连作甜瓜蔓根真菌病害发生与放线菌生物防治研究
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摘要
连作甜瓜蔓根真菌病害是甜瓜设施栽培和专业化种植基地发展的主要限制因素，寻找合适有效的病害防治途径是目前解决甜瓜连作障碍问题的首要任务。国内外关于甜瓜病害防治采取的措施主要有化学农药、嫁接、抗性品种筛选等，但效果均不明显且具有各自的缺点和弊端，如环境污染、农药残留、品质下降、病原菌易变异或者产生抗药性等。随着绿色农业和有机果蔬的兴起，寻找新的环境友好型病害防治措施开始受到人们的广泛关注。“以菌治菌”的生物防治思路有望从源头解决问题。放线菌具有产多种抗生素的功能，能够抑制动物病原菌、防治人畜传染性病害，已在医学上得到广泛应用。放线菌产生的农用抗生素在农业生产上也表现出良好的效果，但放线菌活菌制剂在作物真菌病害防治上的应用研究相对滞后，其作用机理及效果目前仍不清楚。放线菌活菌制剂在甜瓜蔓根真菌病害生物防治上的研究更少。本论文通过对连作甜瓜蔓根真菌病害发生微生态机制，真菌病害生防放线菌筛选鉴定，放线菌防病促生效果，放线菌生防机理以及病原真菌-生防放线菌-甜瓜植株互作机理的系统研究，为防治甜瓜蔓根真菌病害、克服甜瓜连作障碍提供高效生防放线菌菌株，并为后期生防放线菌活菌制剂的商品化生产和实际应用提供理论依据。论文主要研究结果如下：
（1）连作甜瓜蔓根真菌病害发生的微生态机制
连作甜瓜蔓根真菌病害发生时，发病植株及其根际病土中存在多种真菌，包括直接致病真菌、具有侵染性且产生有害毒素但不直接致病真菌以及试验条件下不致病真菌3种类型。从陕西省西安市阎良区连作甜瓜病株及其病土中共获得13株真菌分离物，经形态学和rDNA-ITS序列分析初步鉴定为4个属8个种，包括：甜瓜球腔菌（Didymellabryoniae）、木贼镰刀菌（Fusarium equiseti）、层出镰刀菌（Fusarium proliferatum）、腐皮镰刀菌（Fusarium solani）、葡萄座腔菌（Botryosphaeria dothidea）、细极链格孢菌（Alternaria alternata）、三线镰刀菌（Fusarium tricinctum）和厚垣镰孢菌（Fusariumchlamydosporum）。其中D. bryoniae为甜瓜蔓枯病专性致病菌，F. equiseti、F. proliferatum和F. solani对甜瓜离体根茎段具有侵染性。F. equiseti和F. proliferatum虽未表现出直接致病性，但可通过分泌毒素对甜瓜产生化感危害。其它真菌分离物在试验条件下不致病，但是否产生毒素尚不清楚。
（2）生防放线菌筛选及盆栽防病促生效果
放线菌A12、C28、C13和C27及其无菌发酵滤液对甜瓜球腔菌（D. bryoniae）具有良好的皿内抑制作用。放线菌A12和C28无菌发酵滤液喷施处理能明显降低盆栽甜瓜蔓枯病病害严重程度（DS）及AUDPC值；放线菌C13和C27固态发酵制剂拌土处理，对甜瓜蔓枯病亦表现出良好防效。供试放线菌能明显促进盆栽甜瓜植株生长，在甜瓜植株根区、根表的定殖数约为105～106cfu·g-1干土。放线菌A1、A11和A12能明显抑制木贼镰刀菌（F. equiseti）菌丝生长，其无菌发酵滤液能促进甜瓜胚轴、胚根生长，提高种子简明活力指数。三株放线菌在适宜接种量下与F. equiseti混接，甜瓜叶片叶绿素含量、PPO活性，根系质量、活力均高于未接种对照或者F. equiseti单接。
（3）生防放线菌鉴定
甜瓜蔓枯病生防放线菌A12、C28、C13和C27被依次鉴定为链霉菌属的密旋链霉菌（Streptomyces pactum）、球孢链霉菌球孢亚种（Streptomyces globisporus subsp.globisporus）、纤维黄链霉菌（Streptomyces celluloflavus）和黄三素链霉菌（Streptomycesflavotricini）。两株对F. equiseti具有良好拮抗性的放线菌A1和A11，被初步鉴定为加州链霉菌（Streptomyces californicus）和肉质链霉菌（Streptomyces carnosus）。
（4）生防放线菌作用机理
生防放线菌主要通过5种作用机制对以蔓根真菌病害为主的甜瓜连作障碍进行修复：通过抗菌活性物质抑制病原真菌生长；利用病原真菌菌体的诱导作用合成纤维素酶、几丁质酶及葡聚糖酶等胞外酶，通过酶溶作用破坏病原真菌菌丝体，抑制病原真菌生长；通过分泌植物激素类物质促进甜瓜植株生长，提高果实产量和品质，改善植株生理生化特性；通过提高植物保护性诱导酶活性诱导植株产生系统抗病性，增强植物抗病性及抗逆能力；通过根际土壤微生物区系修复调整植株根际土壤微生态平衡。
生防放线菌A12、C28、C13和C27在温室条件下能促进甜瓜植株生长，增加果实产量，并能提高甜瓜果实可溶性蛋白、维生素C、可溶性糖和固形物等品质指标。放线菌A12和C28处理甜瓜叶片净光合速率提高，有利于光合产物累积。放线菌C13处理，甜瓜植株根系N、P、K含量增加，说明营养元素的吸收能力增强。
