蚕豆油壶菌火肿病（Olpidium viciae Kusano）研究
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摘要
蚕豆油壶菌火肿病(Broad bean blister),由野豌豆油壶菌(Olpidium viciae Kusano)引起,病部细胞增生呈瘤状突起。蚕豆油壶菌火肿病仅在日本东京和中国四川及邻近省的局部地区有发生,该病已成为四川西北高原地区阿坝藏族自治州及邻近地区蚕豆生产上的重要病害。迄今较深入的研究论文仅有3篇,其中日本真菌学家Shunsuke Kusano在1912年对野豌豆油壶菌的生活史和细胞学、野豌豆油壶菌与寄主的关系进行了较系统研究；1984年辛哲生报道了四川省松潘县蚕豆火肿病发生危害与防治的初步研究结果。蚕豆油壶菌火肿病的其它研究未见报道,为此对该病害开展了较为系统深入的研究,并取得了明显进展,在病原生物学、病理组织学、病理生理学和分子生物学研究方面有所创新。
1.蚕豆火肿病的症状
蚕豆火肿病主要危害蚕豆叶和茎,也危害叶柄和果柄,但不危害果荚。病部最初出现稍褪绿的近圆形斑痕,随病情发展病斑表面稍显粗糙,并不断向上增生隆起,形成小的瘤状突起。病瘤单生或群生,可相互愈合成不规则状。受害严重时,病叶畸形,植株矮小。病部形成小的瘤状突起、表面稍显粗糙,是本病的标志性特征,是诊断本病的重要依据。本研究完善了蚕豆油壶菌火肿病的症状特征,明确了蚕豆豆荚上的疱状突起不是本病的症状,从而为准确诊断和防冶蚕豆油壶菌火肿病提供了科学依据。
2.蚕豆火肿病的病原
蚕豆火肿病病原菌寄生在蚕豆叶、茎和果柄表皮细胞内。营养体为单细胞圆形原生质团,营养体整体产果。无性繁殖结构为游动孢子囊,一般为球形,大小变化较大,一般为12.95～62.16μm,平均28.84μm。游动孢子囊萌发产生游动孢子,通过排孢管(孔)释放。游动孢子卵圆形或球形,大小为4.1-5μm,平均为4.47μm。尾部具一根鞭毛,鞭毛长24-3lμm,平均为26.56μm。游动孢子可作为同形游动配子结合,形成双鞭毛圆形的接合子。接合子侵染蚕豆,在表皮细胞内形成休眠孢子囊。休眠孢子囊圆形厚壁,形状大小较一致,大小为19-31μm,平均为25.51μm,平均壁厚3.81μm。一个寄主细胞中的数目可多达60个以上,休眠孢子囊大多形成在病害后期。对该菌的形态学研究表明：四川蚕豆火肿病是由野豌豆油壶菌(Olpidium viciae Kusano)引起。该菌的分类地位属鞭毛菌亚门、壶菌目、油壶菌科、油壶菌属。
对蚕豆火肿病菌的生物学研究结果表明：病斑表皮组织中的游动孢子囊成熟后,遇雨水即可萌发,释放出游动孢子。游动孢子囊萌发的温度范围为0-18。C,在18℃下萌发就受到明显抑制,在20℃、25℃和30。C下不能萌发。光照或黑暗对游动孢子囊萌发没有明显影响。K、Na盐离子溶液对孢子囊萌发有一定抑制作用,浓度分别达到80mmo1/L、100mmol/L时,游动孢子囊萌发就几乎被抑制。pH对游动孢子囊萌发也有一定影响,萌发的pH范围为pH5-8,以pH7-7.5最适。温度影响游动孢子的游动时间长短,在0-30℃范围内,随温度升高,游动时间逐渐缩短。在0-5℃下保持游动时间最长可达72h以上,10℃下可达24h,15℃和20℃下可保持10h,25℃下为3h,30℃下仅可保持游动5min。温度对游动孢子侵染及发病有明显影响,在5℃下能成功侵染,但不产生明显症状。侵染和引起发病的温度范围为10～25℃,30。C及以上温度不能侵染和发病。最适侵染和发病的温度为10-20℃。接种游动孢子引起侵染和发病的保湿时间为12h以上。叶龄影响病菌的侵染和发病,幼叶有利侵染和发病。
病组织中的休眠孢子囊必须经过越夏和越冬后,有水存在时才能萌发产生游动孢子。萌发的温度范围为5-30℃,在17℃左右温度下,经4h即可产生游动孢子。病组织中的休眠孢子囊萌发极不整齐,休眠孢子囊释放游动孢子的方式与游动孢子囊的释放方式相同。
3.蚕豆火肿病的发生规律
四川蚕豆火肿病的发生与分布局限在四川西北部高原的阿坝藏族自治州海拔2400m以上的春播蚕豆生态区。病害的循环和发病特点是：病菌以休眠孢子囊随病残体在土壤中越夏和越冬,作为初侵染源。次年春播蚕豆后,休眠孢子囊萌发产生游动孢子侵染蚕豆幼苗,病害始发期在5月。田间发病后,病菌不断产生游动孢子囊,在有雨露时游动孢子囊萌发释放游动孢子进行再侵染。在蚕豆生长期中病菌的再侵染频繁,病害在田间扩展迅速,在蚕豆进入开花和始荚期,病害进入高峰期；蚕豆进入结荚期后,病菌形成休眠孢子囊,病害发展趋于停止。病菌的游动孢子藉风雨在田间近距离传播。连作和施用带有病残体的厩肥有利于发病。目前生产上种植和供抗性鉴定的40多个蚕豆材料均不同程度感病,缺乏抗病品种。病害传播风险的初步结果表明：病区当年的病残体菌源对盆地内秋播蚕豆区不构成侵染发病的危险。
4.蚕豆火肿病的组织病理学
野豌豆油壶菌侵入蚕豆叶和茎的表皮细胞后,定殖在侵入的细胞内,菌体生长发育,整体产果,形成游动孢子囊和休眠孢子囊。病菌的侵染刺激了侵染点邻近的表皮细胞和其下的叶肉组织或茎的皮层组织细胞,这些已分化的组织细胞转为分生细胞,进行分裂,分裂方式为平周分裂。原有的正常基本组织细胞被未高度分化的薄壁组织细胞取代。这些细胞具有分生能力并不断分裂,细胞体积增大,造成蚕豆病组织细胞发生畸形增生的病理解剖改变,病部呈瘤状隆起。这是蚕豆油壶菌火肿病症状产生的病理组织学机制,也反映了野豌豆油壶菌和蚕豆在细胞和组织水平上的相互作用。
