperature and substrate on interactions between field and storage fungi. Trans Br Mycol Soc 1984;82:83-93.   DOI
114 Marin S, Sanchis V, Saenz R, Ramos AJ, Vinas I, Magan N. Ecological determinants for germination and growth of some Aspergillus and Penicillium spp. from maize grain. J Appl Microbiol 1998;84:25-36.   DOI
115 Young AB, Davis ND, Diener UL. Effect of temperature and moisture on tenuazonic acid production by Alternaria tenuissima. Phytopathology 1980;70:607-9.   DOI
116 Northolt MD, Bullerman LB. Prevention of mold growth and toxin production through control of environmental conditions. J Food Prot 1982;45:519-26.   DOI
117 Sorenson WG, Hesseltine CW, Shotwell OL. Effect of temperature on production of aflatoxin on rice by Aspergillus flavus. Mycopathol Mycol Appl 1967;33:49-55.   DOI
118 Ramos AJ, Labernia N, Marin S, Sanchis V, Magan N. Effect of water activity and temperature on growth and ochratoxin production by three strains of Aspergillus ochraceus on a barley extract medium and on barley grains. Int J Food Microbiol 1998;44:133-40.   DOI
119 Northolt MD, Van Egmond HP, Paulsch WE. Ochratoxin A production by some fungal species in relation to water activity and temperature. J Food Prot 1979;42:485-90.   DOI
120 Marin S, Magan N, Belli N, Ramos AJ, Canela R, Sanchis V. Two-dimensional profiles of fumonisin $B_{1}$ production by Fusarium moniliforme and Fusarium proliferatum in relation to environmental factors and potential for modelling toxin formation in maize grain. Int J Food Microbiol 1999;51:159-67.   DOI
121 Lee HB, Patriarca A, Magan N. Alternaria in food: ecophysiology, mycotoxin production and toxicology. Mycobiology 2015;43: 93-106.   DOI
122 Marin S, Sanchis V, Ramos AJ, Vinas I, Magan N. Environmental factors, in vitro interactions, and niche overlap between Fusarium moniliforme, F. proliferatum, and F. graminearum, Aspergillus and Penicillium species from maize grain. Mycol Res 1991;102:831-7.