摘要
近年来,内生真菌已成为寻找具有生物活性成分的另一重要来源。内生真菌以其丰富的生物多样性和次生代谢物多样性以及在宿主植物-食草动物-环境生态系统中扮演的重要角色而引起众多研究者的兴趣,并已成为当今农业和医药卫生领域新的研究热点。除虫菊(Pyrethrum cinerariifolium Trev.)是世界上著名的杀虫植物,其中主要杀虫活性物质除虫菊素还具有一定的杀菌作用,特别是对真菌的效果明显。植物内生真菌对植物某些药效成分的形成有重要影响。研究表明,内生真菌能产生与宿主相同或相似的具有生理活性的次生代谢产物,且真菌易于培养,可通过育种手段和控制发酵条件等措施来大幅提高其药用成分的含量。因此,利用植物内生真菌发酵生产生物源农药是突破某些植物资源生长周期长、不可再生等限制因素的一条新途径。
而到目前为止,有关除虫菊内生真菌的研究还鲜有报道。本研究以除虫菊植物为材料,从其根、茎、叶和花四部分组织中共分离得到128株内生真菌,并对所分离的内生真菌进行了形态学鉴定和抑菌活性筛选,对其中活性较高的Y1和Y2菌株从其生物学特性、代谢产物的抑菌活性、抑菌活性成分的分离与鉴定等方面进行了较为系统的研究,得出以下主要研究结果及结论:
1.较为系统地研究了除虫菊内生真菌的分布特点。发现除虫菊内生真菌具有较丰富的物种多样性,从除虫菊根、茎、叶、花等组织中共获得内生真菌128株,经形态观察,鉴定出其中的117株,分属于2个亚门、3个纲、7个目、7个科、12个属,分别为镰孢菌属Fusarium、链格孢属Alternaria、腐殖霉属Humicola、内多隔孢属Endophragmiella、黑孢霉属Nigrospora、色串孢属Torula、阜孢霉属Papularia、丝核菌属Rhizoctonia、茎点霉属Phoma、拟盘多毛孢属Pestalotiopsis、毛集座霉属Setodochium和毛壳菌属Chaetomium。117株内生真菌从纲的分类水平上以丝孢纲(Hyphomycetes)为优势类群,占60.9 %,腔孢纲(Coelomycetes)次之,占21.1 %;从目的分类水平上以从梗孢目(Hyphomycetales)和瘤座孢目(Tuberculariales)为优势类群,分别总菌株数的26.6 %和20.3 %;在属的分类水平上以镰孢菌属、腐殖霉属和链格孢属为优势属,分别占总菌株数的20.3 %,13.3 %和13.3 %。这些真菌除了毛壳菌属于子囊菌纲以外,多数属于半知菌亚门真菌。除虫菊内生真菌主要存在于植株的叶和根中,茎部组织中分离到的内生真菌相对较少。镰孢菌属(Fusarium)和腐殖霉属(Humicola)分别是根部和叶部的优势类群。
2.采用抑制菌丝生长速率法测定了128株内生真菌发酵滤液对番茄灰霉病菌、辣椒疫霉病菌、苹果炭疽病菌等6种植物病原真菌的抑制作用,结果表明,在离体条件下,活性菌株(对至少一种供试病原真菌菌丝生长抑制率达到75 %以上的菌株)有56株;其中Y1、Y2菌株发酵液的抑菌活性较好,其稀释10倍的发酵液对6种病原菌菌丝生长的抑制率均较高。上述56株内生真菌菌丝丙酮提取物对6种病原真菌的离体生测结果表明, Y1、Y2、H2三株菌株的菌丝丙酮提取物对6种病原菌均有较好的抑制作用。Y1、Y2菌株发酵滤液5倍稀释液对番茄灰霉病菌和苹果炭疽病菌孢子萌发的抑制作用均在80 %以上;盆栽试验结果表明,Y1、Y2、Y7、H2四株菌株对番茄灰霉病、辣椒疫霉病、黄瓜霜霉病的室内防效在50 %以上,值得进一步深入研究。
3.根据真菌的形态学鉴定方法,对比相关分类系统,并结合分子生物学方法将Y1菌株鉴定为小孢拟盘多毛孢菌(Pestalotiopsis microspora),将Y2菌株鉴定为尖孢镰刀菌(Fusarium oxysporum),其5.8s rDNA序列在Gene Bank上的登录号分别为EU638329和EU152473。
4.初步明确了Y1和Y2菌株的生物学特性。试验结果表明,内生真菌Y1、Y2菌株的培养特性不同,环境和营养因子对该菌的生长繁殖和抑菌活性影响较大。对于Y1菌株:(1)在10～35℃时,菌丝能生长并形成孢子,较适生长繁殖温度为30℃,孢子的致死温度为65℃;(2)在pH 6.5～8.0时,菌丝生长、孢子形成和萌发均良好,较适培养pH值为7.5;(3)能直接利用玉米粉和糊精;(4)氮源以有机态氮为佳,硝态氮对其生长繁殖有抑制作用;(5) Y1菌株以黑暗培养为佳。