5.蚕豆火肿病的病理生理
蚕豆火肿病的症状特点主要是受病部位产生瘤状突起,病部细胞逐渐褐变坏死。这种畸形的病理改变预示着受病菌侵染后,寄主组织中的植物激素发生了异常变化；受病组织逐渐褐变坏死,显示抗氧化保护酶活力改变,细胞受到伤害,最终坏死。为深入了解蚕豆火肿病发生病理改变的生理机制,测定了病组织中的植物激素含量、抗氧化保护酶活性和丙二醛含量。结果表明,病害初期组织中的ZT和KT含量大幅上升,IAA和GA。含量也有一定程度增加,它们共同刺激蚕豆病部细胞大量分裂、增生和体积逐渐膨大,病部开始隆起；在病害初期ABA含量也大量增加,表现为蚕豆幼苗被害,植株矮化畸形、生育期严重滞后,而成株被害,受病叶片提前衰老、易脱落。在病害中后期组织中的GA3含量急剧增加,是健康组织的3倍,ZT和KT仍维持一定幅度增量,但增量率明显降低,使得病害早期大量分裂增生的细胞在病害中后期迅速膨大,病部异常隆起；ABA在病害中后期也有较大增量,使病部细胞褪绿褐变,最终坏死,受病叶片脱落；而IAA在病害中后期呈现负增长,这是寄主感病的生理表现。受病组织中抗氧化保护酶活力测定的结果表明：病、健组织中的SOD、POD活力在病害初期没有明显差异,在病害中后期病组织均显著高于健康组织,说明病菌侵染破坏了寄主本身正常活性氧消除机制的平衡,导致体内活性氧积累,诱导抗氧化保护酶SOD和POD活性增强,是感病的生化表现。而CAT活力在病程初期和中后期,病组织均极显著低于健康组织。己知CAT和H202具有较高的亲和力,CAT活力降低导致组织中的H202积累,加重对组织的损伤。组织中的MDA含量在病害中后期极显著高于健康组织,说明病部组织细胞生物膜损伤加剧,这种不可逆的伤害使细胞最终破坏解体、组织溃烂。这些结果反映了蚕豆火肿病症状产生的病理生理机制,也揭示了蚕豆火肿病病程中组织细胞内主要抗氧化保护酶的病理变化特点。
6.野豌豆油壶菌的分子生物学研究
通过提取病斑组织的总DNA,采用真核生物内转录间隔区通用引物ITS1/ITS4进行扩增,获得了病菌的ITS序列；回收此序列测序,GenBank登记号为HQ677595.1,补充了野豌豆油壶菌的分子生物学研究资料。将病菌与其它真菌进行比对,显示与甘蓝油壶菌(Olpidium brassicae)和瓜类油壶菌(Olpidium bornovanus=O.radical)亲缘关系最近,从分子角度阐释了野豌豆油壶菌与其它油壶菌的分类关系。采集不同地区蚕豆火肿病样本,扩增病菌的ITS序列并进行多态性分析,它们的同源性高达99%以上,说明野豌豆油壶菌ITS序列的保守性高。针对野豌豆油壶菌ITS序列设计一对特征性引物P1/P2,通过优化扩增条件,能够从病斑组织总DNA中特异性扩增病菌的ITS序列,灵敏度达10pg rDNA,对于火肿病的早期检测和病原菌鉴定有实际应用价值。
Broad bean blister is caused by Olpidium viciae Kusano, characterizing by strumae of cell proliferation in lesion spots. Broad bean blister has only been found in Tokyo of Japan, Sichuan and some areas of its neighboring provinces in China. It is the most important disease occurred in broad bean in Northwestern Plateau of Sichuan province, especially in Aba Autonomous Prefecture and its neighboring areas. Till now, only three research papers were found in deep study of broad bean blister. The life-history and cytology of O. viciae, and the host-parasite relationship in Olpidium were studied by Shunsuke Kusano in1912. Broad bean blister was happened in Songpan county, Sichuan province what was reported first in China by Xin Zhesheng in1984. There were not reported on other aspects except studies of the broad bean blister mentioned above. Comprehensive studies of broad bean blister were carried out, some significant progresses and innovations were made especially on pathogen, histopathology, histophysiology and molecular biology.
