一个编码富含丝氨酸蛋白的基因影响大丽轮枝菌的微菌核形成、产孢及致病力

 大丽轮枝菌是一种可在广泛的寄主范围内引发黄萎病的土传植物病原真菌,主要以微菌核的形式在土壤中存活多年,因此鉴定与微菌核形成及致病力相关的基因对防治该病害至关重要。本课题组从前期构建的棉花黄萎病菌T-DNA插入突变体库中筛选到一个微菌核明显减少的突变体,致病力测定结果表明,该突变体致病力明显下降。以棉花黄萎病菌野生型菌株V592的基因组DNA为模板,从棉花黄萎病菌中克隆到被T-DNA插入突变的基因(VdSRP1)编码区全长为415 bp,包含一个外显子,编码一个富含丝氨酸蛋白,与任何已知的注释基因没有显著的序列相似性。为了明确VdSRP1基因在大丽轮枝菌中的功能,利用同源重组的原理及农杆菌介导的遗传转化方法获得了VdSRP1基因的2个敲除体菌株,与棉花黄萎病菌野生型菌株V592相比,VdSRP1基因敲除突变体的微菌核形成明显减少,产孢量及孢子萌发率下降,对棉花的毒力也显著下降。对野生型菌株V592及VdSRP1敲除突变体的转录分析表明,VdSRP1调控一系列与微菌核形成、孢子形成及致病力相关基因的表达。这些结果表明,VdSRP1基因影响大丽轮枝菌的致病力、微菌核形成、产孢及孢子萌发。     

一个编码富含丝氨酸蛋白的基因影响大丽轮枝菌的微菌核形成、产孢及致病力

 大丽轮枝菌是一种可在广泛的寄主范围内引发黄萎病的土传植物病原真菌,主要以微菌核的形式在土壤中存活多年,因此鉴定与微菌核形成及致病力相关的基因对防治该病害至关重要。本课题组从前期构建的棉花黄萎病菌T-DNA插入突变体库中筛选到一个微菌核明显减少的突变体,致病力测定结果表明,该突变体致病力明显下降。以棉花黄萎病菌野生型菌株V592的基因组DNA为模板,从棉花黄萎病菌中克隆到被T-DNA插入突变的基因(VdSRP1)编码区全长为415 bp,包含一个外显子,编码一个富含丝氨酸蛋白,与任何已知的注释基因没有显著的序列相似性。为了明确VdSRP1基因在大丽轮枝菌中的功能,利用同源重组的原理及农杆菌介导的遗传转化方法获得了VdSRP1基因的2个敲除体菌株,与棉花黄萎病菌野生型菌株V592相比,VdSRP1基因敲除突变体的微菌核形成明显减少,产孢量及孢子萌发率下降,对棉花的毒力也显著下降。对野生型菌株V592及VdSRP1敲除突变体的转录分析表明,VdSRP1调控一系列与微菌核形成、孢子形成及致病力相关基因的表达。这些结果表明,VdSRP1基因影响大丽轮枝菌的致病力、微菌核形成、产孢及孢子萌发。     

大丽轮枝菌JR2特异的乙醇脱氢酶的生物学功能研究

大丽轮枝菌是世界范围的植物病原真菌，能够侵染660多种植物，可造成严重的经济损失。大丽轮枝菌基因组分析发现不同菌株含有菌株特异基因，但是大部分基因的生物学功能还不清楚。本研究通过比较大丽轮枝菌VdLs17和JR2的基因组鉴定出1个JR2菌株特异性基因Chr6g02370,该基因编码乙醇脱氢酶，且在侵染过程中被诱导表达。敲除突变体的乙醇脱氢酶酶活性显著降低，但对乙醇等碳源的利用能力并未受到影响。敲除突变体对烟草和番茄的致病力显著降低。表型分析表明敲除该基因降低了黑色素合成能力，但不影响菌丝生长和产孢。研究结果丰富了人们对大丽轮枝菌菌株特异基因生物学功能的认识，为深入开展其功能基因组学研究提供了遗传材料。     

