氧脂素对赭曲霉孢子产生、赭曲霉毒素A合成及粮食受侵染程度的影响
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摘要
为明确氧脂素羟基十八碳二烯酸(hydroxyoctadecaenoic acid,HODE)对不同密度赭曲霉的孢子产生和赭曲霉毒素A(ochratoxin A,OTA)合成的作用,测定不同密度赭曲霉培养过程中9S-HODE、13S-HODE和OTA的产量,添加9S-HODE和13S-HODE后孢子和OTA产量,最后测定不同密度赭曲霉对粮食的侵染程度。结果显示,低接种密度的赭曲霉9S-HODE/13S-HODE比值及OTA产量更高;添加9S-HODE抑制赭曲霉孢子形成、促进OTA合成,13S-HODE作用相反;孢子接种密度为103个/mL和106个/mL的赭曲霉侵染花生、大豆、玉米及小麦后,发芽率分别降低29%、21%、17%、14%和35%、29%、20%、22%。这表明群体密度影响赭曲霉9S-HODE、13S-HODE和OTA产量,9S-HODE和13S-HODE可能是赭曲霉群体感应信号分子,二者在调节赭曲霉孢子产生和OTA合成上作用相反,高群体密度赭曲霉对粮食的侵染能力更强,脂肪及蛋白含量高的粮食可能更易被赭曲霉侵染。
This study was intended to clarify the effect of hydroxyoctadecaenoic acid(HODE) on the spore production and ochratoxin A(OTA) synthesis of Aspergillus ochraceus at different population densities. Here we measured the yields of9 S-HODE, 13 S-HODE and OTA during the culture process of A. ochraceus, and we also determined the production of spores and OTA after adding 9 S-hydroxy-10 E,12 Z-octadecadienoic acid(9 S-HODE) or 13 S-hydroxy-9 Z,11 E-octadecadienoic acid(13 S-HODE) and the infection degree of grain seeds with different densities of A. ochraceus. The results showed that the ratio of 9 S-HODE/13 S-HODE and the yield of OTA were higher at lower inoculation density(103 spores/mL); the addition of 9 S-HODE inhibited the sporulation of A. ochraceus and promoted OTA synthesis, while 13 S-HODE just had the opposite effect. After infection with A. ochraceus at initial spore density of 103 and 106 spores/mL, the germination rates of peanut,soybean, corn and wheat were reduced by 29%, 21%, 17% and 14%; and by 35%, 29%, 20% and 22%, respectively. These findings indicated that the population density of A. ochraceus could affect 9 S-HODE, 13 S-HODE and OTA production.9 S-HODE and 13 S-HODE may both be A. ochraceus quorum sensing signal molecules, which had opposite effects on A.ochraceus spore production and OTA synthesis. High population density of A. ochraceus was more infectious to grain seeds.Seeds with high levels of fat and protein content were more susceptible to A. ochraceus.
This study was intended to clarify the effect of hydroxyoctadecaenoic acid(HODE) on the spore production and ochratoxin A(OTA) synthesis of Aspergillus ochraceus at different population densities. Here we measured the yields of9 S-HODE, 13 S-HODE and OTA during the culture process of A. ochraceus, and we also determined the production of spores and OTA after adding 9 S-hydroxy-10 E,12 Z-octadecadienoic acid(9 S-HODE) or 13 S-hydroxy-9 Z,11 E-octadecadienoic acid(13 S-HODE) and the infection degree of grain seeds with different densities of A. ochraceus. The results showed that the ratio of 9 S-HODE/13 S-HODE and the yield of OTA were higher at lower inoculation density(103 spores/mL); the addition of 9 S-HODE inhibited the sporulation of A. ochraceus and promoted OTA synthesis, while 13 S-HODE just had the opposite effect. After infection with A. ochraceus at initial spore density of 103 and 106 spores/mL, the germination rates of peanut,soybean, corn and wheat were reduced by 29%, 21%, 17% and 14%; and by 35%, 29%, 20% and 22%, respectively. These findings indicated that the population density of A. ochraceus could affect 9 S-HODE, 13 S-HODE and OTA production.9 S-HODE and 13 S-HODE may both be A. ochraceus quorum sensing signal molecules, which had opposite effects on A.ochraceus spore production and OTA synthesis. High population density of A. ochraceus was more infectious to grain seeds.Seeds with high levels of fat and protein content were more susceptible to A. ochraceus.
引文
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[19]REVERBERI M,PUNELLI F,SCARPARI M,et al.Lipoperoxidation affects ochratoxin A biosynthesis in Aspergillus ochraceus,and its interaction with wheat seeds[J].Applied Microbiology&Biotechnology,2010,85(6):1935-1946.DOI:10.1007/s00253-009-2220-4.
[20]PUNELLI F,REVERBERI M,VELEZ H,et al.Characterization of lipoxygenase genes and oxylipin profile in A.ochraceus[C]//Journal of Plant Pathology Edizioni ETS.Peru餫,Italija,2007:56.