1. Symptoms of Broad Bean Blister Disease
Broad bean blister mainly appeared on the leaf and stalk, sometimes also on petiole and carpopodium, but not pod. The early appearance of its symptom was light green nearly round spots, the surfaces of the spots appeared rough with development, proliferated gradually to form intumescent and strumae finally. The strumae was solitary or colonial, was able to fuse into irregular shape. The severe infected plants were pygmyism and leaves were dysmorphosis. Strumae and slightly rough surface were considered as characterization symptom for the broad bean blister, which are the important basis for the diagnosis. This study identified the characterization symptom for diagnosis of broad bean blister, clarified that the blister on bean pod was not the characteristic symptom, which provided scientific basis to diagnose, prevent and control this disease accurately.
2. Etiology of Broad Bean Blister Disease
The pathogen of broad bean blister inhabited in epidermal cells of broad bean leaf, stalk and carpopodium. The fungus body was unicellular and round protoplast, holocarpic reproduction. The vegetative propagation state of O. viciae was zoosporangium, usually sphere, variation in size,12.95～62.16μm in diameter, with an average value of28.84μm. Zoospores were germinated from mature zoosporangium and liberated through spore tubes (holes). Zoospore was oval or spherical,4.1～5μm in size, with an average value of4.47μm. Each zoospore was constituted of a long posterior cilium,24～31μm long with an average value of26.56μm. Zoospore was able to function as motile isogamete and fused together to form zygote with two cilia. Resting sporangium were found in epidermal cells after zygote infected broad bean. Resting sporangium was spherical with thick wall (average value of3.81μm), has approximately uniform size with19～31μm (average value of25.51μm). Resting sporangium was mainly formed at the later period of infection and its number was able to reach more than60in a host cell. According to the morphology, the isolated microorganism was identified as O. viciae which is the disease agent of broad bean blister occurred in Sichuan province. It belongs to the genus Olpidium of the family Olpidiaceae, Chytridiales, Mastigomycotina in taxonomy.
