酿造酱油中黄曲霉毒素B_1的产生及其控制研究
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摘要
作为传统的优势发酵调味品之一，酿造酱油在我国的产量和消费量都很高，是居民餐桌上必不可少的重要调味品，也是发酵食品产业的重要经济支柱。但由于现代环境污染的不断加剧及落后的传统生产技术，使发酵食品的安全性问题尤为突出，人们对酿造酱油的安全性也提出了质疑。酿造酱油从原料的种植、贮存、制曲到酱醅的发酵等过程都不可避免地会受到黄曲霉毒素B_1（AFB_1）的污染，这就使得酿造酱油的终端产品也有污染AFB_1的可能性，会对人类健康和经济发展造成严重危害。因此，为保障我国居民膳食的安全及发酵食品产业稳步快速的发展，对我国现阶段酿造酱油中AFB_1污染现状的调查及如何控制发酵食品中AFB_1污染的研究就显得尤为重要。本论文首先调查了酿造酱油中AFB_1的污染现状，并分析了其对我国人群健康造成的潜在风险程度；然后针对我国酿造酱油中AFB_1的污染问题，与实际生产相结合，分析引起酱油中AFB_1污染的原因，并在此基础上，筛选出能安全、经济、高效控制酿造酱油AFB_1污染的菌株；最后综合分析了该菌株对AFB_1污染的防治机制。具体研究内容如下：
1.采用直接竞争酶联免疫方法分析了我国陕西、山西、山东、辽宁、上海、浙江、广东、江苏8省市209份酿造酱油样品中AFB_1的污染，发现AFB_1在我国部分地区的酿造酱油产品中污染十分广泛，但污染水平较低，符合Normal模型，且未见超标样品，AFB_1平均值为0.40±0.42ng/mL。运用Monte Carlo技术建立了我国部分地区酿造酱油中AFB_1的人群暴露模型和人群风险模型，得出我国部分地区酿造酱油中AFB_1的人群暴露量范围为0~0.385ng/kg bw/day，平均值为0.0417±0.0457ng/kg bw/day，暴露分布符合Logistic模型；引起的人群风险范围为3.87E-6~6.87E-3例癌症/年/100000人，平均值为1.42E-3±1.57E-3例癌症/年/100000人，整个分布符合Logistic模型。总体来说，南方酿造酱油中AFB_1的污染水平、暴露量及人群风险都比北方高。
2.为了确定AFB_1的污染源，找到控制AFB_1的关键点，跟踪监测了酿造酱油一个生产周期中AFB_1及微生物的动态变化。发现酿造过程中AFB_1含量的变化可分为三个阶段：第一阶段从酿造原料到发酵的第4d，AFB_1含量（以干基计）呈下降趋势，并在发酵的第4d降到最低值；第二阶段从发酵的第4d到发酵的第12d，由于产毒菌开始产毒，使AFB_1含量显著增加，并在第12d时达到最大值；第三阶段从发酵的第12d到发酵结束，产毒菌停止生长，同时AFB_1发生生物降解，使AFB_1含量又逐渐下降。结果表明，酿造酱油中有50%的AFB_1来源于原料，17%的AFB_1产生于发酵前期，33%的AFB_1产生于发酵中期。从酱醅中筛选出7株既能抑制黄曲霉（Aspergillus flavus）3.4408产AFB_1，又能降解AFB_1的丝状真菌，其中以FS10的防治效果最好，抑制率和降解率分别为80.33%和57.44%，通过菌株形态学鉴定和18S rDNA分析，确定该菌株为黑曲霉（Aspergillus niger）。
3.将黑曲霉（Aspergillus niger）FS10及其发酵产物与黄曲霉（Aspergillus flavus）3.4408共培养，研究FS10及其发酵产物对黄曲霉生长能力、产孢子能力、产AFB_1能力的影响。结果发现FS10发酵产物能抑制aflR和nor-1产毒基因的表达，并且对黄曲霉生长能力、产孢子能力、产AFB_1能力的抑制效果明显高于FS10，最大抑制率分别40.97%、100%和94.48%，其对黄曲霉3.4408作用的MIC和MFC分别为80μL/mL和320μL/mL，且FS10发酵产物中具有抑菌活性的物质是一种耐热的蛋白质。运用光学显微镜、原子力显微镜和扫描电镜分析FS10发酵产物对黄曲霉孢子萌发、孢子及菌丝体超微结构的影响，发现FS10发酵产物能显著抑制黄曲霉孢子的萌发，引起孢子收缩、塌陷，降低孢子表面的粗糙度，破坏细胞壁结构，导致细胞死亡；同时使黄曲霉菌丝体发生扭曲，收缩变形，严重破坏细胞壁的结构，阻止菌丝体的生长。
4.研究了FS10及其发酵产物对AFB_1的降解效果，结果表明FS10对AFB_1的降解是一种酶促降解。且AFB_1的降解酶是一种不耐热的胞外酶，用80%饱和度的硫酸铵盐析时，粗酶液的酶解作用最强。该酶的最适酶解pH为6.0，最适酶解温度为40℃。通过Ames实验评价了FS10的发酵产物及AFB_1降解产物的致突变性，发现FS10的发酵产物及AFB_1降解产物没有致突变作用。并对AFB_1降解产物进行了荧光光谱和LC-MS分析，发现AFB_1经降解酶作用后荧光强度明显减弱，且产生了分子量为286的物质M。因此，推测AFB_1降解产物的降解途径为：降解酶通过水解作用打开了AFB_1的内酯环结构，生成具有β-酮酸结构的中间产物，然后进一步发生脱羧反应，生成了M，使得AFB_1的毒性大大减弱。