[21]ANTONIOLETTI R,VIGLIANTI A,CRISTOFOLI S,et al.Role of some neosynthesized styryl heterocycles in ochratoxin A control in Aspergillus carbonarius[J].Current Organic Chemistry,2016,20(19):2029-2035.DOI:10.2174/1385272820666160421150917.
[22]晏石娟,黄文洁,刘春明,等.脂肪酸及其氧合物对曲霉属真菌菌丝生长、产孢和黄曲霉毒素合成的影响[J].微生物学报,2017,57(1):24-32.DOI:10.13343/j.cnki.wsxb.20160245.
[23]李彩艳,梁志宏,黄昆仑.黄曲霉群体感应研究进展[J].食品安全质量检测学报,2015(8):3205-3210.
[24]娄庆任,黄家权,王后苗,等.脂氧合酶与作物黄曲霉毒素污染抗性关系研究进展[J].中国油料作物学报,2014,36(1):127-134.DOI:10.7505/j.issn.1007-9084.2014.01.020.
[25]WILSON R A,GARDNER H W,KELLER N P.Cultivar-dependent expression of a maize lipoxygenase responsive to seed infesting fungi[J].Molecular Plant-Microbe Interactions:MPMI,2001,14(8):980.DOI:10.1094/MPMI.2001.14.8.980.
[26]薛鑫,张芊,吴金霞.植物体内活性氧的研究及其在植物抗逆方面的应用[J].生物技术通报,2013(10):6-11.DOI:10.13560/j.cnki.biotech.bull.1985.2013.10.033.
[27]REVERBERI M,ZJALIC S,RICELLI A,et al.Modulation of antioxidant defense in Aspergillus parasiticus is involved in aflatoxin biosynthesis:a role for the ApyapA gene[J].Eukaryotic Cell,2008,7(6):988-1000.DOI:10.1128/EC.00228-07.
[28]PONTS N,PINSON-GADAIS L B M,BARREAU C,et al.Accumulation of deoxynivalenol and its 15-acetylated form is significantly modulated by oxidative stress in liquid cultures of Fusarium graminearum[J].FEMS Microbiology Letters,2006,258(1):102-107.DOI:10.1111/j.1574-6968.2006.00200.x.
[29]REVERBERI M,GAZZETTI K,PUNELLI F,et al.Aoyap1 regulates OTA synthesis by controlling cell redox balance in Aspergillus ochraceus[J].Applied Microbiology&Biotechnology,2012,95(5):1293-1304.DOI:10.1007/s00253-012-3985-4.
[2]OSTRY V,MALIR F,RUPRICH J.Producers and important dietary sources of ochratoxin A and citrinin[J].Toxins,2013,5(9):1574-1586.DOI:10.3390/toxins5091574.
[3]MALIR F,OSTRY V,NOVOTNA E.Toxicity of the mycotoxin ochratoxin A in the light of recent data[J].Toxin Reviews,2013,32(2):19-33.DOI:10.3109/15569543.2013.782504.
[4]张露瀛.赭曲霉毒素A的生物脱毒实例[J].农产品加工,2017(8):46-47.DOI:10.16693/j.cnki.1671-9646(X).2017.08.015.
[5]刘青,邹志飞,余炀炀,等.食品中真菌毒素法规限量标准概述[J].中国酿造,2017,36(1):12-18.DOI:10.11882/j.issn.0254-5071.2017.01.003.
[6]KüGLER S,SEBGHATI T S,EISSENBERG L G,et al.Phenotypic variation and intracellular parasitism by Histoplasma capsulatum[J].Proceedings of the National Academy of Sciences of the United States of America,2000,97(16):8794-8798.DOI:10.1073/pnas.97.16.8794.
[7]WILLIAMS H E,STEELE J C P,CLEMENTS M O,et al.γ-Heptalactone is an endogenously produced quorum-sensing molecule regulating growth and secondary metabolite production by Aspergillus nidulans[J].Applied Microbiology&Biotechnology,2012,96(3):773-781.DOI:10.1007/s00253-012-4065-5.
[8]BROWN S H,ZARNOWSKI R,SHARPEE W C,et al.Morphological transitions governed by density dependence and lipoxygenase activity in Aspergillus flavus[J].Applied&Environmental Microbiology,2008,74(18):5674-5685.DOI:10.1128/AEM.00565-08.
[9]KAI P,BASSLER B L.Quorum sensing signal-response systems in Gram-negative bacteria[J].Nature Reviews Microbiology,2016,14(9):576.DOI:10.1038/nrmicro.2016.89.
[10]李彩艳,梁志宏.细菌和真菌群体感应研究进展[J].延安大学学报(自然科学版),2016,35(2):39-44.DOI:10.13876/J.cnki.ydnse.2016.02.039.
[11]BUROW G B,GARDNER H W,KELLER N P.A peanut seed lipoxygenase responsive to Aspergillus colonization[J].Plant Molecular Biology,2000,42(5):689-701.DOI:10.1023/A:1006361305703.