Biological studies on O. viciae indicated the mature zoosporangium in epidermal cells of disease spots was able to liberate zoospores once in the presence of rain water. The temperature range of germination of zoosporangium was from0to18℃, it will be significantly inhibited below18℃, and was unable to germinate at20℃,25℃and30℃. Light or dark did not result in distinct influence on germination of zoosporangium. K+and Na+inhibited the germination to some degree, and completely inhibited the germination when their concentration reached80or100mmol/L. pH also affected the germination of zoosporangium to some degree, and it was germinated at pH values from5to8, with optimum value between7and7.5. The swarming period of zoospores were depended largely upon the temperature, gradually shortened with temperature increase. The zoospores were able to swim for72h between0and5℃; for24,10, and3h below10,20and25℃, respectively; for only5min when temperature raised to30℃. Temperature apparently affected infection and morbidity of zoospores to plants. Infection and morbidity occurred between10and25℃, no apparent symptom observed at5℃although infection occurred, and both were not observed at30℃or above it. The optimum temperature for infection and morbidity was between10and25℃. The time of keeping high humidity for infection and morbidity of zoospores was12h or longer. Leaf age affected infection and morbidity of zoospores, and young leaves were favorable.
In the presence of water, zoospores were liberated from resting sporangium that lived through the summer and the winter. The temperature range of germination was between5and30℃, and zoospores were liberated in4h at17℃. The speed of germination in resting sporangia was extremely different. However the release mode of zoospores was the same with zoosporangium.
3. Occurences and Dynamic of Broad Bean Blister Disease
Outbreak and distribution of broad bean blister were limited in ecological zone above2400m altitude suitable for spring broad bean growth, Aba Tibetan Autonomous Prefecture, Northwestern Plateau of Sichuan province. Life cycle and clinical characteristics of this disease were as follows:The resting sporangia of O. viciae were lived with sick plant residue through summer and winter together, which was the agent for early infection. Zoospores liberated from resting sporangium would infect seedlings of broad bean in next spring, and then the symptoms of disease were found early in May. Zoosporangiums were constantly reproduced after the disease occurred in field, liberated zoospores with the presence of rain or dew for secondary infection. With highly repeated secondary infection, the disease spread quickly in the field and reached peak outbreak in flowering and pod formation stages, after that formed resting sporangium in following pod stage and then stopped gradually. This disease was able to spread in short distance by wind and rain. Continuous cropping and application of animal manure in which sick plant residue existed could cause this disease seriously. No disease-resistant breed is available in all the44breeds used for planting and disease-resistant identification. Pathogen in the sick plant residue from the same year did not infect autumn broad bean in basin zone according to preliminary epidemiological analysis.
4. Histopathology of Broad Bean Blister Disease
To understand the histopathology of broad bean blister caused by O. viciae, the anatomy of epidermal tissues and cross sections of leaf and stem of broad bean, and the growth of the fungus in host cells were observed with light microscope. The results showed that the pathogen infected and parasitized in the epidermal cells of leaf and stem. The whole thallus of the pathogen was converted into a zoosporangium or a resting sporangium. The host responded quickly to the invasion. Once the fungal body in the individual infected cells began to develop, the epidermal cells adjacent infected cell and differentiated cells (including mesophyll cells of leaf and cortex cells of stem) beneath the immediate area of invasion of the pathogen were stimulated to meristematic activity and converted into meristematic cells. The cells of normal basic tissues were replaced by the cells of largely undifferentiated parenchyma with capacity of division. These cells continued to dividing by periclinal or radial division resulting in hyperplasia at local infected site. This histopathological change of hyperplasia was the mechanism of inducing symptom of broad bean blister. The result also indicated the interaction of the pathogen and broad bean at cytological and histological level.