Y2菌株菌丝生长在20～30℃时较为适宜,最适温度为25℃;分生孢子在4℃～40℃之间均能萌发,且30℃时萌发率最高; 30℃时产孢量最高;60℃处理10 min能完全杀死菌丝体和孢子。Y2菌株对pH值的适应范围很宽,其菌丝在pH 4.0～9.5时均能生长;pH5.5时相对产孢量达到最大;其孢子萌发率在pH5.0最大。之后,随着pH的增加,孢子萌发率逐渐下降。Y2菌株以光照/黑暗交替培养为佳。Y2菌株可以利用多种碳源和氮源生长并产孢,营养来源广泛。但就其产孢情况而言,有机氮源要优于无机氮源,由于供试无机氮源种类较少,此现象是否具有普遍性还要更深入研究。
5.初步研究了Y1、Y2菌株的发酵条件,明确了较佳培养基组成及培养条件。响应面分析方法试验结果表明,Y1菌株较佳培养条件为:以每升培养基中含有D-木糖19.86 g,蛋白胨2.91 g,MgSO4·7H2O 0.305 g,FeSO4 0.0061 g, K2HPO4 1.219 g,KCl 0.610 g为宜,摇瓶发酵较佳条件为:初始pH 7.64,装液量68.81 mL·250 mL~(-1),摇床转速220 r·min~(-1),培养温度26.7℃,接种量9.95 %。优化条件下菌丝体生长量和对番茄灰霉病菌菌丝生长抑菌作用分别达到446.5 mg·100mL~(-1)和95.8 %,较原始培养条件分别增加了约6.99 %和2.46%。
Y2菌株较佳培养条件为:以每升培养基中含有麦芽糖10 g、蛋白胨6 g、K2HPO4 2 g、KCl 1 g、FeSO4 0.02 g、MgSO4?7H2O 1 g为宜;摇瓶发酵较佳条件为:初始pH 7.0,装液量100 mL·250 mL~(-1),摇床转速180 r·min~(-1),培养温度28.0℃,接种量21 %。与初始条件相比,优化条件下菌丝生长量和对番茄灰霉病菌菌丝生长抑制作用分别提高了8.5 %和4.7 %。本研究结果为两株内生真菌的进一步研究提供了有意义的基础参数。
6.以番茄灰霉病菌为示踪生物,对Y1和Y2菌株的次生代谢产物进行了抑菌活性成分的分离,共分离得到13个化合物,其中P2为新化合物。从Y1菌株中新分离出6个化合物,其中2个化合物结构已确定,为3-亚甲基O-β-D-吡喃葡萄糖基-1,2-二酚(P2)和半乳糖醇(dulcitol,P3),1个化合物结构初步确定,为4,4'-N,N-二苯甲酸丙二醇酯(P1),其余3个化合物的结构待鉴定。生测结果表明,在0.25 mg·mL~(-1)供试浓度下,P1和P3对番茄灰霉病菌孢子萌发的抑制率在50 %以上;从Y2菌株中新分离出7个化合物,其中2个结构已确定,分别二十二烷(docosane,F1)和半乳糖醇(dulcitol,F5),2个化合物结构初步确定,分别为4-甲基苯腈(4-methoxybenzonitrile,F3),正十三烷酸乙二醇单酯(2-hydroxyethyl tridecanoate,F2);其余3个化合物结构待鉴定。生测结果表明,在1 mg·mL~(-1)供试浓度下,F2、F3、F4对番茄灰霉病均具有一定的保护作用,7 d室内药效分别达到60 %以上。对Y1、Y2菌株代谢产物中抑菌活性成分的分离与鉴定尚待进一步完成。
An endophyte is a bacterial (in cluding actinomycete) or fungal microorganism, which“colonizes symptomlessly the living, internal tissues of host plant, even though the endophyte may, after an incubation or latency period, cause disease.”Due to the specialized living niches, little knowledge of these“special organisms”has been accumulated. However, much renewed attention is now being paid to the biodiversity, chemistry and bioactivity of functional metabolites and the related physiological and/or ecological role of these microorganisms. Pyrethrum cinerariifolium Trev, a famous traditional Chinese medicinal herb know to be the producer of Pyrethrum, was found to be a widespread insecticide and have antifungal activity. In this paper, we investigated the distribution, bioactivity, biological and chemical characteristics of the representative fungal endophytes colonized inside the leaves of P. cinerariifolium. The results and some points of renovation are discussed as follows:
1. Systemically studied the fungal endophyte distribution and biodiversity of P.cinerariifolium for the first time. A total of 128 fungal were isolated from roots, stems leaves and flowers of P.cinerariifolium and 117 fungal strains were identified after morphology observation.. Among them, these strains were classified into 12 genera: Fusarium, Chaetomium, Phoma, Alternaria, Endophragmiella, Pestalotiopsis, Humicola, Nigrospora, Toruloa, Setodochium, Papularia and Rhizoctonia which showed higher diversity of endophytic fungi in P.cinerariifolium. It was found that the amount, species and distribution of endophytic fungi varied in different parts of P. cinerariifolium.
2. The bioassay activities of extracellular metabolic products from 128 strains endophytic fungi were tested. Antagonistic tests showed that the inhibition rates of the 56 strains were more than 75% against at least one pathogen and among them fermentation of Y1, Y2 have better antagonistic action against six pathogens (Botrytis cinerea, Phytophthora capsic, Colletotrichum gloeosporioides, Exserohilum turcicum, Rhizoctonia cerealis and Fusarium graminearum) at the concentration 10 times of fermentation . Among 56 strains, acetone extraction of Y1, Y2 and H2 showed inhibition activities against the above 6 pathogens. Fermentation of strains Y1, Y2, show inhibition rates higher than 80% towards germination of Botrytis cinerea and Apple anthracnose at the concentration 5 times of fermentation. Results of pot culture experiment in greenhouse indicate that strains Y1, Y2, Y7, H2 showed antifungal activities more than 50% against Botrytis cinerea, Phytophthora capsici and Cucumber downy mildew. The results indicated that the strains of Y1, Y2, H2 and Y7 were broad-spectrum and stronger antifungal activity.