生防菌剂C28和康照在大田条件下对甜瓜植株具有明显的促生作用。两种菌剂育苗接种、育苗移栽双重接种均能提高甜瓜诱导酶活性和可溶性蛋白含量，降低丙二醛含量。生防菌剂处理，甜瓜诱导抗病性增强，膜脂过氧化程度减弱，有利于植株形成抵御外界病害侵染的系统保护屏障。生防菌剂育苗接种、育苗移栽双重接种配施微量元素能促进甜瓜植株生长，对甜瓜抗病性和抗逆性影响不明显或具有负效应。
生防放线菌A12、C28、C13和C27对温室甜瓜根区和根表土壤细菌、放线菌生长繁殖具有明显促进作用。放线菌A12、C28和C27处理，甜瓜植株根区土壤病原性真菌数量降低。甜瓜根域土壤微生物由“真菌型”向“细菌型”转变，说明生防放线菌具有调节和改良甜瓜根域土壤微生态、降低真菌病害发生风险的效应。
（5）病原真菌-生防放线菌-甜瓜植株互作机理
瓜类作物病原真菌菌体可诱导供试生防放线菌合成并分泌纤维素酶、几丁质酶及葡聚糖酶等胞外酶，进而溶解病原真菌菌体，即通过酶溶抑菌作用抑制病原真菌生长繁殖。层出镰刀菌（F. proliferatum）菌体作为唯一碳源，能够明显增强拮抗放线菌产纤维素酶活性。以木贼镰刀菌（F. equiseti）菌体为唯一碳源时，放线菌A12和C28的几丁质酶活性较高。
具有侵染性的非致病真菌能够通过分泌毒素对甜瓜植株产生双重化感作用。木贼镰刀菌（F. equiseti）和层出镰刀菌（F. proliferatum）产生的毒素能明显抑制甜瓜种子萌发和胚轴、胚根生长，引起甜瓜幼苗根系丙二醛、可溶性蛋白含量及细胞膜相对透性增加。但两株镰刀菌毒素在一定稀释倍数下对甜瓜幼苗根系诱导酶活性具有提高作用，即能增强甜瓜植株系统抗病性。
Fungal diseases of the tendril and root of continuous cropping melon plants are themajor limiting factors of the development of melon protected cultivation and specializedplanting base. Finding effective disease control approaches is the first priority to solve theseproblems. The current measures for controlling melon fungal diseases at home and abroadmainly include chemical pesticides, grafting, resistant varieties screening and so on. Thesemeasures show little effect, while have their own shortcomings and drawbacks, such asenvironment pollution, pesticide residue, pathogens variation and drug resistance. With thedevelopment of green agriculture and organic fruit and vegetables, the new environment-friendly measures such as biocontrol methods attract widespread attention, and will be themain trend in current and future. The biocontrol idea ‘to prevent and cure fungal diseases byantagonistic microorganisms’ was a new way to solve these problems. Actinomycetes werewidely used in medicine for their inhibition effect on animal pathogens, human and livestockdiseases via antibiotics production. The agricultural antibiotics also played an important partin agricultural production, while the applications of live actinomycetes preparations on plantsfungal diseases control were somewhat slow, their action mechanisms and effects remainedunclear. There were still few researches about the biocontrol potential of live actinomycetespreparations on fungal diseases of melon tendril and root.
