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Biodiversity of Aspergillus section Flavi in the United States: a review. 美国黄曲霉(Aspergillus Flavi)生物多样性综述。
Pub Date : 2007-10-01 Epub Date: 2007-08-24 DOI: 10.1080/02652030701510012
Bruce W Horn

Fungi belonging to Aspergillus section Flavi are of great economic importance in the United States due to their ability to produce toxic and carcinogenic aflatoxins in agricultural commodities. Development of control strategies against A. flavus and A. parasiticus, the major aflatoxin-producing species, is dependent upon a basic understanding of their diversity in agricultural ecosystems. This review summarizes our current knowledge of species and population diversity in the United States in relation to morphology, mycotoxin production and genetic characters. The high genetic diversity in populations of aflatoxigenic fungi is a reflection of their versatile habits in nature, which include saprotrophic colonization of plant debris in soil and parasitism of seeds and grain. Genetic variation within populations may originate from a cryptic sexual state. The advent of intensive monoculture agriculture not only increases population size but also may introduce positive selective pressure for aflatoxin production due to its link with pathogenicity in crops. Important goals in population research are to determine how section Flavi diversity in agricultural ecosystems is changing and to measure the direction of this evolution.

黄曲霉属黄曲霉科真菌在美国具有重要的经济意义,因为它们能够在农产品中产生有毒和致癌的黄曲霉毒素。黄曲霉和寄生黄曲霉是主要的黄曲霉毒素产生物种,其防治策略的制定取决于对其在农业生态系统中的多样性的基本认识。这篇综述总结了我们目前对美国物种和种群多样性的认识,包括形态、霉菌毒素产生和遗传特征。产黄曲霉毒素真菌种群的高度遗传多样性反映了它们在自然界的多种习性,包括在土壤中腐殖植物残骸和寄生种子和谷物。种群内的遗传变异可能源于一种隐蔽的性状态。集约化单一栽培农业的出现不仅增加了种群规模,而且由于黄曲霉毒素与作物致病性的联系,可能会给黄曲霉毒素的生产带来积极的选择压力。种群研究的重要目标是确定农业生态系统中黄花多样性的变化情况,并测量这种变化的方向。
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引用次数: 101
Aflatoxin formation and gene expression in response to carbon source media shift in Aspergillus parasiticus. 碳源介质转移对寄生曲霉黄曲霉毒素形成及基因表达的影响。
Pub Date : 2007-10-01 DOI: 10.1080/02652030701579454
J R Wilkinson, J Yu, H K Abbas, B E Scheffler, H S Kim, W C Nierman, D Bhatnagar, T E Cleveland

Aflatoxins are toxic and carcinogenic polyketide metabolites produced by fungal species, including Aspergillus flavus and A. parasiticus. The biosynthesis of aflatoxins is modulated by many environmental factors, including the availability of a carbon source. The gene expression profile of A. parasiticus was evaluated during a shift from a medium with low concentration of simple sugars, yeast extract (YE), to a similar medium with sucrose, yeast extract sucrose (YES). Gene expression and aflatoxins (B1, B2, G1, and G2) were quantified from fungal mycelia harvested pre- and post-shifting. When compared with YE media, YES caused temporary reduction of the aflatoxin levels detected at 3-h post-shifting and they remained low well past 12 h post-shift. Aflatoxin levels did not exceed the levels in YE until 24 h post-shift, at which time point a tenfold increase was observed over YE. Microarray analysis comparing the RNA samples from the 48-h YE culture to the YES samples identified a total of 2120 genes that were expressed across all experiments, including most of the aflatoxin biosynthesis genes. One-way analysis of variance (ANOVA) identified 56 genes that were expressed with significant variation across all time points. Three genes responsible for converting norsolorinic acid to averantin were identified among these significantly expressed genes. The potential involvement of these genes in the regulation of aflatoxin biosynthesis is discussed.

