#对本文贡献相同
#These authors contributed equally to this work.
近年来,由土传致病链霉菌引起的疮痂病在我国马铃薯产区大面积暴发,严重影响块茎的商品价值,由于缺乏有效防控措施,其危害程度呈逐年加重趋势。然而拮抗菌可有效抑制病原菌的繁殖,其防控效果优异,对环境友好,逐渐引起了人们的关注。
分离筛选对致病链霉菌具有拮抗功能的细菌,鉴定其分类学名,评价其主要生物学特性,为研发高效防病菌剂提供菌种资源。
以疮痂链霉菌为靶向微生物,从马铃薯疮痂病高发土样中筛选拮抗菌株,通过形态学、生理生化和分子生物学试验确定其分类地位;用盆栽试验初步判定其对马铃薯疮痂病的防控效果,研究其广谱抗病性和耐盐碱特性,结合生产中常用化学农药的敏感性测定,综合评价其应用前景。
筛选出一株对致病疮痂链霉菌(
菌株HD9-1鉴定为枯草芽孢杆菌,对马铃薯疮痂病防控效果较好,且具有良好的环境适应性,可作为防治马铃薯土传病害的高效复合菌剂的备选菌株。
The soil-borne potato common scab (CS), caused by
To isolate and identify antagonistic bacterial isolate against
Antagonistic isolates were screened from soil with high incidence of potato CS. The taxonomic status was confirmed with morphological, physiological, biochemical, and molecular methods. The control effect was analyzed by pot trials. Then, the potential application of the isolate was evaluated by broad-spectrum resistance (BSR) test, salt-alkali tolerance assay, and sensitivity tests to chemical fungicides.
HD9-1 with significant antagonistic effect on
HD9-1, a
马铃薯作为重要的粮食作物,具有极大的市场需求量,约70%的马铃薯产区集中分布于西北、西南、华北、东北等地,是当地农民的主粮作物和重要经济收入来源[
马铃薯疮痂病由多种致病链霉菌(
本研究以
疮痂链霉菌(
细菌理化性质检测试剂盒,北京兰伯瑞生物科技有限公司。显微镜:显微镜数码采集装置购自北京长恒荣创科技有限公司;日立冷场发射扫描电子显微镜购自Hitachi公司。Biolog GENIII鉴定板1030,北京兰伯瑞生物科技有限公司。
PDA培养基(g/L):土豆200.0,琼脂15.0,葡萄糖20.0,调整其pH值至7.0−7.2。LB培养基(g/L):蛋白胨10.0,酵母提取物5.0,NaCl 10.0,调整其pH值至7.0−7.2。King氏培养基(g/L):蛋白胨20.000,K2HPO4 1.725,丙三醇0.015,MgSO4
取1 g土样置于50 mL离心管,加入10 mL蒸馏水,振荡使土壤均匀悬浮。吸取100 µL 10−3、10−4、10−5稀释液滴至LB固体培养基并均匀涂布,28 ℃培养24 h,挑选单菌落划线纯化,将获得的菌株接种于LB液体培养基中,于37 ℃、200 r/min振荡培养过夜备用。
取100 µL疮痂链霉菌孢子悬浮液均匀涂布于PDA固体培养基,吸取6 µL细菌培养液滴于上述培养皿中央,28 ℃共培养7 d后测量抑菌圈直径。在4 ℃、4 500 r/min离心15 min,分别收集参试菌株的培养上清液和菌体沉淀,用等量无菌水重新悬浮菌体,以LB液体培养基为对照,将培养上清液、菌体悬浮液和培养液同时进行平板对峙试验,培养48 h后观察并记录抑菌圈的直径。
将待测菌液均匀涂布于LB固体培养基,37 ℃培养24 h,观察菌落的形态特征,并用显微镜数码采集装置LV2000拍照记录形态特征,采用日立冷场发射扫描电子显微镜SU8010观察菌体形态,测量10个菌体大小。
将待测菌液滴至载玻片上自然晾干,轻微烘烤固定菌体,滴加草酸铵结晶紫试剂,染色1 min后用水冲洗,碘液染色1 min再次冲洗,用95%乙醇脱色,滴加番红复染3−5 min,水洗后进行镜检。
采用Biolog GENIII鉴定板1030鉴定菌株可利用碳源;用试剂盒测试细菌的理化性质(具体操作见说明书);接触酶试验参照《常见细菌系统鉴定手册》[