As one of the traditional fermented condiments in China, the fermented soy saucehas high output and consumption. It is not only an indispensable condiment on thetable of Chinese residents, but also an important economic pillar of the fermented foodindustry. However, due to the growing environmental pollution and backwardproduction technology, the food safety issue of fermented food is outstandingly, thesafety of fermented soy sauce is also questioned by people. From the cultivation andstorage of raw materials to koji and the fermentation process of paste, it is inevitably tobe contaminated by aflatoxin B_1(AFB_1). Consequently, the fermented soy sauce islikely to be contaminated with AFB_1, which results in serious harm to human healthand economic development. Therefore, in order to protect the dietary safety of Chinesepeople and the steady and rapid development of fermented food industry, it isparticularly important to survey the AFB_1comtamination in fermnented soy sauce andcontrol AFB_1contamination. In this study, the AFB_1contamination status in fermentedsoy sauce was investigated and its potential risk to people health was analyzed. Then,with the actual production process, the reasons which cause AFB_1contamination infermented soy sauce were analyzed. Based on the results, the strain that could safely,economic, efficient control AFB_1contamination was isolated. Finally, the mechanismof biocontrol AFB_1was comprehensively analyzed and the major studies are asfollows:
1. The contamination of AFB_1in209fermented soy sauce samples came from8provinces of Shannxi, Shanxi, Shandong, Liaoning, Jiangsu, Zhejiang, Shanghai andGuangdong was detected by direct competitive enzyme-linked immunosorbent(ELISA). It was found that AFB_1contamination in fermented soy sauce in China iswidespread with lower level. The concentration of all samples was below the Chineseregulation (5ng/mL) with the mean of0.40±0.42ng/mL and fits to the Normaldistribution model. Based on the data of soy sauce intake and the value of hepatitis Bsurface antigen-positive rate (HBsAg+), the exposure and population risk models ofAFB_1through the fermented soy sauce were estimated by probabilistic approach usingthe Monte Carlo method. Results showed that the exposure is from0to0.177ng/kgbw/day with the mean of0.0417±0.0457ng/kg bw/day and fits to the Logisticdistribution model, the population risk is from3.87E-6to6.87E-3cancers/year/100,000people with the mean of1.42E-3±1.57E-3cancers/year/100,000people and fits to the Logistic distribution model. On the whole, there is high level ofAFB_1contamination, dietary intake and potential risk for cancer in south samples.