黄曲霉毒素是由黄曲霉和寄生蜂等真菌产生的有毒致癌多酮代谢产物。黄曲霉毒素的生物合成受到许多环境因素的调节,包括碳源的可用性。研究了从低浓度单糖酵母提取物(YE)培养基到含蔗糖酵母提取物蔗糖(YES)培养基中寄生蜂基因表达谱的变化。基因表达和黄曲霉毒素(B1, B2, G1和G2)在移植物前和移植物后采集的真菌菌丝中进行定量分析。与YE培养基相比,YES在移位后3小时检测到的黄曲霉毒素水平暂时降低,并且在移位后12小时仍保持较低水平。直到轮班后24小时,黄曲霉毒素水平才超过YE的水平,在这个时间点上,YE的黄曲霉毒素水平增加了10倍。微阵列分析比较了48小时YE培养液和YES培养液的RNA样本,共鉴定出2120个基因在所有实验中表达,包括大多数黄曲霉毒素生物合成基因。单因素方差分析(ANOVA)鉴定出56个基因在所有时间点上表达有显著差异。在这些显著表达的基因中,鉴定出三个负责将去盐酸转化为averantin的基因。讨论了这些基因在黄曲霉毒素生物合成调控中的潜在作用。
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引用次数: 38
Gene profiling for studying the mechanism of aflatoxin biosynthesis in Aspergillus flavus and A. parasiticus. 黄曲霉和寄生蜂黄曲霉毒素合成机制研究的基因谱分析。
Pub Date : 2007-10-01 DOI: 10.1080/02652030701513800
Jiujiang Yu, Catherine M Ronning, Jeffery R Wilkinson, Bruce C Campbell, Gary A Payne, Deepak Bhatnagar, Thomas E Cleveland, William C Nierman

Aflatoxins are toxic and carcinogenic polyketide metabolites produced by certain fungal species, including Aspergillus flavus and A. parasiticus. Many internal and external factors, such as nutrition and environment affect aflatoxin biosynthesis; therefore, we analyzed the transcriptome of A. flavus using expressed sequence tags (ESTs) from a normalized cDNA expression library constructed from mycelia harvested under several conditions. A total of 7218 unique ESTs were identified from 26,110 sequenced cDNA clones. Functional classifications were assigned to these ESTs and genes, potentially involved in the aflatoxin contamination process, were identified. Based on this EST sequence information, a genomic DNA amplicon microarray was constructed at The Institute for Genomic Research (TIGR). To identify potential regulatory networks controlling aflatoxin contamination in food and feeds, gene expression profiles in aflatoxin-supportive media versus non-aflatoxin-supportive media were evaluated in A. flavus and A. parasiticus. Genes consistently expressed in several aflatoxin-supportive media are reported.

黄曲霉毒素是由某些真菌种类产生的有毒和致癌的多酮代谢产物,包括黄曲霉和寄生蜂。黄曲霉毒素的生物合成受到营养、环境等多种内外部因素的影响;因此,我们利用在不同条件下收获的菌丝构建的标准化cDNA表达文库中的表达序列标签(est)分析了黄曲霉的转录组。从26,110个cDNA克隆中共鉴定出7218条独特的ESTs。对这些est进行了功能分类,并确定了可能参与黄曲霉毒素污染过程的基因。基于这些EST序列信息,在基因组研究所(TIGR)构建了基因组DNA扩增子微阵列。为了确定控制食品和饲料中黄曲霉毒素污染的潜在调控网络,我们在黄曲霉毒素支持培养基和非黄曲霉毒素支持培养基中对黄曲霉和寄生蜂的基因表达谱进行了评估。在几种黄曲霉毒素支持介质中一致表达的基因有报道。
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引用次数: 24
Analysis of aflatoxin regulatory factors in serial transfer-induced non-aflatoxigenic Aspergillus parasiticus. 系列转移诱导的非产黄曲霉寄生的黄曲霉毒素调控因子分析。
Pub Date : 2007-10-01 DOI: 10.1080/02652030701564563
S P Kale, J W Cary, N Hollis, J R Wilkinson, D Bhatnagar, J Yu, T E Cleveland, J W Bennett