脂肽类抗生素fengycin对大丽轮枝菌孢子萌发和微菌核形成的影响

 枯草芽胞杆菌NCD-2菌株能够产生脂肽类抗生素fengycin和surfactin。本研究分别比较了NCD-2野生型菌株、丧失fengycin合成的突变子MF和丧失surfactin合成的突变子MS对大丽轮枝菌的抑菌活性。结果表明,NCD-2菌株和突变子MS周围能形成清晰的透明圈,而突变子MF周围透明圈模糊,说明fengycin对大丽轮枝菌具有抑菌活性。不同浓度的脂肽提取物对大丽轮枝菌孢子萌发的抑制试验表明,NCD-2野生型菌株的脂肽提取物能抑制大丽轮枝菌孢子萌发,在0.1 mg·mL^-1浓度下对孢子萌发的抑制率为46.30%,且随着脂肽浓度的提高抑制效果更加明显;同野生型菌株相比,突变子MF脂肽提取物显著降低了对大丽轮枝菌孢子萌发的抑制效果,在0.1 mg·mL^-1浓度下对孢子萌发的抑制率仅为18.48%,而突变子MS的脂肽提取物对大丽轮枝菌孢子萌发抑制效果与野生型菌株没有显著差异。通过测试脂肽提取物对微菌核形成的影响,发现野生型菌株和突变子MS脂肽提取物均能够显著抑制微菌核的形成,而突变子MF几乎丧失了对微菌核形成的抑制作用。RT-qPCR结果证明,野生型菌株和突变子MS脂肽提取物能抑制微菌核形成相关基因的表达,而突变子MF丧失了对微菌核形成相关基因表达的抑制作用。以上结果证明,fengycin在NCD-2菌株抑制大丽轮枝菌孢子萌发和微菌核形成中发挥主要作用。     

脂肽类抗生素fengycin对大丽轮枝菌孢子萌发和微菌核形成的影响

 枯草芽胞杆菌NCD-2菌株能够产生脂肽类抗生素fengycin和surfactin。本研究分别比较了NCD-2野生型菌株、丧失fengycin合成的突变子MF和丧失surfactin合成的突变子MS对大丽轮枝菌的抑菌活性。结果表明,NCD-2菌株和突变子MS周围能形成清晰的透明圈,而突变子MF周围透明圈模糊,说明fengycin对大丽轮枝菌具有抑菌活性。不同浓度的脂肽提取物对大丽轮枝菌孢子萌发的抑制试验表明,NCD-2野生型菌株的脂肽提取物能抑制大丽轮枝菌孢子萌发,在0.1 mg·mL^-1浓度下对孢子萌发的抑制率为46.30%,且随着脂肽浓度的提高抑制效果更加明显;同野生型菌株相比,突变子MF脂肽提取物显著降低了对大丽轮枝菌孢子萌发的抑制效果,在0.1 mg·mL^-1浓度下对孢子萌发的抑制率仅为18.48%,而突变子MS的脂肽提取物对大丽轮枝菌孢子萌发抑制效果与野生型菌株没有显著差异。通过测试脂肽提取物对微菌核形成的影响,发现野生型菌株和突变子MS脂肽提取物均能够显著抑制微菌核的形成,而突变子MF几乎丧失了对微菌核形成的抑制作用。RT-qPCR结果证明,野生型菌株和突变子MS脂肽提取物能抑制微菌核形成相关基因的表达,而突变子MF丧失了对微菌核形成相关基因表达的抑制作用。以上结果证明,fengycin在NCD-2菌株抑制大丽轮枝菌孢子萌发和微菌核形成中发挥主要作用。     

匐柄霉对大丽轮枝菌生长及微菌核形成的影响

在离体条件下及棉花植株内测试了匐柄霉对大丽轮枝菌的抑制作用。结果表明,在离体条件下,匐柄霉菌的菌丝体或其培养滤液,对大丽轮枝菌的生长和微菌核形成均有影响。将形态发生变异,不能形成微菌核的大丽轮枝菌菌株转移至ＰＤＡ培养基后,其微菌核形成能力不再恢复。匐柄霉培养滤液的抑菌物质受热不稳定,１００℃处理１０分钟后活性丧失。抑菌物质经硫酸铵、乙醇沉淀初步测定,属非蛋白次生代谢产物。匐柄霉经接种可进入棉株体内,定殖率为７５％～１００％,分布于棉苗根、茎的维管束组织中。与轮枝菌先后或同时混合接种,匐柄霉可减轻棉花黄萎病的症状,以棉苗栽种前用匐柄霉菌分生孢子悬浮液沾根处理效果最明显。     