2. To determine the source of AFB_1in fermented soy sauce and find the critical control point, the dynamic changes of AFB_1concentration and microbiology weremonitored during the process. Changes in AFB_1content is divided into three stages: thefirst stage is from raw materials to the4day of fermentation, AFB_1content (dry basis)decreased and got the minimum value. The second stage is from the4day to12day offermentation, toxigenic fungi began to produceAFB_1. So the AFB_1content increasedsignificantly, and reached a maximum value in the12day of fermentation. The thirdstage is from the fermentation of the12day to the end of fermentation, toxigenic fungistoped growing and AFB_1was degraded, so the content of AFB_1was decreasedgradually. Results revealed that50%of AFB_1is caused by raw materials,17%of AFB_1and33%of AFB_1is produced in early and middle fermentation period, respectively.Seven strains were isolated from fermented paste, which can inhibit AFB_1biosynthesisby Aspergillus flavus3.4408and degrade AFB_1simultaneously. For the seven strains,FS10has the strongest ability to control AFB_1contaminaton with80.33%of inhibitionand57.44%of degradation, respectively. So it was selected for further study. FS10wasidentified as Aspergillus niger by morphology and18S rDNA sequence.
3. To elucidate the biocontrol mechanism, the ability of Aspergillus niger FS10andits fermentation products against growth, sporulation and AFB_1production bytoxigenic Aspergillus flavus3.4408was evaluated in vitro. Results showed that FS10fermentation products could restrain the expression of aflR and nor-1, and its ability toinhibit mycelia growth of Aspergillus flavus, sporulation and AFB_1production weresignificantly higher than FS10, with the maximal inhibition of40.97%、100%and94.48%, respectively. The MIC and MFC of FS10fermentation products againstAspergillus flavus were80μL/mL and320μL/mL, respectively. Results indicated thatthe active substance is a heat-resistant protein. In addition, the effect of FS10fermentation products to spore germination, ultrastructure monphology of spore andmycelium were also studied by optical microscopy, atomic force microscopy (AFM)and scanning electron microscopy (SEM). Result showed that FS10fermentationproducts could significantly inhibit spore germination. Morphology of spores wasserious damaged accompanying with spore analosis and cell wall collapse, resulted ininhibition of spore germination. Forthermore, it made mycelium of Aspergillus flavusto distort, shrink; then the cell wall structure was serious damaged to prevent thegrowth of mycelium.
4. The degradation of AFB_1by FS10and its fermentation products was studied.Results showed that AFB_1degradation is an enzymatic degradation by extracellularenzyme with poor thermal stability produced by FS10. When FS10fermentationproducts were salted by80%saturation of ammonium sulfate, the degradation activitywas strongest. The optimum enzymatic pH and temperature are6.0and40℃, respectively. The mutagenicity of FS10fermentation products and AFB_1degradationproducts were analyzed by Ames test. The results indicated that the fermentationproducts and degradation products did not have the ability of mutagenicity. In addition,the fluorescence spectrometry and LC-MS of AFB_1degradation product was studied.Results showed that fluorescence intensity of AFB_1was significantly reduced by thedegradation enzyme, and a substance with molecular weight of286was formed. Theprobable degradation pathway as follow: lactone ring structure of AFB_1was opened bythe degradation enzyme; then, an intermediate product with β-keto structure wasformed; finally, M was formed after decarboxylation reaction, cause greatly reductionof AFB_1toxicity.
1.采用直接竞争酶联免疫方法分析了我国陕西、山西、山东、辽宁、上海、浙江、广东、江苏8省市209份酿造酱油样品中AFB_1的污染，发现AFB_1在我国部分地区的酿造酱油产品中污染十分广泛，但污染水平较低，符合Normal模型，且未见超标样品，AFB_1平均值为0.40±0.42ng/mL。运用Monte Carlo技术建立了我国部分地区酿造酱油中AFB_1的人群暴露模型和人群风险模型，得出我国部分地区酿造酱油中AFB_1的人群暴露量范围为0~0.385ng/kg bw/day，平均值为0.0417±0.0457ng/kg bw/day，暴露分布符合Logistic模型；引起的人群风险范围为3.87E-6~6.87E-3例癌症/年/100000人，平均值为1.42E-3±1.57E-3例癌症/年/100000人，整个分布符合Logistic模型。总体来说，南方酿造酱油中AFB_1的污染水平、暴露量及人群风险都比北方高。