[12]LI C,SONG Y,XIONG L,et al.Initial spore density has an influence on ochratoxin A content in Aspergillus ochraceus CGMCC 3.4412in PDB and its interaction with seeds[J].Toxins,2017,9(4):146.DOI:10.3390/toxins9040146.
[13]TSITSIGIANNIS D I,KELLER N P.Oxylipins as developmental and host-fungal communication signals[J].Trends in Microbiology,2007,15(3):109-118.DOI:10.1016/j.tim.2007.01.005.
[14]姜春姣,江芸,耿志明,等.亚油酸氧化产物--羟基十八碳二烯酸的研究进展[J].食品科学,2018,39(7):278-284.DOI:10.7506/spkx1002-6630-201807041.
[15]HELL K,MUTEGI C,FANDOHAN P,et al.Aflatoxin control and prevention strategies in maize for sub-Saharan Africa[J].Julius-KühnArchiv,2011,5(5):459-466.DOI:10.5897/AJMR10.009.
[16]YAN S,LIANG Y,ZHANG J,et al.Autoxidated linolenic acid inhibits aflatoxin biosynthesis in Aspergillus flavus,via oxylipin species[J].Fungal Genetics&Biology,2015,81(4):229-237.DOI:10.1016/j.fgb.2014.11.005.
[17]CALVO A M,HINZE L L,GARDNER H W,et al.Sporogenic effect of polyunsaturated fatty acids on development of Aspergillus spp.[J].Applied&Environmental Microbiology,1999,65(8):3668-3673.DOI:10.1089/oli.1.1999.9.359.
[18]徐博恒,谢林香,张燎原.真菌群体感应现象的研究进展[J].微生物学杂志,2 0 1 4,3 4(5):1 0 0-1 0 5.D O I:1 0.3 9 6 9/j.issn.1005-7021.2014.05.019.
[19]REVERBERI M,PUNELLI F,SCARPARI M,et al.Lipoperoxidation affects ochratoxin A biosynthesis in Aspergillus ochraceus,and its interaction with wheat seeds[J].Applied Microbiology&Biotechnology,2010,85(6):1935-1946.DOI:10.1007/s00253-009-2220-4.
[20]PUNELLI F,REVERBERI M,VELEZ H,et al.Characterization of lipoxygenase genes and oxylipin profile in A.ochraceus[C]//Journal of Plant Pathology Edizioni ETS.Peru餫,Italija,2007:56.
[21]ANTONIOLETTI R,VIGLIANTI A,CRISTOFOLI S,et al.Role of some neosynthesized styryl heterocycles in ochratoxin A control in Aspergillus carbonarius[J].Current Organic Chemistry,2016,20(19):2029-2035.DOI:10.2174/1385272820666160421150917.
[22]晏石娟,黄文洁,刘春明,等.脂肪酸及其氧合物对曲霉属真菌菌丝生长、产孢和黄曲霉毒素合成的影响[J].微生物学报,2017,57(1):24-32.DOI:10.13343/j.cnki.wsxb.20160245.
[23]李彩艳,梁志宏,黄昆仑.黄曲霉群体感应研究进展[J].食品安全质量检测学报,2015(8):3205-3210.
[24]娄庆任,黄家权,王后苗,等.脂氧合酶与作物黄曲霉毒素污染抗性关系研究进展[J].中国油料作物学报,2014,36(1):127-134.DOI:10.7505/j.issn.1007-9084.2014.01.020.
[25]WILSON R A,GARDNER H W,KELLER N P.Cultivar-dependent expression of a maize lipoxygenase responsive to seed infesting fungi[J].Molecular Plant-Microbe Interactions:MPMI,2001,14(8):980.DOI:10.1094/MPMI.2001.14.8.980.
[26]薛鑫,张芊,吴金霞.植物体内活性氧的研究及其在植物抗逆方面的应用[J].生物技术通报,2013(10):6-11.DOI:10.13560/j.cnki.biotech.bull.1985.2013.10.033.
[27]REVERBERI M,ZJALIC S,RICELLI A,et al.Modulation of antioxidant defense in Aspergillus parasiticus is involved in aflatoxin biosynthesis:a role for the ApyapA gene[J].Eukaryotic Cell,2008,7(6):988-1000.DOI:10.1128/EC.00228-07.
[28]PONTS N,PINSON-GADAIS L B M,BARREAU C,et al.Accumulation of deoxynivalenol and its 15-acetylated form is significantly modulated by oxidative stress in liquid cultures of Fusarium graminearum[J].FEMS Microbiology Letters,2006,258(1):102-107.DOI:10.1111/j.1574-6968.2006.00200.x.
[29]REVERBERI M,GAZZETTI K,PUNELLI F,et al.Aoyap1 regulates OTA synthesis by controlling cell redox balance in Aspergillus ochraceus[J].Applied Microbiology&Biotechnology,2012,95(5):1293-1304.DOI:10.1007/s00253-012-3985-4.