5. Pathological physiology of Broad Bean Blister Disease
Broad bean blister was characterized by strumae as major symptom in infected spots, caused brown stain and necrosis of infected cells. The abnormal pathological change indicated phytohormones changed in host tissues, the activity of antioxidant enzymes in disease tissues changed evidenced by brown stain and necrosis of infected tissues, and then hurt cell to death. In order to deep understand the histophysiology of broad bean blister, concentrations of phytohormones, activity of antioxidant enzyme and concentrations of malonaldehyde in diseased tissues were tested in this study. Results showed that concentrations of ZT and KT increased greatly, together with slight increase in IAA and GA3in early stage. They stimulated cells that were divided and proliferated in diseased tissue, which beginning make the spot verrucose in the early stage. At the same time, increased ABA hurt seedling of broad bean plants, caused them to dwarfing and dysmorphia, delayed their growth stage; damaged old plants and made their leaf pro-senescence and easy to abscission as results. Concentrations of GA3increased dramatically in the late-mid-period of infection, higher than that in health tissue by3times, slight increases were observed for both ZT and KT, as a result, cells what were divided and proliferated in early stage were swelled quickly, which making spot verrucose seriously. ABA increased to some degree in these stages, caused infected cells chlorisis and brown stain, Ultimately Death and leaf senescence. However, concentrations of IAA decreased in the late-mid-period, which was reflected the physiological response of suscept. In the early stage, the analysis of antioxidant enzymes activity in diseased tissues indicated that there are no significant differences of both SOD and POD in diseased tissue compared with health tissue as control; however both of them significantly increased in diseased tissues in late-mid-stages. Because active oxygen removing system was disrupted and active oxygen accumulated in the cells of spot. So, the increased activity of SOD and POD was the biochemical characteristic by infection. Activity of CAT in diseased tissues what in early or in late-mid-stage was lower than that in health tissues. It is well known CAT has high affinity to H2O2, so the decreased CAT led to accumulation of H2O2and then damaged the infected tissues deeply. The concentrations of MDA was higher significantly in diseased tissues in late-mid-stage indicated damages to cellular membrane were intensified, and the inreversible damages disrupted cell and ulcerated tissues finally. These studies uncovered the histophysiological mechanism of symptom caused by broad bean blister, and also discovered the change characteristics of major antioxidant enzymes in diseased tissues in the whole process of broad bean blister.
6. Molecular biology of O. viciae
The Internal Transcribed Spacer (ITS) was amplified with universal primers ITS1/ITS4using total DNA isolated from infected tissues, further identified by sequencing and deposited at NCBI GenBank under accession number HQ677595.1, which added new information of molecular biology for O. viciae. It was most closely related to Olpidium brassicae and Olpidium bornovanus=O radical, elucidated that taxonomic relationships of O. viciae and other Olpidiums at molecular biology level. The ITS sequences were amplified, compared and analyzed for polymorphism using broad bean blister samples collected from different regions, and the homology was reached above99%, which indicates the ITS sequence of O. viciae is highly conservative. Using designed primers of P1/P2, ITS of O. viciae was amplified specifically from diseased tissues with sensitivity of10pg rDNA after optimizing the amplification conditions, and it had practical value for application in early detection and pathogenic identification of broad bean blister when the characteristic symptom did not appear.