3. The complete 5.8S rDNA sequence of Y1 and Y2 strain were cloned and sequenced respectively, and phylogenetic tree based on 5.8S rDNA sequences and analyses sequence homology were constructed. The characteristics of morphology showed that strain Y1 was identified as Pestalotiopsis sp., the analysis of DNA sequences showed that the sequence homology of Y1 strain was 100% with Pestalotiopsis microspora, and the 5.8s rDNA sequences number was EU638329 on Gene Bank。
The characteristics of morphology showed that the macroconidium was falciform with five or seven diaphragm; microconidium has zero or one diaphragm. The analysis of DNA sequences showed that the sequence homology of Y2 strain was 100% with Fusarium oxysporum. The characteristics of morphology and the analysis of DNA sequences suggest that the strain of Y2 was identified as Fusarium oxysporum, and the 5.8s rDNA sequences number was EU152473 on Gene Bank。
4.The biology characteristics of the two strains are preliminary defined. Results show that the effect of environmental and nutritional factors on its growth and reproduction was remarkable. To strain Y1 :ⅰ) Temperature from 10℃to 35℃was suitable for mycelial growth, germination and sporulations. The optimum temperature was 30℃for sporulation, and the lethal temperature was 65℃when treated for 10 minutes.ⅱ) pH value ranged from 6.5 to 8.0 was favorable for mycelial growth, sporulation and spore germination, and the optimum pH was 7.5.ⅲ) The best carbon sources for mycelial growth and sporulations were corn flour and dextrin.ⅳ) As to nitrogen sources, the fungus utilized organic nitrogen very well, while NO3–N (nitrate) inhibited its growth and sporulations. To strain Y2:ⅰ) Temperature from 20℃to 33℃were suitable for mycelial growth, sporulations and germination. The optimum temperature was 30℃for sporulation, and the lethal temperature was 60℃treated for 10 minutes.ⅱ) pH value ranged from 4.0 to 9.5 was favorable for mycelial growth, sporulation and spore germination. The optimum pH was 5.5.ⅲ) The best carbon sources for mycelial growth and sporulations were corn flour or dextrin.ⅳ) As to nitrogen sources, the fungus utilized organic nitrogen very well.
5. The fermentation conditions of strains Y1 and Y2 were studied and better components of culture media and conditions were determined. The response surface methodology indicated that basic culture media of strainY1 were D-xylose 19.86 g·L-1, peptone 2.91 g·L-1, MgSO4?7H2O 0.305 g·L-1,FeSO4 0.0061 g·L-1 , K2HPO4 1.219 g·L-1 ,KCl 0.610 g·L-1and optimal incubation conditions of initial pH , medium volume in flask, shaking speed, temperature and inoculation amount were 7.64, 68.81 mL·250 mL~(-1), 220 r·min~(-1), 26.7℃and 9.95%, respectively. Under the optimal condition, the biomass weight and antibiotic activity were enhanced by 6.99% and 2.46% respectively compared with that under the initial condition.
The single factor experiment indicated that basic culture media of Y2 were maltose 10 g, peptone 6 g, KH2PO4 2 g, KCl 1 g, FeSO4 0.02 g, MgSO4?7H2O 1 g every liter; and optimal incubation conditions of initial pH , medium volume in flask ,shaking speed , temperature and inoculation amount were7.0 , 100 mL·250 mL~(-1), 180 r·min~(-1) , 28.0℃and 21% , respectively. Under the optimal condition, the biomass weight and antibiotic activity were 897.5 mg·1000 mL~(-1) and 96.6%, and were enhanced by 8.5% and 4.7% respectively compared with that under the initial condition.
6. In the process of isolation of intracellular metabolic components, bioactivity is tested using Botrytis cinerea to detect antifungal and antibacterial activities, respectively. Thirteen antifungal chemical components are isolated from intracellular metabolic products of Y1 and Y2. Six chemical components are newly isolated from intracellular metabolic products of Y1, the chemical structure of one of them (P2 and P3) is clarified, which is dulcitol and 3-methylbenzene-1,2-diol- dextrose indicant (new chemical component), and the chemical structure of P1 would be Benzoic acid 4,4’-N,N-cyclic propylene ester.The bioassay results indicate that these two compounds inhibit spore of Botrytis cinerea germination, and at 0.25 mg·mL~(-1), the inhibitory rates of P1, P2 and P3 against bourgeon of the spore of Botrytis cinerea are more than 50%. Seven antifungal chemical components are isolated from metabolic products of Y2, the chemical structure of the four (F1, F2, F3, F5) is clearly explained, which may be 4-methoxybenzonitrile,2-hydroxyethyl tridecanoate,docosane and dulcitol. The bioassay results indicate that the inhibitory rates of F2, F3and F4 against Botrytis cinerea are more than 60% at 1 mg·mL~(-1) after 7d. Hence, isolation and identification of antifungal active components of Y1 and Y2 need to be studied further.
引文
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