土壤大丽轮枝菌微菌核的快速定量检测

 微菌核是大丽轮枝菌在土壤中的主要存活结构和黄萎病的初侵染来源。对土壤中大丽轮枝菌微菌核进行定量是黄萎病监测和预警的基础。本研究以大丽轮枝菌Internal Transcribed Spacer (ITS)区特异性引物对P1/P2扩增产物的重组质粒为标准品,构建SYBR Green I实时荧光定量PCR反应的标准曲线,结合土样水筛法建立了土壤大丽轮枝菌微菌核定量检测体系。同时,建立了土壤中微菌核数量与棉花黄萎病发病率的关系模型。结果表明,实时定量PCR检测灵敏度比常规PCR高10倍,检测下限为1个微菌核/克土,在5.54×10²～5.54×10⁷copies范围内,DNA拷贝数的对数值与Ct值具有良好的线性关系。建立的土壤中微菌核个数n与Ct值之间的关系为n=e^7.3-Ct/3.905。温室人工接种微菌核数量与棉花黄萎病发病率间的线性关系为y=2.710n+0.251。     

拮抗性链霉菌对大丽轮枝菌微菌核形成与萌发的影响

为探索拮抗性链霉菌对棉花黄萎病病原大丽轮枝菌Verticillium dahliae Kleb.的抑菌机制,采用菌丝生长速率法、微菌核萌发法研究了6株拮抗链霉菌无菌发酵滤液对大丽轮枝菌生长、微菌核形成与萌发的影响。链霉菌无菌发酵滤液对大丽轮枝菌菌落生长、菌核形成和微菌核萌发有明显抑制作用。其中菌株B49的抑菌效果最好,5倍稀释发酵液培养14天时对菌落生长的抑菌率达69.7%;菌株B49、D184和Act12的5倍稀释发酵液对微菌核形成的抑制率达100%;将经B49、D184和Act12发酵液处理后丧失形成微菌核能力的大丽轮枝菌菌株转接至不含发酵液的PDA培养基,连续传代至第5代,其仍然不能恢复形成微菌核的能力;微菌核在含有菌株D184 5倍稀释发酵液的培养基上培养168 h时,萌发率仅为38.3%。     

江苏省大丽轮枝菌的培养、遗传及致病特性分析

由大丽轮枝菌引起的黄萎病是我国棉花和茄子的主要病害之一,对棉花和茄子的生产造成巨大损失。为了研究大丽轮枝菌的群体遗传变异以及对棉花和茄子的交互致病性,本文对分离自江苏省的63个棉花黄萎菌和10个茄子黄萎菌进行了培养、遗传和致病特性分析。根据菌株在PDA培养基上生长时形成微菌核的多少来划分培养类型,结果菌核型占83.6%,成为主要的培养类型。用PCR技术检测菌株的致病类型、交配型以及是否具有无毒基因Ave1,结果落叶型菌株占86.3%,为优势种群,但是10株茄子黄萎菌100%都是非落叶型菌株;供试的所有江苏菌株交配型都是MAT1-2型,并且都没有无毒基因Ave1。选择江苏省的6个棉花黄萎菌和4个茄子黄萎菌在室内苗期接种棉花和茄子,进行交互致病性测定,结果这10个菌株都可以侵染棉花和茄子,而且来源于不同寄主作物的菌系之间致病力分化明显,表现在不同菌株对同一寄主的致病力不同,同一菌株对不同寄主的致病力也不同。研究结果为深入研究大丽轮枝菌群体遗传结构和制定黄萎病防治措施提供了理论支持。     

多菌灵、三环唑对大丽轮枝菌微菌核、黑色素形成及致病力的影响

 在离体和活体条件下测定了多菌灵、三环唑对大丽轮枝菌的微菌核黑色素形成的影响,以及经2种杀菌剂处理产生的形态变异菌株对棉苗的致病力。结果表明:多菌灵在培养基内含量超过0.1μg/ml时,即可抑制大丽轮枝菌微菌核的形成,随着培养基内多菌灵浓度的增加,微菌核形成时间逐渐延长,形成量逐渐减少;三环唑浓度为0.5μg/ml时,可抑制微菌核的黑色素形成,微菌核从黑色变为浅红至红褐色。三环唑对微菌核黑色素形成呈可逆抑制,变色的微菌核菌落移入不含药的培养基后,大多可恢复形成黑色素。培养基内三环唑浓度的提高,也可抑制大丽轮枝菌微菌核的形成;因2种杀菌剂的抑制而丧失形成微菌核的白色菌丝体移入不含药的培养基,微菌核形成能力也不能恢复。多菌灵和三环唑经棉株吸收后均能抑制植株内大丽轮枝菌微菌核的形成,但对微菌核色素的形成未见有明显影响。2种杀菌剂处理产生的形态变异菌株的致病力与野生型菌株致病力差异不显著。     