Aflatoxins (AFs) are carcinogenic secondary metabolites of Aspergillus parasiticus. In previous studies, non-toxigenic A. parasiticus sec- (for secondary metabolism negative) variants were generated through serial transfer of mycelia from their toxigenic sec+ (for secondary metabolism positive) parents for genetic and physiological analysis for understanding regulation of AF biosynthesis. Previous studies have shown no difference in the DNA sequence of aflR, a positive regulator of AF production, in the sec+ and sec- strains. In this study, AflJ, another positive regulator of AF production, laeA, a global regulator of secondary metabolism, and the intergenic region between aflR and aflJ, were analysed to determine if they play a role in establishment of the sec- phenotype. The study showed that while this sequence identity extended to the aflJ as well as the aflJ-aflR intergenic region, expression of aflR in the sec- strain was several fold lower than that observed in the sec+ strain, while aflJ expression was barely detectable in the sec- strain. Western blot analysis indicated that despite AflR protein being present in the sec- strain, no toxin production resulted. Introduction of a second copy of aflR into the sec- strain increased aflR expression, but did not restore AF production. Lastly, reverse transcription-PCR analysis revealed that laeA was expressed in both sec+ and sec- strains. These results suggest that although aflR, aflJ and laeA are necessary for AF production, they are not sufficient. We propose that the aflR and aflJ expression may be regulated by element(s) downstream from laeA or from pathways not influenced by laeA.

黄曲霉毒素(AFs)是寄生曲霉的致癌次生代谢产物。在以往的研究中,寄生蜂的非产毒性sec-(次级代谢阴性)变异是通过产毒性sec+(次级代谢阳性)亲本的菌丝连续转移产生的,用于遗传和生理分析,以了解AF生物合成的调控。先前的研究表明,在sec+和sec-菌株中,aflR的DNA序列没有差异,aflR是AF产生的正调节因子。本研究分析了AF的另一个正调节因子AflJ、次级代谢的全局调节因子laeA以及aflR和AflJ之间的基因间区域,以确定它们是否在sec表型的建立中起作用。研究表明,虽然该序列的同一性扩展到aflJ和aflJ-aflR基因间区,但sec-株中aflR的表达量比sec+株低几倍,而aflJ在sec-株中几乎没有表达。Western blot分析表明,该菌株虽然含有AflR蛋白,但不产生毒素。在第二株中引入第二拷贝aflR增加了aflR的表达,但没有恢复AF的产生。反转录pcr分析表明,laeA在sec+和sec-菌株中均有表达。这些结果表明,虽然aflR、aflJ和laeA是AF产生的必要条件,但它们并不充分。我们认为,aflR和aflJ的表达可能受到laeA下游的元件或不受laeA影响的通路的调控。
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引用次数: 18
Functional characterization, sequence comparisons and distribution of a polyketide synthase gene required for perithecial pigmentation in some Fusarium species. 一些镰刀菌种表皮色素沉着所需的聚酮合成酶基因的功能特征、序列比较和分布。
Pub Date : 2007-10-01 DOI: 10.1080/02652030701546495
R H Proctor, R A E Butchko, D W Brown, A Moretti