In this paper, the author systematically studied the microecology mechnism of theoccurrence of melon fungal diseases, screening and identification of antagonisticactinomycetes, biocontrol and growth promotion effect of actinomycetes, biocontrolmechnism as well as the interaction mechnism among pathogenic fungi, biocontrolactinomycetes and melon plants. Biocontrol actinomycetes with high efficiency and stabilitywere obtained for controlling melon fungal diseases and overcoming melon continuouscropping obstacles. The related studies about biocontrol mechanisms provided solid scientific basis for the commercially production and practical application of the biocontrol preparationsin the near future. The main contributions of this research were summarized as follows:
(1) Microecology mechnism of fungal diseases occurrence in tendril and root ofcontinuous cropping melon plants
Various fungi were obtained from the infected melon plants and their rhizosphere soils,when the fungal diseases of tendril and root of continuous cropping melon plants occurred,which including direct pathogenic fungi, indirect pathogenic fungi but with infection andtoxin action, as well as indirect pathogenic fungi. From the infected melon plants and theirrhizosphere soils,13fungi isolates were collected and further attributed to4genus and8species, according to morphological characters and rDNA-ITS sequence analysis. Thesuperior fungi mainly included Didymella bryoniae, Fusarium equiseti, Fusariumproliferatum, Fusarium solani, Botryosphaeria dothidea, Alternaria alternata, Fusariumtricinctum and Fusarium chlamydosporum. Among which, D. bryoniae was the pathogencaused melon gummy stem blight, F. equiseti and F. proliferatum showed melon root andstem infection ability in vitro. Other fungi isolates had no pathogenicity to melon plants, theirtoxin production ability were unknown.
(2) Disease biocontrol and growth promotion effect of antagonistic actinomycetes
Actinomycetes A12, C28, C13, C27and their sterile culture filtrates showed favorableantagonistic effect against D. bryoniae in Petri dish. Sterile culture filtrates of actinomycetesA12and C28expressed significant reduction effect on the disease severity (DS) and AUDPCvalue of melon plants; solid-state fermentation preparations of actinomycetes C13and C27also showed obvious biocontrol effect on melon gummy stem blight. The four actinomycetesdisplayed definite growth promotion effect and substantial survival ability (105～106cfu·g-1dry soil) in the rhizosphere and on the rhizoplane of melon plants in pots. Actinomycetes A1,A11and A12showed evident mycelia inhibition effect on F. equiseti. The hypocotyl, radiclelength, and simple vigor index of melon seeds were all increased by their sterile culturefiltrates. Compared with the control or F. equiseti single inoculation, the chlorophyll contentsand PPO activity of melon leaves, root weights and activities were all improved, when F.equiseti were inoculated with the3antagonistic actinomycetes.
(3) Identification of the biocontrol actinomycetes
Actinomycetes A12, C28, C13and C27, with biocontrol effect on melon gummy stemblight, were subsequently identified as Streptomyces pactum, Streptomyces globisporus subsp.globisporus, Streptomyces celluloflavus and Streptomyces flavotricini. Actinomycetes A1andA11, with obvious antagonistic effect against F. equiseti, were identified as Streptomycescalifornicus and Streptomyces carnosus, respectively.
