解淀粉芽胞杆菌L19对马铃薯枯萎病菌的抑制及植株的促生作用

doi:10.16420/j.issn.0513-353x.2022-0876

摘要/Abstract

摘要：

为获得对马铃薯枯萎病菌具有优良拮抗效果，并对马铃薯植株具有促生作用的双重功能生防菌株，采用平板对峙方法测定分离筛选得到的菌株L19发酵产物对马铃薯枯萎病菌的生长抑制作用，利用PCR技术和气相色谱—质谱联用技术（GC-MS）分别对其脂肽类抗生素和挥发性化合物进行检测鉴定，采用温室盆栽试验和叶绿素荧光检测技术研究了其对马铃薯植株的促生作用以及对光合作用的影响。通过拮抗试验共分离得到25株生防菌株，其中菌株L19可显著抑制尖孢镰刀菌菌丝的生长，经形态学及gyrB序列分析，鉴定其为解淀粉芽胞杆菌（Bacillus amyloliquefaciens）。光学显微观察发现，L19发酵液致使马铃薯枯萎病菌菌丝扭曲、黏连、膨大和泡状化等畸形，菌丝丧失穿透力，并且随着发酵液浓度的增加，其孢子萌发率降低。L19含有sfp、ituA、fenB、bacD、mycB等脂肽合成基因；可产生12种挥发性化合物，芳香族含量所占比例为47.42%，其次为醇类和酮类，其中2-乙基甲苯为主要挥发性物质，占比20.91%；具有产生纤维素酶的能力。盆栽试验表明，不同浓度L19发酵液可增加马铃薯植株的生长指数，其中5 × 10⁷CFU · mL^-1菌液处理后其须根直径增长率高达78.57%，根系形态优于对照植株，促生效果明显。叶绿素荧光参数分析表明，不同浓度L19菌株发酵液处理后植株叶片的OJIP曲线在O、J（2 ms）、I（30 ms）、P点的荧光强度较对照均存在显著差异；施用不同浓度发酵液后，马铃薯叶片光合指数单位反应中心吸收光能（ABS/RC）、单位反应中心捕获用于还原Q_A能量（TR_o/RC）、单位面积的热耗散（DI_o/CS_o）分别增加了16.32% ~ 28.38%，说明L19发酵液能提高植株叶片的光能利用率。上述研究结果表明，L19分泌的次级代谢产物对马铃薯枯萎病菌具有生长抑制作用，同时发酵液还对马铃薯植株具有良好的促生效果，提高植株叶片的光合作用，具有良好的应用前景。

关键词: 马铃薯, 枯萎病, 尖孢镰刀菌, 解淀粉芽胞杆菌, 拮抗作用, 促生, 生物防治

Abstract:

In order to obtain a dual functional biocontrol strain with excellent antagonistic effect against Fusarium oxysporum in potato and promoting the growth of potato plants，the inhibition effect of secondary metabolites produced by strain L19 on the growth of Fusarium oxysporum was determined by the plate confrontation method. The lipopeptides and volatile compounds released by strain L19 were detected and identified by PCR and gas chromatography-mass spectrometry（GC-MS），respectively. The greenhouse pot experiment and chlorophyll fluorescence detection technology were used to explore the effect of strain L19 on the promotion of potato plants growth and leaf photosynthesis，respectively. The results showed that a total of 25 biocontrol strains were isolated，among which strain L19 could significantly inhibit the growth of Fusarium oxysporum hyphae. According to morphology and gyrB sequence analysis，the strain was identified as Bacillus amyloliquefaciens. The secondary metabolites produced by strain L19 caused the deformation of hyphae，such as twisting，adhesion，expansion and bubbling. The hyphae treated with B. amyloliquefaciens L19 metabolites lost penetration. With fermentation liquid concentration increasing，the spore germination rate of F. oxysporum was lower. B. amyloliquefaciens L19 contained lipopeptide synthesis genes such as sfp，ituA，fenB，bacD and mycB. Moreover，B. amyloliquefaciens L19 produced 12 volatile compounds，among them，the aromatic content accounted for 47.42%，followed by alcohols and ketones. 2-Ethyltoluene was the main volatile compound，with relative content of 20.91%. B. amyloliquefaciens L19 showed the ability to produce cellulase. The pot experiment showed that compared to the control group，the growth index of potato plants significantly increased after applying different concentrations of L19 fermentation broth. The growth rate of root circumference was as high as 78.57%，and the root morphology was better treated by 5 × 10⁷ CFU · mL^-1 L19 fermentation. The analysis of chlorophyll fluorescence parameters showed that there were significant differences in the fluorescence intensity of OJIP curve at O，J（2 ms），I（30 ms）and P points between the treated groups and the control groups. After the application of different concentrations of fermentation broth，the absorption of light energy per unit reaction center（ABS/RC），the energy captured by the unit reaction center for reduction of QA（TRo/RC），and the heat dissipation per unit area（DIo/CSo）of potato leaf photosynthetic index increased by 16.32%-28.38%，respectively，B. amyloliquefaciens L19 fermentation liquid can improve the light energy utilization rate of plant leaves. Therefore，the secondary metabolites secreted by B. amyloliquefaciens L19 showed strong inhibition effect on F. oxysporum，and the fermentation broth also had the growth promoting effect on potato plants. Moreover，B. amyloliquefaciens L19 could improve the photosynthesis of plant leaves and has good application prospects.