Polyketides are a structurally diverse class of secondary metabolites produced by bacteria, fungi, plants and animals. The fungal genus Fusarium includes agronomically important plant pathogenic and mycotoxin-producing species and produces numerous polyketides. The study further characterized a polyketide synthase-encoding gene (PKS3 = PGL1) that was previously identified in F. graminearum and F. verticillioides. Disruption of the F. verticillioides PGL1 indicated that it is required for the production of the dark pigment in perithecial walls, as previously shown in F. graminearum. A third PGL1 orthologue was identified in the genomic sequence of N. haematococca (anamorph F. solani f. sp. pisi). Analysis of the carboxy-terminal end of the deduced PGL1 protein indicated that it had a functional domain related to dehydrogenases/reductases that is sometimes present in non-ribosomal peptide synthetases. Comparison of the genomic regions flanking PGL1 in F. graminearum, F. verticillioides and N. haematococca revealed that the extent of gene synteny in this region was greater between F. graminearum and F. verticillioides than between either of these species and N. haematococca. Southern blot analysis indicated that PGL1 occurs widely within the genus Fusarium including species with no known sexual stage.

聚酮是一类结构多样的次级代谢产物,由细菌、真菌、植物和动物产生。镰刀菌属真菌包括农艺学上重要的植物致病性和产生霉菌毒素的物种,并产生许多聚酮。该研究进一步鉴定了先前在F. graminearum和F. verticillioides中发现的聚酮合成酶编码基因(PKS3 = PGL1)。verticillioides PGL1的破坏表明,正如之前在F. graminearum中所显示的那样,PGL1是生产周壁黑色色素所必需的。在赤球奈索菌(anamorph F. solani F. sp. pisi)基因组序列中鉴定出第三个PGL1同源基因。对推断出的PGL1蛋白的羧基末端的分析表明,它具有与脱氢酶/还原酶相关的功能域,这种功能域有时存在于非核糖体肽合成酶中。通过对小麦赤霉病菌、黄萎病菌和红葡萄球菌PGL1侧翼基因组区域的比较,发现小麦赤霉病菌和黄萎病菌的PGL1侧翼基因同源程度大于赤霉病菌和红葡萄球菌。Southern blot分析表明,PGL1广泛存在于镰刀菌属中,包括不知道有性期的物种。
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引用次数: 44
Wheat kernel black point and fumonisin contamination by Fusarium proliferatum. 小麦籽粒黑点与富马菌素污染。
Pub Date : 2007-10-01 DOI: 10.1080/02652030701513834
A E Desjardins, M Busman, R H Proctor, R Stessman

Fusarium proliferatum is a major cause of maize ear rot and fumonisin contamination and also can cause wheat kernel black point disease. The primary objective of this study was to determine whether nine F. proliferatum strains from wheat from Nepal can cause black point and fumonisin contamination in wheat kernels. For comparison, the study included three Fusarium strains from US maize. In test 1, all the strains but one produced significant symptoms of kernel black point; two strains decreased kernel yield; and four strains contaminated kernels with fumonisins B(1), B(2) and B(3) as determined by liquid chromatography-mass spectroscopy. Strain Ggm202 from Nepal, which produced the highest levels of fumonisins (mean = 49 microg g(-1)) on five wheat cultivars in test 1, was confirmed to produce fumonisins (mean = 38 microg g(-1)) on two cultivars in test 2. The data indicate a potential for fumonisin contamination of wheat infected with F. proliferatum.

增殖镰刀菌是玉米穗腐病和伏马菌素污染的主要原因,也可引起小麦籽粒黑点病。本研究的主要目的是确定来自尼泊尔小麦的9株增芽孢杆菌是否能引起小麦籽粒中的黑点和伏马菌素污染。为了进行比较,该研究包括了来自美国玉米的三种镰刀菌菌株。在试验1中,除1株外,其余菌株均出现了显著的籽粒黑点症状;2株籽粒产量下降;用液相色谱-质谱联用法测定了4株感染伏马菌素B(1)、B(2)和B(3)的菌株。来自尼泊尔的菌株Ggm202在试验1中对5个小麦品种产生最高水平的伏马菌素(平均为49微克(-1)),在试验2中被证实对2个小麦品种产生伏马菌素(平均为38微克(-1))。这些数据表明,富马菌素污染小麦感染了增芽孢杆菌。
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引用次数: 66
Rapid polymerase chain reaction (PCR)-single-stranded conformational polymorphism (SSCP) screening method for the identification of Aspergillus section Nigri species by the detection of calmodulin nucleotide variations. 快速聚合酶链反应(PCR)-单链构象多态性(SSCP)筛选法检测钙调素核苷酸变异对黑曲霉种进行鉴定。
Pub Date : 2007-10-01 DOI: 10.1080/02652030701551834
A Susca, G Stea, G Perrone