(4) Biocontrol mechnisms of the actinomycetes
The biocontrol actinomycetes showed good repair effect on the continuous croppingobstacle of melon plants from the following5action mechnisms: fungal mycelia growthinhibition effect via antimicrobial active substances; extracellular enzymes production andfungal mycelia degradation effect through these enzymes; growth and yield promotion, fruitquality improvement, as well as plant physiological and biochemical metabolism regulationby means of hormone-like substances; plant systematic disease resistance induction effect viaprotective enzymes improvement; rhizosphere soil microecosystem repair and regulation.
Biocontrol actinomycetes A12, C28, C13and C27showed favorable growth and yieldpromotion effect on melon plants in greenhouse. Fruit quality such as soluble protein, vitaminC, soluble sugar and solids contents were improved in the presence of these actinomycetes.The net photosynthetic rate was also increased by A12and C28treatments, which waspropitious to the accumulation of photosynthetic product. The root absorption ability ofnutrient elements N, P, K was improved by actinomycete C13treatment.
Two biocontrol preparations C28and Kangzhao showed obvious growth promotioneffect on melon plants in field. The induced enzymes activities and soluble protein contentsof melon plants were improved, and MDA contents were decreased by biocontrolpreparations treatments. With the enhanced disease resistance and decreased lipidperoxidation, melon plants have formed a protective barrier to resist disease attack. Themicroelements admixture of zine, manganese and titanium displayed significantly growthpromotion effect on melon plants, when biocontrol preparations C28and Kangzhao wereapplied. Application of microelements admixture with biocontrol preparations showed littledistinct variety even negative effect on the disease resistance of melon plants.
There was an upward trend about the soil bacteria and actinomycetes amounts in therhizosphere and on the rhizoplane soil of melon plants after treated by biocontrolactinomycetes A12, C28, C13and C27. The pathogenic fungi quantities in the rhizospheresoil of melon plants were decreased by A12and C28treatments. Microorganisms in the rootdomain soil of melon plants were transited from “fungal-type” to “bacteria-type”, whichmeant the biocontrol actinomycetes displayed a positive effect on the regulation of soilmicroecosystem.
(5) Interaction mechnism among pathogenic fungi, biocontrol actinomycetes and melonplants
Mycelia preparation of the pathogenic fungi (MPPF) of cucurbit plants showedsynchronous induction effect on extracellular enzymes produced by antagonisitcactinomycetes, fungal mycelia degadation and inhibition were the combined action of these enzymes. The MPPF served as good carbon source to enhance cellulase activities, especiallythose from F. proliferatum. The MPPF from F. equiseti obviously improved chitinaseactivities of actinomycetes A12and C28.
