Isolation and Identification of Lactiplantibacillus plantarum ST3.5 and Its Inhibitory Effect on Mold
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摘要: 乳酸菌因具有拮抗霉菌等有害微生物的潜力，有望成为下一代安全、稳定的生物抗菌剂。本研究筛选获得了一株抑制霉菌活性较好的菌株ST3.5，基于16S rRNA测序鉴定为植物乳杆菌，并对其耐酸耐胆盐、抑制病原菌、耐药等特性进行分析。进一步对植物乳杆菌ST3.5的无细胞上清液进行酸处理、热处理及酶处理，分析其中主要的抑霉菌物质，通过高效液相色谱（HPLC）测定上清液中有机酸含量，扫描电镜（SEM）观察其对霉菌菌丝的破坏情况，并以玉米粒为实际样本进行生物防治试验。结果表明，植物乳杆菌ST3.5具有良好的耐酸特性，耐胆盐能力较弱，对庆大霉素、卡那霉素等抗生素具有耐药性，对氨苄西林、氯霉素等抗生素敏感，可以抑制致病菌的生长。有机酸分析发现植物乳杆菌ST3.5无细胞上清液中乳酸含量最高（22.02±0.23）g/L，其次是柠檬酸（4.99±0.04）g/L和乙酸（3.67±0.06）g/L。SEM结果显示上清液对霉菌菌丝有破坏作用。此外，生物防治实验证实了植物乳杆菌ST3.5无细胞上清液能够抑制玉米表面霉菌的生长。综上所述，植物乳杆菌ST3.5能够抑制霉菌生长，可用于开发生物防治制剂，以最大限度地减少霉菌污染并保障食品安全。Abstract: Lactic acid bacteria (LAB) has the potential to antagonize harmful microorganisms such as molds, which is expected to be safe and stable biological antimicrobial agents of the next generation. In present study, a strain ST3.5 with inhibiting mold activity was screened and obtained, which was identified as Lactiplantibacillus plantarum (L. plantarum) based on 16S rRNA sequencing. And its characteristics of acid resistance, bile salt resistance, inhibition of pathogenic bacteria, and drug resistance were analyzed. Furtherly, the cell-free supernatant (CFS) of L. plantarum ST3.5 was treated with acid, heat and enzyme, which the main antifungal substances were analyzed. The content of organic acids in supernatant was analyzed by high performance liquid chromatography (HPLC) and the destruction of mold mycelium was observed by scanning electron microscope (SEM). Meanwhile, the biological control experiment was carried out with corn kernels as an actual sample. The results showed that L. plantarum ST3.5 had good acid resistance and weak bile salt resistance. The strain was resistant to antibiotics (e.g., gentamicin and kanamycin) and sensitive to antibiotics (e.g., ampicillin and chloramphenicol), which could inhibit the growth of pathogenic bacteria. Analysis of organic acid found that the content of lactic acid in the CFS was the highest (22.02±0.23) g/L, followed by citric acid (4.99±0.04) g/L and acetic acid (3.67±0.06) g/L. The SEM results demonstrated the CFS could damage mold hyphae. Additionally, the biocontrol experiment confirmed that the CFS of L. plantarum ST3.5 could inhibit the mold residing growth of the corn surface. In conclusion, L. plantarum ST3.5 can inhibit the growth of mold, which can be used to develop biological control agents to minimize mold pollution and ensure food safety.
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Key words:
- lactic acid bacteria /
- Lactiplantibacillus plantarum /
- screening /
- identification /
- mold inhibitory /
- corn kernels
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图 1 菌株ST3.5抑制产黄青霉和杂色曲霉效果图
Figure 1. Effect diagram of strain ST3.5 inhibiting P. chrysogenum and A. versicolor
注：A：产黄青霉；B：ST3.5抑制产黄青霉；C：杂色曲霉；D：ST3.5抑制杂色曲霉。
图 3 分离菌株 ST3.5 镜检结果（1000×）
Figure 3. Results of microscopic examination of isolated strain ST3.5(1000×)