2.为了确定AFB_1的污染源，找到控制AFB_1的关键点，跟踪监测了酿造酱油一个生产周期中AFB_1及微生物的动态变化。发现酿造过程中AFB_1含量的变化可分为三个阶段：第一阶段从酿造原料到发酵的第4d，AFB_1含量（以干基计）呈下降趋势，并在发酵的第4d降到最低值；第二阶段从发酵的第4d到发酵的第12d，由于产毒菌开始产毒，使AFB_1含量显著增加，并在第12d时达到最大值；第三阶段从发酵的第12d到发酵结束，产毒菌停止生长，同时AFB_1发生生物降解，使AFB_1含量又逐渐下降。结果表明，酿造酱油中有50%的AFB_1来源于原料，17%的AFB_1产生于发酵前期，33%的AFB_1产生于发酵中期。从酱醅中筛选出7株既能抑制黄曲霉（Aspergillus flavus）3.4408产AFB_1，又能降解AFB_1的丝状真菌，其中以FS10的防治效果最好，抑制率和降解率分别为80.33%和57.44%，通过菌株形态学鉴定和18S rDNA分析，确定该菌株为黑曲霉（Aspergillus niger）。
3.将黑曲霉（Aspergillus niger）FS10及其发酵产物与黄曲霉（Aspergillus flavus）3.4408共培养，研究FS10及其发酵产物对黄曲霉生长能力、产孢子能力、产AFB_1能力的影响。结果发现FS10发酵产物能抑制aflR和nor-1产毒基因的表达，并且对黄曲霉生长能力、产孢子能力、产AFB_1能力的抑制效果明显高于FS10，最大抑制率分别40.97%、100%和94.48%，其对黄曲霉3.4408作用的MIC和MFC分别为80μL/mL和320μL/mL，且FS10发酵产物中具有抑菌活性的物质是一种耐热的蛋白质。运用光学显微镜、原子力显微镜和扫描电镜分析FS10发酵产物对黄曲霉孢子萌发、孢子及菌丝体超微结构的影响，发现FS10发酵产物能显著抑制黄曲霉孢子的萌发，引起孢子收缩、塌陷，降低孢子表面的粗糙度，破坏细胞壁结构，导致细胞死亡；同时使黄曲霉菌丝体发生扭曲，收缩变形，严重破坏细胞壁的结构，阻止菌丝体的生长。
4.研究了FS10及其发酵产物对AFB_1的降解效果，结果表明FS10对AFB_1的降解是一种酶促降解。且AFB_1的降解酶是一种不耐热的胞外酶，用80%饱和度的硫酸铵盐析时，粗酶液的酶解作用最强。该酶的最适酶解pH为6.0，最适酶解温度为40℃。通过Ames实验评价了FS10的发酵产物及AFB_1降解产物的致突变性，发现FS10的发酵产物及AFB_1降解产物没有致突变作用。并对AFB_1降解产物进行了荧光光谱和LC-MS分析，发现AFB_1经降解酶作用后荧光强度明显减弱，且产生了分子量为286的物质M。因此，推测AFB_1降解产物的降解途径为：降解酶通过水解作用打开了AFB_1的内酯环结构，生成具有β-酮酸结构的中间产物，然后进一步发生脱羧反应，生成了M，使得AFB_1的毒性大大减弱。
As one of the traditional fermented condiments in China, the fermented soy saucehas high output and consumption. It is not only an indispensable condiment on thetable of Chinese residents, but also an important economic pillar of the fermented foodindustry. However, due to the growing environmental pollution and backwardproduction technology, the food safety issue of fermented food is outstandingly, thesafety of fermented soy sauce is also questioned by people. From the cultivation andstorage of raw materials to koji and the fermentation process of paste, it is inevitably tobe contaminated by aflatoxin B_1(AFB_1). Consequently, the fermented soy sauce islikely to be contaminated with AFB_1, which results in serious harm to human healthand economic development. Therefore, in order to protect the dietary safety of Chinesepeople and the steady and rapid development of fermented food industry, it isparticularly important to survey the AFB_1comtamination in fermnented soy sauce andcontrol AFB_1contamination. In this study, the AFB_1contamination status in fermentedsoy sauce was investigated and its potential risk to people health was analyzed. Then,with the actual production process, the reasons which cause AFB_1contamination infermented soy sauce were analyzed. Based on the results, the strain that could safely,economic, efficient control AFB_1contamination was isolated. Finally, the mechanismof biocontrol AFB_1was comprehensively analyzed and the major studies are asfollows:
1. The contamination of AFB_1in209fermented soy sauce samples came from8provinces of Shannxi, Shanxi, Shandong, Liaoning, Jiangsu, Zhejiang, Shanghai andGuangdong was detected by direct competitive enzyme-linked immunosorbent(ELISA). It was found that AFB_1contamination in fermented soy sauce in China iswidespread with lower level. The concentration of all samples was below the Chineseregulation (5ng/mL) with the mean of0.40±0.42ng/mL and fits to the Normaldistribution model. Based on the data of soy sauce intake and the value of hepatitis Bsurface antigen-positive rate (HBsAg+), the exposure and population risk models ofAFB_1through the fermented soy sauce were estimated by probabilistic approach usingthe Monte Carlo method. Results showed that the exposure is from0to0.177ng/kgbw/day with the mean of0.0417±0.0457ng/kg bw/day and fits to the Logisticdistribution model, the population risk is from3.87E-6to6.87E-3cancers/year/100,000people with the mean of1.42E-3±1.57E-3cancers/year/100,000people and fits to the Logistic distribution model. On the whole, there is high level ofAFB_1contamination, dietary intake and potential risk for cancer in south samples.