Single-stranded conformational polymorphism (SSCP) analysis for genetic diversity studies has been widely applied to detect indirectly sequence differences up to a single base in amplified DNA fragments of the same length, representing an alternative to gene sequencing. In this study SSCP analysis was used to detect sequence variations contained in an about 180-bp region of the calmodulin gene in order to identify Aspergillus section Nigri species. The method described shows that fluorescence-based SSCP analysis by capillary electrophoresis is cheaper and faster than direct sequencing, and suitable for computer-assisted analyses allowing discrimination between the Aspergillus species belonging to the Nigri section: A. aculeatus, Aspergillus 'atypic uniseriate', A. brasiliensis, A. carbonarius, A. ellipticus, A. foetidus, A. heteromorphus, A. ibericus, A. japonicus, A. niger, and A. tubingensis.

单链构象多态性(SSCP)分析已广泛应用于遗传多样性研究,用于间接检测相同长度的扩增DNA片段中单个碱基的序列差异,是基因测序的一种替代方法。本研究采用SSCP分析法检测钙调蛋白基因约180 bp区域的序列变异,以鉴定Aspergillus section Nigri种。所描述的方法表明,基于荧光的毛细管电泳SSCP分析比直接测序更便宜、更快,并且适用于计算机辅助分析,可以区分属于黑曲霉组的曲霉种:A. aculeatus、A. carbonarius、A. ellipticus、A. foetidus、A. heteromorphus、A. ibericus、A. japonicus、A. niger和A. tubingensis。
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引用次数: 20
Immunochemical methods for rapid mycotoxin detection: evolution from single to multiple analyte screening: a review. 快速检测真菌毒素的免疫化学方法:从单一到多种分析物筛选的演变:综述。
Pub Date : 2007-10-01 DOI: 10.1080/02652030701557179
I Y Goryacheva, S De Saeger, S A Eremin, C Van Peteghem

This review focuses on recent developments in immunochemical methods for detection of mycotoxins, with a particular emphasis on simultaneous multiple analyte determination. This includes high-throughput instrumental analysis for the laboratory environment (microtitre plate enzyme-linked immunoabsorbant assay (ELISA), different kinds of immunosensors, fluorescence polarization immunoassay, and capillary electrophoretic immunoassay), as well as rapid visual tests for on-site testing (lateral-flow, dipstick, flow-through and column tests). For each type of immunoassay, perspectives for multiple analyte application are discussed and examples cited.

本文综述了真菌毒素免疫化学检测方法的最新进展,特别强调了同时检测多种分析物的方法。这包括用于实验室环境的高通量仪器分析(微滴板酶联免疫吸附测定法(ELISA)、不同种类的免疫传感器、荧光极化免疫测定法和毛细管电泳免疫测定法),以及用于现场测试的快速视觉测试(横向流动、试纸、流动和柱测试)。对于每种类型的免疫分析,多分析物应用的前景进行了讨论,并列举了例子。
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引用次数: 143
Application of a liquid chromatography-tandem mass spectrometric method to multi-mycotoxin determination in raw cereals and evaluation of matrix effects. 液相色谱-串联质谱法在谷物原料中多种霉菌毒素测定及基质效应评价中的应用。
Pub Date : 2007-10-01 DOI: 10.1080/02652030701510004
Michael Sulyok, Rudolf Krska, Rainer Schuhmacher