图 5 L. plantarum ST3.5的生长曲线及发酵液pH变化曲线
Figure 5. Growth curve and pH change curve of fermentation broth of L. plantarum ST3.5
注：不同小写字母表示不同时间pH的差异显著（P<0.05）；不同大写字母表示不同时间OD600 nm值的差异显著（P<0.05）。
图 6 不同处理的L. plantarum ST3.5-CFS对霉菌的抑制效果
Figure 6. Inhibitory effect of different treated L. plantarum ST3.5-CFS on mold
注：*，#：P<0.05 ；**，##：P<0.01；***，###：P<0.001表示两菌抑菌效果与CFS组相比的显著性。
图 7 L. plantarum ST3.5-CFS中的有机酸含量
Figure 7. Content of organic acids in L. plantarum ST3.5-CFS
注：不同字母表示不同有机酸含量的差异显著（P<0.05）。
图 8 霉菌菌丝扫描电镜图
Figure 8. Scanning electron microscope image of mold mycelium
注：A：产黄青霉；B：杂色曲霉；a：对照组；b：CFS处理组；1：菌丝形态（标尺：30 μm）；2：菌丝形态（标尺：5 μm）；3：菌丝尖端（标尺：5 μm）。
图 9 ST3.5-CFS处理前后的玉米粒对霉菌的抑制效果图
Figure 9. Inhibition effect of corn kernels treated with ST3.5-CFS on mold
注：A、B是产黄青霉的对照组和处理组，C、D是杂色曲霉的对照组和处理组。
表 1 菌株对产黄青霉和杂色曲霉的抑制效果
Table 1. Inhibitory effect of strain on P. chrysogenum and A. versicolor
序号 菌种编号 抑制产
黄青霉抑制杂
色曲霉序号 菌种编号 抑制产
黄青霉抑制杂
色曲霉1 H1.1 ++ ++ 18 S4.2 +++ ++ 2 H1.2 （−） ++ 19 ST1.1 ++ +++ 3 H1.3 ++ + 20 ST1.2 +++ ++ 4 H1.4 （−） ++ 21 ST1.3 ++ +++ 5 H2.2 ++ + 22 ST2.1 ++ +++ 6 H3 ++ ++ 23 ST2.2 +++ ++ 7 H6 + ++ 24 ST2.3 （−） +++ 8 L2.2 ++ +++ 25 ST2.4 +++ ++ 9 L2.5.1 +++ ++ 26 ST3.4 （−） +++ 10 L4.1 +++ ++ 27 ST3.5 +++ +++ 11 L4.3 +++ − 28 ST4.3 （−） + 12 L4.5 +++ ++ 29 ST4.4 +++ ++ 13 L5.1 ++ （−） 30 ST4.5 +++ ++ 14 L5.2 ++ （−） 31 ST4.6 （−） + 15 L5.3 − ++ 32 ST5.3 （−） （−） 16 L5.5 （−） ++ 33 ST5.4 ++ +++ 17 L6 （−） +++ 34 ST5.5 + ++ 注：“−”，没有抑制作用；“（−）”，微弱抑制，每个乳酸菌线周围有占总平板面积0.1%~10%的抑菌圈；“+”，每个乳酸菌线周围有占总平板面积10%~30%的抑菌圈；“++”，每个乳酸菌线周围有占总平板面积30%~50%的抑菌圈；“+++”，每个乳酸菌线周围有占总平板面积50%以上的抑菌圈。 
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表 2 L. plantarum ST3.5的耐酸耐胆盐效果
Table 2. Acid and bile salt tolerance of L. plantarum ST3.5
酸耐受性 活菌数（log CFU/mL） 0 h 1 h 3 h pH5 9.16±0.11bB 9.46±0.04bA 9.83±0.21aA pH4 8.94±0.02bB 9.63±0.33aA 9.48±0.02aAB pH3 8.98±0.14bB 9.31±0.02aA 9.33±0.21aB pH2 9.61±0.05aA 7.26±0.24bB 4.48±0.00cC 胆盐耐受性 活菌数（log CFU/mL） 0 h 1 h 3 h 0.1% 9.10±0.04aA 9.11±0.33aA 9.31±0.05aA 0.2% 6.78±0.16aB 6.78±0.79aB 6.09±0.98aB 0.3% 5.94±0.34aC 5.51±0.20bC ND 注：a、b、c表示相同的酸度或胆盐浓度条件下不同的处理时间的差异显著性（P<0.05）；A、B、C表示相同的时间下不同的酸度或胆盐浓度的差异显著性（P<0.05）。
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表 3 L. plantarum ST3.5对致病菌的抑制效果
Table 3. Inhibitory effect of L. plantarum ST3.5 on pathogenic bacterias
菌株 抑菌圈（mm） 大肠杆菌 20.00±0.00bc 金黄色葡萄球菌 25.00±0.00a 铜绿假单胞菌 18.33±0.58c 单细胞增生李斯特菌 21.00±1.73b 注：a、b、c表示对不同致病菌的抑菌圈大小的差异显著性（P<0.05）。
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表 4 L. plantarum ST3.5对不同抗生素敏感性
Table 4. Sensitivity of L. plantarum ST3.5 to different antibiotics
分类 抗生素 敏感性 分类 抗生素 敏感性 氨基糖苷类 阿米卡星 S 青霉素类 青霉素 R 庆大霉素 R 苯唑西林 R 卡那霉素 R 氨苄西林 S 新霉素 R 羧苄西林 S 链霉素 R 哌拉西林 S 大环内酯类 麦迪霉素 S 喹诺酮类 环丙沙星 R 红霉素 S 诺氟沙星 R 头孢菌素类 头孢呋辛 S 氧氟沙星 I 头孢他啶 S 其他 万古霉素 R 头孢唑林 S 多粘菌素B R 头孢哌酮 I 呋喃唑酮 I 头孢拉定 S 氯霉素 S 头孢曲松 S 利福平 S 头孢氨苄 S 复方新诺明 R 头孢噻肟 R 克林霉素 R 四环素类 多西环素 S 四环素 R 米诺环素 R 注：S：敏感；I：中度敏感；R：耐药。
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