将待测菌株分别接种至pH值为3.0、5.0、7.0、9.0、11.0,NaCl含量为1%、3%、5%、7%、9%、11%的LB培养基中,37 ℃、200 r/min振荡培养12 h后测定其
参考李建萍[
分别在PDA培养基上接种大丽轮枝菌、立枯丝核菌和尖孢镰刀菌,在其两侧间隔1 cm处分别接种6 μL HD9-1培养液,并以单独接种病原菌的处理为对照,28 ℃培养4 d观察抑菌效果。抑菌率(%)=(对照组面积−处理组面积)/对照组面积×100。
将细菌培养液送至北京博迈德生物工程股份有限公司,运用16S rRNA基因片段通用引物27F (5′-AGAGTTTGATCCTGGCTCAG-3′)和1492R (5′-CTACGGCTACCTTGTTACGA-3′)进行PCR。PCR反应体系(20 µL):2×T5 Super PCR Mix 10 µL,HD9-1基因组DNA (200 ng/μL) 2 µL,正、反向引物(10 µmol/L)各1 µL,ddH2O 6 µL。PCR反应条件:95 ℃ 5 min;95 ℃ 30 s,55 ℃ 30 s,72 ℃ 2 min,35个循环;72 ℃ 10 min。测序结果在National Center for Biotechnology Information (NCBI)上通过BLASTn进行比对查找与HD9-1相似性较高的序列,用MEGA 7.0软件构建菌株系统发育树。
在直径大小为17 cm的花盆中装满蛭石,每盆移栽马铃薯品种夏波蒂的脱毒试管苗3株,置于光照强度为50 000 lx、光周期为L16 h/D8 h的条件下培养。10 d后在每株幼苗的根部注射1×107 CFU/mL的
病害分级标准:马铃薯皮完好,无病斑,为0级;表皮病斑面积为大于0小于1/6,为1级;表皮病斑面积为大于1/6小于1/3,为2级;表皮病斑面积为大于1/3小于1/2,为3级;表皮病斑面积大于1/2,为4级。
发病率(%)=发病块茎数/调查总块茎数×100;
病情指数=∑(各病级块茎数×该病级数代表值)/(调查个体总和×最高病级数)×100%;
相对防效(%)=(对照组病情指数–处理组病情指数)/对照组病情指数×100。
参照李振东等[
在直径大小为17 cm的花盆中装满蛭石,每盆移栽马铃薯品种夏波蒂的脱毒试管苗3株,置于光照强度为50 000 lx、光周期为L16 h/D8 h的条件下培养7 d后每盆均匀浇施1×108 CFU/mL的拮抗菌悬浮液30 mL,以加入30 mL无菌水处理为对照,分别于室温培养0 d、14 d时统计马铃薯株高。
采用Origin 2019b和Excel 2016软件处理相关数据。
从采自广东省惠州市惠东县的薯田土壤中分离出一株对
菌株HD9-1对疮痂链霉菌的抑制效果
The inhibitory effect of screening strain HD9-1 on
HD9-1培养上清液、菌体悬浮液和培养液同时进行平板对峙试验,抑菌效果如
如
菌株HD9-1形态学特征
Morphological characteristics of HD9-1.
菌株HD9-1的生理生化特性
Biochemical properties of HD9-1
Test items | Reaction |
+: Positive; –: Negative. | |
2-nitrophenyl | + |
Arginine | + |
Lysine | + |
Ornithine | + |
Sodium citrate | + |
Sodium thiosulfate | – |
Uridine | + |
Tryptophan | + |
Melibiose | – |
Amygdalin | + |
Pyruvate | + |
Kohn gelatin | + |
Glucose | – |
Mannitol | – |
Inositol | – |
Sorbitol | – |
Rhamnose | – |
Sucrose | + |
Arabinose | – |
以HD9-1的基因组DNA为模板,利用通用引物扩增目的基因片段,序列已上传至NMDC数据库(登录号:NMDCN00011OJ)。其序列在National Center for Biotechnology Information (NCBI)进行比对,发现与
基于16S rRNA基因序列构建的菌株HD9-1系统发育树
Phylogenetic tree of HD9-1 constructed based on the 16S rRNA gene sequences.
盆栽试验共收获马铃薯微型薯31粒(
菌株HD9-1的盆栽抑病试验结果
The results of potted disease inhibition test of HD9-1.