Key words: potato, Fusarium wilt, Fusarium oxysporum, Bacillus amyloliquefaciens, antagonism, growth promoting effect, biological control

强然, 张岱, 杨喆, 陈明玥, 赵景, 朱杰华. 解淀粉芽胞杆菌L19对马铃薯枯萎病菌的抑制及植株的促生作用[J]. 园艺学报, 2024, 51(4): 875-892.

QIANG Ran, ZHANG Dai, YANG Zhe, CHEN Mingyue, ZHAO Jing, ZHU Jiehua. Effects of Bacillus amyloliquefaciens‘L19’on Fusarium oxysporum Inhibition and Plants Growth Promotion in Potato[J]. Acta Horticulturae Sinica, 2024, 51(4): 875-892.

图/表 15

图1 对峙培养中菌株L19对马铃薯枯萎病原菌Fusarium oxysporum的抑制作用

Fig. 1 Inhibitory effect of strain L19 on Fusarium oxysporum in confrontation culture

表1 脂肽基因引物

Table 1 Lipopeptide gene primer

基因 Gene	引物序列（5′-3′） Primer sequences	扩增长度/bp Amplification growth
sfp	ATGAAGATTTACGGAATTTA；TTATAAAAGCTCTTCGTACG	675
ituA	ATGTATACCAGTCAATTCC；GATCCGAAGCTGACAATAG	1 150
fenB	CTATAGTTTGTTGACGGCTC；CAGCACTGGTTCTTGTCGCA	1 400
bacD	TCACCCGTTTCTGATTGCC；TTTGCGATGACCTTGATGTGG	540
mycB	ATGTCGGTGTTTAAAAATCAAGTAACG；TTAGGACGCCAGCAGTTCTTCTATTGA	2 024

图2 菌株L19菌落形态学（A）和革兰氏染色（B）鉴定

Fig. 2 Morphology of strain L19 colonies（A）and Gram stain（B）identification

图3 菌株L19 gyrB基因系统发育树

Fig. 3 Phylogenetic tree of strain L19 gyrB gene

图4 菌株L19发酵无菌上清液对尖孢镰刀菌菌丝的致畸作用

Fig. 4 The supernatant of strain L19 caused Fusarium oxysporum mycelial malformation

图5 菌株L19挥发性化合物对枯萎病原菌菌落及菌丝的影响

Fig. 5 Effect of volatile compounds of strain L19 on the colony and hypha of Fusarium wilt pathogen

图6 菌株L19挥发性气体对枯萎病原菌菌丝穿透性的影响

Fig. 6 Effect of volatiles of strain L19 on hyphal penetration of Fusarium wilt pathogen

图7 菌株L19次级代谢产物对尖孢镰刀菌孢子萌发的影响不同小写字母表示各处理间差异显著。

Fig. 7 Effect of secondary metabolite produced by strain L19 on spore germination rate of Fusarium oxysporum Different lowercase letters indicate significant difference between different treatments.