Indirect pathogenic fungi with infection ability showed double allelopathic effect onmelon plants via toxin production. F. equiseti and F. proliferatum showed obvious inhibitioneffect on seeds germination and the growth of melon plants. The MDA, soluble proteincontents and cell membrane permeability of melon roots were increased in the presence ofFusarium toxins. The induced protective enzymes of melon roots were improved whentreated by the toxins at stated dilutions, which strengthened the disease resistance of melonplants.
（1）连作甜瓜蔓根真菌病害发生的微生态机制
连作甜瓜蔓根真菌病害发生时，发病植株及其根际病土中存在多种真菌，包括直接致病真菌、具有侵染性且产生有害毒素但不直接致病真菌以及试验条件下不致病真菌3种类型。从陕西省西安市阎良区连作甜瓜病株及其病土中共获得13株真菌分离物，经形态学和rDNA-ITS序列分析初步鉴定为4个属8个种，包括：甜瓜球腔菌（Didymellabryoniae）、木贼镰刀菌（Fusarium equiseti）、层出镰刀菌（Fusarium proliferatum）、腐皮镰刀菌（Fusarium solani）、葡萄座腔菌（Botryosphaeria dothidea）、细极链格孢菌（Alternaria alternata）、三线镰刀菌（Fusarium tricinctum）和厚垣镰孢菌（Fusariumchlamydosporum）。其中D. bryoniae为甜瓜蔓枯病专性致病菌，F. equiseti、F. proliferatum和F. solani对甜瓜离体根茎段具有侵染性。F. equiseti和F. proliferatum虽未表现出直接致病性，但可通过分泌毒素对甜瓜产生化感危害。其它真菌分离物在试验条件下不致病，但是否产生毒素尚不清楚。
（2）生防放线菌筛选及盆栽防病促生效果
放线菌A12、C28、C13和C27及其无菌发酵滤液对甜瓜球腔菌（D. bryoniae）具有良好的皿内抑制作用。放线菌A12和C28无菌发酵滤液喷施处理能明显降低盆栽甜瓜蔓枯病病害严重程度（DS）及AUDPC值；放线菌C13和C27固态发酵制剂拌土处理，对甜瓜蔓枯病亦表现出良好防效。供试放线菌能明显促进盆栽甜瓜植株生长，在甜瓜植株根区、根表的定殖数约为105～106cfu·g-1干土。放线菌A1、A11和A12能明显抑制木贼镰刀菌（F. equiseti）菌丝生长，其无菌发酵滤液能促进甜瓜胚轴、胚根生长，提高种子简明活力指数。三株放线菌在适宜接种量下与F. equiseti混接，甜瓜叶片叶绿素含量、PPO活性，根系质量、活力均高于未接种对照或者F. equiseti单接。
（3）生防放线菌鉴定
甜瓜蔓枯病生防放线菌A12、C28、C13和C27被依次鉴定为链霉菌属的密旋链霉菌（Streptomyces pactum）、球孢链霉菌球孢亚种（Streptomyces globisporus subsp.globisporus）、纤维黄链霉菌（Streptomyces celluloflavus）和黄三素链霉菌（Streptomycesflavotricini）。两株对F. equiseti具有良好拮抗性的放线菌A1和A11，被初步鉴定为加州链霉菌（Streptomyces californicus）和肉质链霉菌（Streptomyces carnosus）。
（4）生防放线菌作用机理
生防放线菌主要通过5种作用机制对以蔓根真菌病害为主的甜瓜连作障碍进行修复：通过抗菌活性物质抑制病原真菌生长；利用病原真菌菌体的诱导作用合成纤维素酶、几丁质酶及葡聚糖酶等胞外酶，通过酶溶作用破坏病原真菌菌丝体，抑制病原真菌生长；通过分泌植物激素类物质促进甜瓜植株生长，提高果实产量和品质，改善植株生理生化特性；通过提高植物保护性诱导酶活性诱导植株产生系统抗病性，增强植物抗病性及抗逆能力；通过根际土壤微生物区系修复调整植株根际土壤微生态平衡。
生防放线菌A12、C28、C13和C27在温室条件下能促进甜瓜植株生长，增加果实产量，并能提高甜瓜果实可溶性蛋白、维生素C、可溶性糖和固形物等品质指标。放线菌A12和C28处理甜瓜叶片净光合速率提高，有利于光合产物累积。放线菌C13处理，甜瓜植株根系N、P、K含量增加，说明营养元素的吸收能力增强。
生防菌剂C28和康照在大田条件下对甜瓜植株具有明显的促生作用。两种菌剂育苗接种、育苗移栽双重接种均能提高甜瓜诱导酶活性和可溶性蛋白含量，降低丙二醛含量。生防菌剂处理，甜瓜诱导抗病性增强，膜脂过氧化程度减弱，有利于植株形成抵御外界病害侵染的系统保护屏障。生防菌剂育苗接种、育苗移栽双重接种配施微量元素能促进甜瓜植株生长，对甜瓜抗病性和抗逆性影响不明显或具有负效应。
生防放线菌A12、C28、C13和C27对温室甜瓜根区和根表土壤细菌、放线菌生长繁殖具有明显促进作用。放线菌A12、C28和C27处理，甜瓜植株根区土壤病原性真菌数量降低。甜瓜根域土壤微生物由“真菌型”向“细菌型”转变，说明生防放线菌具有调节和改良甜瓜根域土壤微生态、降低真菌病害发生风险的效应。
（5）病原真菌-生防放线菌-甜瓜植株互作机理
瓜类作物病原真菌菌体可诱导供试生防放线菌合成并分泌纤维素酶、几丁质酶及葡聚糖酶等胞外酶，进而溶解病原真菌菌体，即通过酶溶抑菌作用抑制病原真菌生长繁殖。层出镰刀菌（F. proliferatum）菌体作为唯一碳源，能够明显增强拮抗放线菌产纤维素酶活性。以木贼镰刀菌（F. equiseti）菌体为唯一碳源时，放线菌A12和C28的几丁质酶活性较高。
具有侵染性的非致病真菌能够通过分泌毒素对甜瓜植株产生双重化感作用。木贼镰刀菌（F. equiseti）和层出镰刀菌（F. proliferatum）产生的毒素能明显抑制甜瓜种子萌发和胚轴、胚根生长，引起甜瓜幼苗根系丙二醛、可溶性蛋白含量及细胞膜相对透性增加。但两株镰刀菌毒素在一定稀释倍数下对甜瓜幼苗根系诱导酶活性具有提高作用，即能增强甜瓜植株系统抗病性。