菌株HD9-1的防治效果统计
Control effect statistics of HD9-1
Treatment | Number of grains | Sickness | Occurrence rate (%) | Disease index | Control effects (%) | ||||
0 | 1 | 2 | 3 | 4 | |||||
/: The effectiveness of control 1 and control 2 does not need to be calculated. | |||||||||
Control 1 | 10 | 11 | 0 | 0 | 0 | 0 | 0.00 | 0.00 | / |
Control 2 | 11 | 1 | 0 | 2 | 1 | 7 | 90.91 | 79.55 | / |
HD9-1 | 10 | 3 | 4 | 1 | 1 | 1 | 70.00 | 32.50 | 59.15 |
菌体悬浮液分别经37、60、80 ℃处理30 min后,平均活菌数分别为1.6×109、6.0×108、1.2×106 CFU/mL,100 ℃处理30 min后仍有1.5×104 CFU/mL的菌体存活(
HD9-1的高温耐受性
High temperature tolerance of HD9-1
Temperature (℃) | Colony number (CFU/mL) |
表中数据格式为平均数±标准偏差 |
|
37 | 1.6×109±8.2×106 |
60 | 6.0×108±8.1×105 |
80 | 1.2×106±3.2×103 |
100 | 1.5×104±4.1×103 |
HD9-1拮抗菌耐盐碱范围
Salt-alkali tolerance range of HD9-1.
多种化学农药对HD9-1生长的抑制效果
Inhibitory effects of pesticides on HD9-1
Fungicide | Manufacturer | Potency (%) | Tolerance |
+:拮抗性细菌生长没有影响;–:抑制拮抗性细菌生长;/:H2O的防效,不需计算 |
|||
H2O | / | / | + |
Zhongshengmycin | Fujian Kaili Biological Products Limited Company | 0.10 | + |
Mancozeb | Sichuan Runer Technology Limited Company | 0.10 | – |
Difenoconazole | Hebei Zhongbao Green Crop Technology Limited Company | 0.10 | + |
Flutamicarb | Bayer Crop Science (China) Limited Company | 0.10 | + |
Kasugamycin | Shaanxi McColo Biological Limited Company | 0.10 | + |
Flusilazole | Jiangsu Jiannong Plant Protection Limited Company | 0.01 | + |
Pyrazole ether fungus ester | Jinan Zhongke Green Biological Engineering Limited Company | 0.05 | + |
Azoxystrobin | Xianzhengda Nantong Crop Protection Limited Company | 0.10 | + |
Fluridylamine | Ningbo Shiyuan Jinniu Agricultural Technology Limited Company | 0.10 | – |
Thiophanate-methyl | Jinan Taihe Chemical Limited Company | 0.10 | + |
菌株HD9-1的广谱抑菌效果
Wide-spectrum resistance of strain HD9-1. Control: The plate without HD9-1; T: The plate with HD9-1.
由此可知,HD9-1可以拮抗多种病原菌,其广谱拮抗性有待进一步挖掘。
IAA的测定通常采用SalKowski比色法。
菌株HD9-1代谢产物中生长素测定
Auxin assay in strain HD9-1 metabolites.
盆栽试验结果表明,添加HD9-1培养14 d后,土豆苗株高为13.82 cm,较对照增高了2.88 cm;生长速度为0.47 cm/d,高出对照0.13 cm/d (
菌株HD9-1的促生能力分析
Analysis of growth-promoting ability of strain HD9-1.
在马铃薯生产中,种植户为了追求产量和效益而过量施肥用药的现象普遍存在。化肥与农药的不合理施用导致土壤生态失衡,以疮痂病为代表的土传病害逐年加剧,不仅造成巨大的经济损失,也严重威胁核心产区产业(尤其种薯产业)的可持续发展。人们开展马铃薯疮痂病防控研究已有100多年历史,但至今尚未取得重大突破,未开发出可规模化应用的特效化学药剂及其他措施[
使用化学药剂及微生物制剂是目前缓解马铃薯疮痂病危害的主要方法,研究者利用棉隆和辣根素水剂熏蒸土壤,对疮痂病的防效可达80%左右,但是防治效果与土壤质地密切相关,且随着熏蒸次数的增加,土壤中有毒物质含量也显著升高,严重影响农产品品质[
本文从广东薯田土壤中分离、筛选的枯草芽孢杆菌HD9-1,对
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