2. To determine the source of AFB_1in fermented soy sauce and find the critical control point, the dynamic changes of AFB_1concentration and microbiology weremonitored during the process. Changes in AFB_1content is divided into three stages: thefirst stage is from raw materials to the4day of fermentation, AFB_1content (dry basis)decreased and got the minimum value. The second stage is from the4day to12day offermentation, toxigenic fungi began to produceAFB_1. So the AFB_1content increasedsignificantly, and reached a maximum value in the12day of fermentation. The thirdstage is from the fermentation of the12day to the end of fermentation, toxigenic fungistoped growing and AFB_1was degraded, so the content of AFB_1was decreasedgradually. Results revealed that50%of AFB_1is caused by raw materials,17%of AFB_1and33%of AFB_1is produced in early and middle fermentation period, respectively.Seven strains were isolated from fermented paste, which can inhibit AFB_1biosynthesisby Aspergillus flavus3.4408and degrade AFB_1simultaneously. For the seven strains,FS10has the strongest ability to control AFB_1contaminaton with80.33%of inhibitionand57.44%of degradation, respectively. So it was selected for further study. FS10wasidentified as Aspergillus niger by morphology and18S rDNA sequence.
3. To elucidate the biocontrol mechanism, the ability of Aspergillus niger FS10andits fermentation products against growth, sporulation and AFB_1production bytoxigenic Aspergillus flavus3.4408was evaluated in vitro. Results showed that FS10fermentation products could restrain the expression of aflR and nor-1, and its ability toinhibit mycelia growth of Aspergillus flavus, sporulation and AFB_1production weresignificantly higher than FS10, with the maximal inhibition of40.97%、100%and94.48%, respectively. The MIC and MFC of FS10fermentation products againstAspergillus flavus were80μL/mL and320μL/mL, respectively. Results indicated thatthe active substance is a heat-resistant protein. In addition, the effect of FS10fermentation products to spore germination, ultrastructure monphology of spore andmycelium were also studied by optical microscopy, atomic force microscopy (AFM)and scanning electron microscopy (SEM). Result showed that FS10fermentationproducts could significantly inhibit spore germination. Morphology of spores wasserious damaged accompanying with spore analosis and cell wall collapse, resulted ininhibition of spore germination. Forthermore, it made mycelium of Aspergillus flavusto distort, shrink; then the cell wall structure was serious damaged to prevent thegrowth of mycelium.
4. The degradation of AFB_1by FS10and its fermentation products was studied.Results showed that AFB_1degradation is an enzymatic degradation by extracellularenzyme with poor thermal stability produced by FS10. When FS10fermentationproducts were salted by80%saturation of ammonium sulfate, the degradation activitywas strongest. The optimum enzymatic pH and temperature are6.0and40℃, respectively. The mutagenicity of FS10fermentation products and AFB_1degradationproducts were analyzed by Ames test. The results indicated that the fermentationproducts and degradation products did not have the ability of mutagenicity. In addition,the fluorescence spectrometry and LC-MS of AFB_1degradation product was studied.Results showed that fluorescence intensity of AFB_1was significantly reduced by thedegradation enzyme, and a substance with molecular weight of286was formed. Theprobable degradation pathway as follow: lactone ring structure of AFB_1was opened bythe degradation enzyme; then, an intermediate product with β-keto structure wasformed; finally, M was formed after decarboxylation reaction, cause greatly reductionof AFB_1toxicity.
引文
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