Weed hosts of <Emphasis Type="Italic">Verticillium dahliae</Emphasis> in cotton fields in Turkey and characterization of <Emphasis Type="Italic">V. dahliae</Emphasis> isolates from weeds

A weed survey conducted in 2004 and 2005 in Aydin province of Turkey showed that Solanum nigrum, Xanthium strumarium, Amaranthus retroflexus, Portulaca oleracea, Sonchus oleraceus and Datura stramonium were the most prevalent weeds in the cotton fields exhibiting Verticillium wilt. Verticillium dahliae Kleb. was recovered from A. retroflexus and X. strumarium in those cotton fields. This is the first report of V. dahliae occurring naturally in A. retroflexus in Turkey. Pathogenicity tests on cotton and weeds showed that the virulence of V. dahliae isolates from weeds was higher on cotton plants than on weeds, with the disease severity ranging from 31.7% to 98.0%. Disease severity of V. dahliae isolates was 54.7–93.9% on eggplant, 23.7–51.6% on cucumber and 11.0–16.4% on tomato, whereas it did not cause any disease symptoms, or only low levels, on pepper and bell pepper. Two vegetative compatibility groups (VCGs) were identified among seven tested weed isolates: VCG2A (two isolates) and VCG2B (three isolates) using international reference strains.     

新疆棉花黄萎病的发生现状及其病原菌的分子鉴定与ISSR分析

为掌握新疆主要植棉区棉花黄萎病的发生现状及其病原菌大丽轮枝菌Verticillium dahliae的落叶型菌系分布以及遗传变异情况,于2015年对26个新疆主要植棉区棉花黄萎病的发生情况进行了随机调查,统计新疆大丽轮枝菌的培养性状,利用大丽轮枝菌落叶型特异引物D1/D2、INTD2F/INTD2R与非落叶型特异性引物ND1/ND2、INTNDF/INTNDR对新疆大丽轮枝菌菌系进行互补鉴定,并对部分菌系的遗传变异进行简单序列重复区间(inter simple sequence repeat,ISSR)分析。结果表明:2015年新疆棉花黄萎病发病田比例为54.0%,其中病情指数在10.0以上的发病田与2013年持平,而病情指数在20.0以上的严重发病田比例为10.8%,比2013年增加3.8个百分点;新疆大丽轮枝菌的培养性状以菌核型为主,比例为70.1%,菌丝型与中间型比例分别为13.4%和16.5%;新疆大丽轮枝菌落叶型菌系比例为53.2%,26株菌株的来源地全部检出落叶型菌系;聚类分析结果显示,当遗传相似系数为0.66时,新疆大丽轮枝菌落叶型与非落叶型菌系聚为2个谱系,菌系地理来源、培养性状与大丽轮枝菌的遗传分化无明显相关性。     

胶孢炭疽菌G蛋白信号调控因子CgRGS3的生物学功能

G蛋白信号调控因子(regulators of G-protein signaling,RGS)是G蛋白的一类负调控因子,在植物病原真菌生长发育及致病过程中起着重要作用。为探究胶孢炭疽病菌1个RGS基因CgRGS3的生物学功能,利用PCR技术扩增CgRGS3基因并进行生物信息学分析,通过同源重组的方法获得该基因的敲除突变体,并在突变体的基础上获得互补株,通过表型分析确定CgRGS3的生物学功能。结果表明:CgRGS3编码1个含有367个氨基酸的蛋白,包含1个RGS功能域和3个跨膜区域,表型分析发现与野生型菌株相比,敲除突变体营养生长缓慢,分生孢子产量附着胞形成率显著降低,对Cu~(2+)、Fe~(2+)、Zn~(2+)离子更加敏感,细胞壁完整性发生改变,黑色素产量降低及致病力减弱等。由此可见,CgRGS3参与调控胶孢炭疽菌的营养生长,分生孢子产生及附着胞的分化、细胞壁完整性、黑色素产生及致病性等。     