Fungal diseases of the tendril and root of continuous cropping melon plants are themajor limiting factors of the development of melon protected cultivation and specializedplanting base. Finding effective disease control approaches is the first priority to solve theseproblems. The current measures for controlling melon fungal diseases at home and abroadmainly include chemical pesticides, grafting, resistant varieties screening and so on. Thesemeasures show little effect, while have their own shortcomings and drawbacks, such asenvironment pollution, pesticide residue, pathogens variation and drug resistance. With thedevelopment of green agriculture and organic fruit and vegetables, the new environment-friendly measures such as biocontrol methods attract widespread attention, and will be themain trend in current and future. The biocontrol idea ‘to prevent and cure fungal diseases byantagonistic microorganisms’ was a new way to solve these problems. Actinomycetes werewidely used in medicine for their inhibition effect on animal pathogens, human and livestockdiseases via antibiotics production. The agricultural antibiotics also played an important partin agricultural production, while the applications of live actinomycetes preparations on plantsfungal diseases control were somewhat slow, their action mechanisms and effects remainedunclear. There were still few researches about the biocontrol potential of live actinomycetespreparations on fungal diseases of melon tendril and root.
In this paper, the author systematically studied the microecology mechnism of theoccurrence of melon fungal diseases, screening and identification of antagonisticactinomycetes, biocontrol and growth promotion effect of actinomycetes, biocontrolmechnism as well as the interaction mechnism among pathogenic fungi, biocontrolactinomycetes and melon plants. Biocontrol actinomycetes with high efficiency and stabilitywere obtained for controlling melon fungal diseases and overcoming melon continuouscropping obstacles. The related studies about biocontrol mechanisms provided solid scientific basis for the commercially production and practical application of the biocontrol preparationsin the near future. The main contributions of this research were summarized as follows:
(1) Microecology mechnism of fungal diseases occurrence in tendril and root ofcontinuous cropping melon plants
Various fungi were obtained from the infected melon plants and their rhizosphere soils,when the fungal diseases of tendril and root of continuous cropping melon plants occurred,which including direct pathogenic fungi, indirect pathogenic fungi but with infection andtoxin action, as well as indirect pathogenic fungi. From the infected melon plants and theirrhizosphere soils,13fungi isolates were collected and further attributed to4genus and8species, according to morphological characters and rDNA-ITS sequence analysis. Thesuperior fungi mainly included Didymella bryoniae, Fusarium equiseti, Fusariumproliferatum, Fusarium solani, Botryosphaeria dothidea, Alternaria alternata, Fusariumtricinctum and Fusarium chlamydosporum. Among which, D. bryoniae was the pathogencaused melon gummy stem blight, F. equiseti and F. proliferatum showed melon root andstem infection ability in vitro. Other fungi isolates had no pathogenicity to melon plants, theirtoxin production ability were unknown.