1.蚕豆火肿病的症状
蚕豆火肿病主要危害蚕豆叶和茎,也危害叶柄和果柄,但不危害果荚。病部最初出现稍褪绿的近圆形斑痕,随病情发展病斑表面稍显粗糙,并不断向上增生隆起,形成小的瘤状突起。病瘤单生或群生,可相互愈合成不规则状。受害严重时,病叶畸形,植株矮小。病部形成小的瘤状突起、表面稍显粗糙,是本病的标志性特征,是诊断本病的重要依据。本研究完善了蚕豆油壶菌火肿病的症状特征,明确了蚕豆豆荚上的疱状突起不是本病的症状,从而为准确诊断和防冶蚕豆油壶菌火肿病提供了科学依据。
2.蚕豆火肿病的病原
蚕豆火肿病病原菌寄生在蚕豆叶、茎和果柄表皮细胞内。营养体为单细胞圆形原生质团,营养体整体产果。无性繁殖结构为游动孢子囊,一般为球形,大小变化较大,一般为12.95～62.16μm,平均28.84μm。游动孢子囊萌发产生游动孢子,通过排孢管(孔)释放。游动孢子卵圆形或球形,大小为4.1-5μm,平均为4.47μm。尾部具一根鞭毛,鞭毛长24-3lμm,平均为26.56μm。游动孢子可作为同形游动配子结合,形成双鞭毛圆形的接合子。接合子侵染蚕豆,在表皮细胞内形成休眠孢子囊。休眠孢子囊圆形厚壁,形状大小较一致,大小为19-31μm,平均为25.51μm,平均壁厚3.81μm。一个寄主细胞中的数目可多达60个以上,休眠孢子囊大多形成在病害后期。对该菌的形态学研究表明：四川蚕豆火肿病是由野豌豆油壶菌(Olpidium viciae Kusano)引起。该菌的分类地位属鞭毛菌亚门、壶菌目、油壶菌科、油壶菌属。
对蚕豆火肿病菌的生物学研究结果表明：病斑表皮组织中的游动孢子囊成熟后,遇雨水即可萌发,释放出游动孢子。游动孢子囊萌发的温度范围为0-18。C,在18℃下萌发就受到明显抑制,在20℃、25℃和30。C下不能萌发。光照或黑暗对游动孢子囊萌发没有明显影响。K、Na盐离子溶液对孢子囊萌发有一定抑制作用,浓度分别达到80mmo1/L、100mmol/L时,游动孢子囊萌发就几乎被抑制。pH对游动孢子囊萌发也有一定影响,萌发的pH范围为pH5-8,以pH7-7.5最适。温度影响游动孢子的游动时间长短,在0-30℃范围内,随温度升高,游动时间逐渐缩短。在0-5℃下保持游动时间最长可达72h以上,10℃下可达24h,15℃和20℃下可保持10h,25℃下为3h,30℃下仅可保持游动5min。温度对游动孢子侵染及发病有明显影响,在5℃下能成功侵染,但不产生明显症状。侵染和引起发病的温度范围为10～25℃,30。C及以上温度不能侵染和发病。最适侵染和发病的温度为10-20℃。接种游动孢子引起侵染和发病的保湿时间为12h以上。叶龄影响病菌的侵染和发病,幼叶有利侵染和发病。
病组织中的休眠孢子囊必须经过越夏和越冬后,有水存在时才能萌发产生游动孢子。萌发的温度范围为5-30℃,在17℃左右温度下,经4h即可产生游动孢子。病组织中的休眠孢子囊萌发极不整齐,休眠孢子囊释放游动孢子的方式与游动孢子囊的释放方式相同。
3.蚕豆火肿病的发生规律
四川蚕豆火肿病的发生与分布局限在四川西北部高原的阿坝藏族自治州海拔2400m以上的春播蚕豆生态区。病害的循环和发病特点是：病菌以休眠孢子囊随病残体在土壤中越夏和越冬,作为初侵染源。次年春播蚕豆后,休眠孢子囊萌发产生游动孢子侵染蚕豆幼苗,病害始发期在5月。田间发病后,病菌不断产生游动孢子囊,在有雨露时游动孢子囊萌发释放游动孢子进行再侵染。在蚕豆生长期中病菌的再侵染频繁,病害在田间扩展迅速,在蚕豆进入开花和始荚期,病害进入高峰期；蚕豆进入结荚期后,病菌形成休眠孢子囊,病害发展趋于停止。病菌的游动孢子藉风雨在田间近距离传播。连作和施用带有病残体的厩肥有利于发病。目前生产上种植和供抗性鉴定的40多个蚕豆材料均不同程度感病,缺乏抗病品种。病害传播风险的初步结果表明：病区当年的病残体菌源对盆地内秋播蚕豆区不构成侵染发病的危险。
4.蚕豆火肿病的组织病理学
野豌豆油壶菌侵入蚕豆叶和茎的表皮细胞后,定殖在侵入的细胞内,菌体生长发育,整体产果,形成游动孢子囊和休眠孢子囊。病菌的侵染刺激了侵染点邻近的表皮细胞和其下的叶肉组织或茎的皮层组织细胞,这些已分化的组织细胞转为分生细胞,进行分裂,分裂方式为平周分裂。原有的正常基本组织细胞被未高度分化的薄壁组织细胞取代。这些细胞具有分生能力并不断分裂,细胞体积增大,造成蚕豆病组织细胞发生畸形增生的病理解剖改变,病部呈瘤状隆起。这是蚕豆油壶菌火肿病症状产生的病理组织学机制,也反映了野豌豆油壶菌和蚕豆在细胞和组织水平上的相互作用。
5.蚕豆火肿病的病理生理