Vegetative compatibility groups inVerticillium dahliae isolates from cotton in Turkey

Verticillium dahliae Kleb. with a complicated genetic diversity is a widely distributed major pathogen resulting in cotton wilt, which causes high economic losses in cotton lint production in the cotton belt of Turkey. A collection of 70 TurkishV. dahliae isolates (68 from wilted cotton plants in 28 districts and two from watermelon plants in two districts) were tested for vegetative compatibility by observing heterokaryon formation among complementary nitrate-nonutilizing (nit) mutants. The mutants were tested against international reference tester isolates and also were paired with one another. Thirty-nine isolates were assigned to vegetative compatibility group (VCG) 2B, 19 to VCG2A and three to VCG4B. One isolate was self-incompatible and eight others could not be assigned to any of the identified VCGs because theirnit mutants showed negative reactions with the tester isolates of four VCGs or theirnit mutants reverted back to the wild type. This is the first report of VCGs inV. dahliae from cotton in Turkey.     

Mating type gene MAT1-2-1 is common among Japanese isolates of Verticillium dahliae

Mating type genes of Verticillium dahliae, a wilt pathogen affecting many plant species, were identified to examine sexual recombination between Japanese pathotypes. We amplified a DNA sequence encoding high mobility group (HMG) box from V. dahliae using PCR. A cloned genomic DNA fragment included a sequence homologous to MAT1-2-1 gene. Despite that sequence's presence in all V. dahliae isolates we used, MAT1-1-1 (an opposite mating type gene) was never amplified. We concluded that V. dahliae is potentially heterothallic. Furthermore, sexual bias practically obviates sexual recombination between Japanese pathotypes. This report describes, for the first time, a mating type gene of phytopathogenic Verticillium.     

Differentiation ofVerticillium dahliae populations on the basis of vegetative compatibility and pathogenicity on cotton

Complementary auxotrophic nitrate-nonutilizing (nit) mutants were used to investigate vegetative compatibility within 27 strains ofVerticillium dahliae isolated from several hosts originating from Africa, Asia, Europe and the United States. Using about 500nit mutants generated from these strains, three vegetative compatibility groups, 1, 2 and 4, were identified. Simultaneously, virulence of each strain was assessed on cultivars ofGossypium hirsutum, G. barbadense andG. arboreum, based upon Foliar Alteration Index (FAI) and Browning Index (BI) estimation. The strains in VCG1 were of both the cotton-defoliating pathotype and race 3 (on cotton) but were non pathogenic on tomato; those in VCG2 and VCG4 were of the nondefoliating pathotype and belonged to different races on cotton and on tomato. Hyaline mutants deriving from parental wild-type strain showed differences in pathogenicity but were always assigned to the parental VCG. A relationship was established between VCGs and the taxonomic position of host plants. Data fromnit pairings indicated that the sub-populations ofV. dahliae (VCGs) may not be completely isolated genetically.     

棉花内生真菌CEF-373菌株对棉花黄萎病的防效及其作用机理

为明确棉花内生真菌CEF-373菌株对棉花黄萎病的防效及其作用机理,利用圆盘滤膜法和平板对扣培养法测定菌株CEF-373对大丽轮枝菌Verticillium dahliae菌丝生长的抑制作用,测定其对棉花黄萎病的温室和田间防效,并通过活性氧含量及防御基因表达情况来分析其作用机理。结果表明,菌株CEF-373的挥发性代谢产物和非挥发性代谢产物对大丽轮枝菌菌丝的生长均有显著抑制作用,抑制率最高分别可达37.75%和100.00%。用1×107CFU/mL的菌株CEF-373分生孢子悬浮液灌根后,对棉花黄萎病的温室防效可达71.12%,用质量比为3%的菌株CEF-373固体菌剂拌土栽培后,对棉花黄萎病的温室防效可达62.74%,防治作用显著。菌株CEF-373的发酵液滴灌和微生物肥料处理40 d后,对棉花黄萎病的田间防效达到最大,分别为36.23%和27.71%,而在后期有所降低。菌株CEF-373可以诱导棉花叶片中细胞活性氧的爆发;且菌株CEF-373成功激活了苯丙氨酸解氨酶、过氧化物酶、多酚氧化酶、几丁质酶和病程相关蛋白基因PR10的表达,对大丽轮枝菌的侵染具有抵御作用。表明棉花内生真菌CEF-373菌株通过抑制大丽轮枝菌生长以及诱导寄主系统抗病性来有效防治棉花黄萎病,具有较好的生物防治应用前景。     