(2) Disease biocontrol and growth promotion effect of antagonistic actinomycetes
Actinomycetes A12, C28, C13, C27and their sterile culture filtrates showed favorableantagonistic effect against D. bryoniae in Petri dish. Sterile culture filtrates of actinomycetesA12and C28expressed significant reduction effect on the disease severity (DS) and AUDPCvalue of melon plants; solid-state fermentation preparations of actinomycetes C13and C27also showed obvious biocontrol effect on melon gummy stem blight. The four actinomycetesdisplayed definite growth promotion effect and substantial survival ability (105～106cfu·g-1dry soil) in the rhizosphere and on the rhizoplane of melon plants in pots. Actinomycetes A1,A11and A12showed evident mycelia inhibition effect on F. equiseti. The hypocotyl, radiclelength, and simple vigor index of melon seeds were all increased by their sterile culturefiltrates. Compared with the control or F. equiseti single inoculation, the chlorophyll contentsand PPO activity of melon leaves, root weights and activities were all improved, when F.equiseti were inoculated with the3antagonistic actinomycetes.
(3) Identification of the biocontrol actinomycetes
Actinomycetes A12, C28, C13and C27, with biocontrol effect on melon gummy stemblight, were subsequently identified as Streptomyces pactum, Streptomyces globisporus subsp.globisporus, Streptomyces celluloflavus and Streptomyces flavotricini. Actinomycetes A1andA11, with obvious antagonistic effect against F. equiseti, were identified as Streptomycescalifornicus and Streptomyces carnosus, respectively.
(4) Biocontrol mechnisms of the actinomycetes
The biocontrol actinomycetes showed good repair effect on the continuous croppingobstacle of melon plants from the following5action mechnisms: fungal mycelia growthinhibition effect via antimicrobial active substances; extracellular enzymes production andfungal mycelia degradation effect through these enzymes; growth and yield promotion, fruitquality improvement, as well as plant physiological and biochemical metabolism regulationby means of hormone-like substances; plant systematic disease resistance induction effect viaprotective enzymes improvement; rhizosphere soil microecosystem repair and regulation.
Biocontrol actinomycetes A12, C28, C13and C27showed favorable growth and yieldpromotion effect on melon plants in greenhouse. Fruit quality such as soluble protein, vitaminC, soluble sugar and solids contents were improved in the presence of these actinomycetes.The net photosynthetic rate was also increased by A12and C28treatments, which waspropitious to the accumulation of photosynthetic product. The root absorption ability ofnutrient elements N, P, K was improved by actinomycete C13treatment.
Two biocontrol preparations C28and Kangzhao showed obvious growth promotioneffect on melon plants in field. The induced enzymes activities and soluble protein contentsof melon plants were improved, and MDA contents were decreased by biocontrolpreparations treatments. With the enhanced disease resistance and decreased lipidperoxidation, melon plants have formed a protective barrier to resist disease attack. Themicroelements admixture of zine, manganese and titanium displayed significantly growthpromotion effect on melon plants, when biocontrol preparations C28and Kangzhao wereapplied. Application of microelements admixture with biocontrol preparations showed littledistinct variety even negative effect on the disease resistance of melon plants.
There was an upward trend about the soil bacteria and actinomycetes amounts in therhizosphere and on the rhizoplane soil of melon plants after treated by biocontrolactinomycetes A12, C28, C13and C27. The pathogenic fungi quantities in the rhizospheresoil of melon plants were decreased by A12and C28treatments. Microorganisms in the rootdomain soil of melon plants were transited from “fungal-type” to “bacteria-type”, whichmeant the biocontrol actinomycetes displayed a positive effect on the regulation of soilmicroecosystem.
(5) Interaction mechnism among pathogenic fungi, biocontrol actinomycetes and melonplants
Mycelia preparation of the pathogenic fungi (MPPF) of cucurbit plants showedsynchronous induction effect on extracellular enzymes produced by antagonisitcactinomycetes, fungal mycelia degadation and inhibition were the combined action of these enzymes. The MPPF served as good carbon source to enhance cellulase activities, especiallythose from F. proliferatum. The MPPF from F. equiseti obviously improved chitinaseactivities of actinomycetes A12and C28.
Indirect pathogenic fungi with infection ability showed double allelopathic effect onmelon plants via toxin production. F. equiseti and F. proliferatum showed obvious inhibitioneffect on seeds germination and the growth of melon plants. The MDA, soluble proteincontents and cell membrane permeability of melon roots were increased in the presence ofFusarium toxins. The induced protective enzymes of melon roots were improved whentreated by the toxins at stated dilutions, which strengthened the disease resistance of melonplants.
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