图8 菌株L19脂肽合成基因的PCR产物

Fig. 8 PCR product electrophoresis of lipopeptide synthesis gene of strain L19 M：DM2000 marker；1：sfp；2：ituA；3：fenB；4：bacD；5：mycB.

表2 菌株L19的GC-MS分析

Table 2 GC-MS analysis of strain L19

序号Code	化合物 Compound	CAS编号 CAS number	化学式 Chemical formula	保留时间/min Retention time	匹配度 Match	反匹配度 Inverse match	相对含量/% Relative content
1	4-辛醇 4-Octanol	589-62-8	C₈H₁₈O	16.66	883	1 359	1.53
2	环己醇 Cyclohexanol	108-93-0	C₆H₁₂O	17.16	878	1 374	1.29
3	1-辛醇 1-Octanol	111-87-5	C₈H₁₈O	22.16	933	1 530	9.82
4	1-癸醇 1-Decanol	112-30-1	C₁₀H₂₂O	28.16	952	1 732	5.79
5	正丙苯 N-propylbenzene	103-65-1	C₉H₁₂	10.76	903	1 187	1.91
6	3-乙基甲 3-Ethyltoluene	620-14-4	C₉H₁₂	11.26	893	1 202	12.60
7	1,3,5-三甲苯 1,3,5-Trimethylbenzene	108-67-8	C₉H₁₂	11.96	917	1 222	11.99
8	2-乙基甲苯1-Ethyl-2-methylbenzene	611-14-3	C₉H₁₂	14.96	930	1 308	20.91
9	4-甲基-2-庚醇 4-Methyl-2-heptanol	56298-90-9	C₈H₁₈O	15.86	879	1 335	1.25
10	2-壬酮 2-Nonanone	821-55-6	C₉H₁₈O	16.86	931	1 366	14.36
11	2-十一酮 2-Undecanone	112-12-9	C₁₁H₂₂O	23.66	932	1 577	16.36
12	9-十烯-1-醇9-Decenyl-1- alcohol	13019-22-2	C₁₀H₂₀O	29.66	904	1 787	2.18

图9 菌株L19产生的次级代谢产物纤维素酶（A）、蛋白酶（B）和果胶酶（C）的检测

Fig. 9 Detection of the secondary metabolites cellulase（A），protease（B）and pectinase（C）produced by strain L19

表3 不同浓度解淀粉芽胞杆菌L19发酵液灌根7 d后马铃薯各项生理指标

Table 3 Physiological indexes of potato after root irrigation with different concentrations of strain L19 fermentation broth for seven days

发酵液/（CFU · mL^-1） Fermentation broth	株高/cm Plant height	须根直径/mm Fibrous root diameter	地上部鲜质量/g Aboveground fresh mass	地下部鲜质量/g Underground fresh mass
0（H₂O对照Control）	27.40 ± 2.24 b	0.42 ± 0.05 b	46.74 ± 8.83 a	7.54 ± 4.09 a
5 × 10³	34.10 ± 4.39 a	0.64 ± 0.07 a	59.56 ± 14.37 a	9.80 ± 4.26 a
5 × 10⁵	35.50 ± 3.18 a	0.69 ± 0.14 a	62.34 ± 17.00 a	8.95 ± 2.38 a
5 × 10⁷	34.80 ± 2.14 a	0.75 ± 0.12 a	55.63 ± 6.38 a	8.21 ± 1.97 a

图10 不同浓度解淀粉芽胞杆菌L19发酵液对马铃薯须根的影响

Fig. 10 Effects of different concentrations of Bacillus amyloliquefaciens L19 fermentation broth on potato root circumference

图11 马铃薯幼苗经不同浓度L19发酵液灌根处理后叶片的OJIP曲线

Fig. 11 OJIP curves of potato seedlings after root irrigation with different concentrations of L19 fermentation broth

图12 不同浓度解淀粉芽胞杆菌L19发酵液灌根对马铃薯叶片叶绿素荧光参数的影响

Fig. 12 Effects of different concentrations of Bacillus amyloliquefaciens L19 fermentation broth on chlorophyll fluorescence parameters of potato leaves

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