Comparative Study of Genetic Diversity and Pathogenicity Among Populations of Verticillium Dahliae from Cotton in Spain and Israel

Genetic diversity and phenotypic diversity in Verticillium dahliae populations on cotton were studied among 62 isolates from Spain and 49 isolates from Israel, using vegetative compatibility grouping (VCG), virulence and molecular assays. In Spain, defoliating V. dahliae isolates (D pathotype) belong to VCG1, and non-defoliating isolates (ND) belong to VCG2A (often associated with tomato) and VCG4B (often associated with potato). The D pathotype was not identified in Israel. The ND pathotype in Israel is comprised of VCG2B and VCG4B. Isolates in VCG2B and VCG4B ranged in virulence from weakly virulent to highly virulent. The highly virulent isolates induced either partial defoliation or no defoliation. Virulence characteristics varied with inoculation method and cotton cultivar. Highly virulent isolates from Israel were as virulent as D isolates from Spain under conditions conducive to severe disease. The D pathotype is pathologically and genetically homogeneous, whereas the ND pathotype is heterogeneous with respect to virulence, VCG, and molecular markers based on single-primer RAPD and on PCR primer pairs.     

Genetic variability and virulence among Verticillium albo-atrum isolates from hop

Verticillium wilt, caused by Verticillium albo-atrum or V. dahliae, is an important disease of many worldwide crop species. In Europe, V. albo-atrum isolates infecting hop express different levels of virulence, inducing mild or lethal disease syndromes, and it is therefore an attractive model for studying the virulence of this pathogen. In this work, eleven amplified fragment length polymorphism (AFLP) primer combinations were used to analyze genetic variability among 55 V. albo-atrum hop isolates from four European hop growing regions, as well as isolates from other hosts and V. dahliae isolates. Cluster analysis divided V. albo-atrum and V. dahliae isolates into two well-separated groups. Within the V. dahliae cluster, isolates were separated without host specific grouping, although no host adapted isolates were included. In V. albo-atrum, the alfalfa isolates were distinct from isolates of other hosts, where a high association with virulence was observed in hop and tomato isolates. All lethal hop isolates were genetically different from mild hop isolates. The lethal hop isolates from England and Slovenia expressed the same virulence phenotype, although they showed a different AFLP pattern. The mild hop isolates formed two subgroups, to which isolates clustered irrespective of geographical location. These data suggest multiple origins of V. albo-atrum hop isolates, and the possible appearance of new virulent isolates in the future in other hop growing regions.     

棉花黄萎病拮抗细菌H14的筛选鉴定及其拮抗机理分析

为筛选对大丽轮枝菌Verticillium dahliae Kleb.具有拮抗活性的根际细菌,以大丽轮枝菌为靶菌,分离获得棉花根际细菌,利用平板对峙法和琼脂扩散法筛选具有较高拮抗活性的菌株,采用室内盆栽法测定筛选所得菌株对棉花黄萎病的防效,并通过形态特征、生理生化特性和16S r DNA基因序列分析确立其分类地位,采用底物降解法和抗菌肽基因克隆法检测其产生水解酶和抗菌肽的能力。结果显示,试验共分离获得182株棉花根际细菌,筛选得到3株对大丽轮枝菌抑菌率大于50.00%且抑菌圈直径大于15 mm的菌株,其中菌株H14的抑菌率和抑菌圈直径最大,分别为54.25%和18.10 mm。该菌能在0～9%NaCl和pH 4.5～9.0的NB培养基上生长;经形态特征观察、生理生化试验及16S rDNA基因序列分析,最终将其鉴定为莫哈韦芽胞杆菌Bacillus mojavensis;菌株H14对大丽轮枝菌孢子萌发的抑制率和对棉花黄萎病的盆栽防效分别为89.55%和74.57%;菌株H14能合成蛋白酶,含有srfAA、fenD、bacA脂肽类抗生素合成基因。表明莫哈韦芽胞杆菌菌株H14能够合成蛋白酶和脂肽类拮抗物质,具有良好的抑菌和防病能力。