蚕豆火肿病的症状特点主要是受病部位产生瘤状突起,病部细胞逐渐褐变坏死。这种畸形的病理改变预示着受病菌侵染后,寄主组织中的植物激素发生了异常变化；受病组织逐渐褐变坏死,显示抗氧化保护酶活力改变,细胞受到伤害,最终坏死。为深入了解蚕豆火肿病发生病理改变的生理机制,测定了病组织中的植物激素含量、抗氧化保护酶活性和丙二醛含量。结果表明,病害初期组织中的ZT和KT含量大幅上升,IAA和GA。含量也有一定程度增加,它们共同刺激蚕豆病部细胞大量分裂、增生和体积逐渐膨大,病部开始隆起；在病害初期ABA含量也大量增加,表现为蚕豆幼苗被害,植株矮化畸形、生育期严重滞后,而成株被害,受病叶片提前衰老、易脱落。在病害中后期组织中的GA3含量急剧增加,是健康组织的3倍,ZT和KT仍维持一定幅度增量,但增量率明显降低,使得病害早期大量分裂增生的细胞在病害中后期迅速膨大,病部异常隆起；ABA在病害中后期也有较大增量,使病部细胞褪绿褐变,最终坏死,受病叶片脱落；而IAA在病害中后期呈现负增长,这是寄主感病的生理表现。受病组织中抗氧化保护酶活力测定的结果表明：病、健组织中的SOD、POD活力在病害初期没有明显差异,在病害中后期病组织均显著高于健康组织,说明病菌侵染破坏了寄主本身正常活性氧消除机制的平衡,导致体内活性氧积累,诱导抗氧化保护酶SOD和POD活性增强,是感病的生化表现。而CAT活力在病程初期和中后期,病组织均极显著低于健康组织。己知CAT和H202具有较高的亲和力,CAT活力降低导致组织中的H202积累,加重对组织的损伤。组织中的MDA含量在病害中后期极显著高于健康组织,说明病部组织细胞生物膜损伤加剧,这种不可逆的伤害使细胞最终破坏解体、组织溃烂。这些结果反映了蚕豆火肿病症状产生的病理生理机制,也揭示了蚕豆火肿病病程中组织细胞内主要抗氧化保护酶的病理变化特点。
6.野豌豆油壶菌的分子生物学研究
通过提取病斑组织的总DNA,采用真核生物内转录间隔区通用引物ITS1/ITS4进行扩增,获得了病菌的ITS序列；回收此序列测序,GenBank登记号为HQ677595.1,补充了野豌豆油壶菌的分子生物学研究资料。将病菌与其它真菌进行比对,显示与甘蓝油壶菌(Olpidium brassicae)和瓜类油壶菌(Olpidium bornovanus=O.radical)亲缘关系最近,从分子角度阐释了野豌豆油壶菌与其它油壶菌的分类关系。采集不同地区蚕豆火肿病样本,扩增病菌的ITS序列并进行多态性分析,它们的同源性高达99%以上,说明野豌豆油壶菌ITS序列的保守性高。针对野豌豆油壶菌ITS序列设计一对特征性引物P1/P2,通过优化扩增条件,能够从病斑组织总DNA中特异性扩增病菌的ITS序列,灵敏度达10pg rDNA,对于火肿病的早期检测和病原菌鉴定有实际应用价值。
Broad bean blister is caused by Olpidium viciae Kusano, characterizing by strumae of cell proliferation in lesion spots. Broad bean blister has only been found in Tokyo of Japan, Sichuan and some areas of its neighboring provinces in China. It is the most important disease occurred in broad bean in Northwestern Plateau of Sichuan province, especially in Aba Autonomous Prefecture and its neighboring areas. Till now, only three research papers were found in deep study of broad bean blister. The life-history and cytology of O. viciae, and the host-parasite relationship in Olpidium were studied by Shunsuke Kusano in1912. Broad bean blister was happened in Songpan county, Sichuan province what was reported first in China by Xin Zhesheng in1984. There were not reported on other aspects except studies of the broad bean blister mentioned above. Comprehensive studies of broad bean blister were carried out, some significant progresses and innovations were made especially on pathogen, histopathology, histophysiology and molecular biology.