A multi-analyte method for the liquid chromatography-tandem mass spectrometric determination of mycotoxins in crude grain extracts without clean-up has been applied to the analysis of spelt, rice and barley. Method performance characteristics were determined after spiking blank samples at multiple levels and were found to be comparable for all investigated matrices as regards linearity (linear calibration functions were obtained for all analyte/matrix combinations except for moniliformin), precision (coefficient of variations <6%) and sensitivity. Matrix-induced signal suppression/enhancement was studied in detail and varied significantly between the investigated matrices, as well as between individual samples (relative standard deviation was as high as 40% within three rice varieties) and individual toxins. It was concluded that a reliable quantitative analysis using matrix-matched calibration requires careful consideration of the model matrix, which should match the investigated samples as close as possible.

建立了一种无需清洗的液相色谱-串联质谱法测定粗谷物提取物中真菌毒素的方法,并应用于小麦、大米和大麦的分析。在多个水平的空白样品中添加峰后确定了方法的性能特征,并发现所有被调查的矩阵在线性(除moniliformin外,所有分析物/矩阵组合都获得了线性校准函数),精度(变异系数)方面具有可比性
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引用次数: 93
Apyap1 affects aflatoxin biosynthesis during Aspergillus parasiticus growth in maize seeds. Apyap1对玉米种子曲霉寄生生长过程中黄曲霉毒素合成的影响。
Pub Date : 2007-10-01 DOI: 10.1080/02652030701553244
M Reverberi, S Zjalic, F Punelli, A Ricelli, A A Fabbri, C Fanelli

It is demonstrated that, in fungal cells grown in synthetic media, the Apyap1 gene is implicated in the modulation of aflatoxin biosynthesis following the perturbation of redox balance. This study suggests that an association between oxidative stress and aflatoxin biosynthesis also occurs in maize seeds. We used DeltaApyap1, a strain in which the gene Apyap1 was disrupted, to verify whether this oxidative stress-related transcription factor, by affecting cell redox balance, can have a role in the modulation of aflatoxin synthesis. The amount of hydroperoxides (ROOH) produced by wild type (WT) and DeltaApyap1, both grown in potato dextrose broth, was assayed in the filtrate. In maize seeds (30 g), inoculated with WT and DeltaApyap1conidia and incubated at 30 degrees C for 15 days, lipoxygenase activity (LOX), lipoperoxides (LOOH) production, fungal growth and aflatoxin biosynthesis was analysed. It was observed that DeltaApyap1 released more hydroperoxides in the culture media and more aflatoxins in seeds, possibly through stronger stimulation of LOX, which, in turn led to greater LOOH production in the seeds. On the basis of the results, a hypothesis regarding strategies to control aflatoxin synthesis is formulated.

研究表明,在合成培养基中生长的真菌细胞中,Apyap1基因与氧化还原平衡被扰乱后黄曲霉毒素生物合成的调节有关。这项研究表明,氧化应激与黄曲霉毒素生物合成之间的关联也发生在玉米种子中。我们使用Apyap1基因被破坏的菌株DeltaApyap1来验证这种与氧化应激相关的转录因子是否可以通过影响细胞氧化还原平衡来调节黄曲霉毒素的合成。对野生型(WT)和DeltaApyap1在马铃薯葡萄糖肉汤中培养的滤液中产生的氢过氧化物(ROOH)进行了测定。在玉米种子(30 g)中,接种WT和deltaapyap1分生孢子,在30℃下培养15天,分析了脂氧合酶活性(LOX)、脂过氧化物(LOOH)的产生、真菌生长和黄曲霉毒素的生物合成。结果表明,DeltaApyap1在培养基中释放了更多的氢过氧化物,在种子中释放了更多的黄曲霉毒素,这可能是通过更强的LOX刺激,从而导致种子中产生更多的LOOH。在此基础上,提出了控制黄曲霉毒素合成策略的设想。
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引用次数: 52
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