1. Symptoms of Broad Bean Blister Disease
Broad bean blister mainly appeared on the leaf and stalk, sometimes also on petiole and carpopodium, but not pod. The early appearance of its symptom was light green nearly round spots, the surfaces of the spots appeared rough with development, proliferated gradually to form intumescent and strumae finally. The strumae was solitary or colonial, was able to fuse into irregular shape. The severe infected plants were pygmyism and leaves were dysmorphosis. Strumae and slightly rough surface were considered as characterization symptom for the broad bean blister, which are the important basis for the diagnosis. This study identified the characterization symptom for diagnosis of broad bean blister, clarified that the blister on bean pod was not the characteristic symptom, which provided scientific basis to diagnose, prevent and control this disease accurately.
2. Etiology of Broad Bean Blister Disease
The pathogen of broad bean blister inhabited in epidermal cells of broad bean leaf, stalk and carpopodium. The fungus body was unicellular and round protoplast, holocarpic reproduction. The vegetative propagation state of O. viciae was zoosporangium, usually sphere, variation in size,12.95～62.16μm in diameter, with an average value of28.84μm. Zoospores were germinated from mature zoosporangium and liberated through spore tubes (holes). Zoospore was oval or spherical,4.1～5μm in size, with an average value of4.47μm. Each zoospore was constituted of a long posterior cilium,24～31μm long with an average value of26.56μm. Zoospore was able to function as motile isogamete and fused together to form zygote with two cilia. Resting sporangium were found in epidermal cells after zygote infected broad bean. Resting sporangium was spherical with thick wall (average value of3.81μm), has approximately uniform size with19～31μm (average value of25.51μm). Resting sporangium was mainly formed at the later period of infection and its number was able to reach more than60in a host cell. According to the morphology, the isolated microorganism was identified as O. viciae which is the disease agent of broad bean blister occurred in Sichuan province. It belongs to the genus Olpidium of the family Olpidiaceae, Chytridiales, Mastigomycotina in taxonomy.
Biological studies on O. viciae indicated the mature zoosporangium in epidermal cells of disease spots was able to liberate zoospores once in the presence of rain water. The temperature range of germination of zoosporangium was from0to18℃, it will be significantly inhibited below18℃, and was unable to germinate at20℃,25℃and30℃. Light or dark did not result in distinct influence on germination of zoosporangium. K+and Na+inhibited the germination to some degree, and completely inhibited the germination when their concentration reached80or100mmol/L. pH also affected the germination of zoosporangium to some degree, and it was germinated at pH values from5to8, with optimum value between7and7.5. The swarming period of zoospores were depended largely upon the temperature, gradually shortened with temperature increase. The zoospores were able to swim for72h between0and5℃; for24,10, and3h below10,20and25℃, respectively; for only5min when temperature raised to30℃. Temperature apparently affected infection and morbidity of zoospores to plants. Infection and morbidity occurred between10and25℃, no apparent symptom observed at5℃although infection occurred, and both were not observed at30℃or above it. The optimum temperature for infection and morbidity was between10and25℃. The time of keeping high humidity for infection and morbidity of zoospores was12h or longer. Leaf age affected infection and morbidity of zoospores, and young leaves were favorable.
In the presence of water, zoospores were liberated from resting sporangium that lived through the summer and the winter. The temperature range of germination was between5and30℃, and zoospores were liberated in4h at17℃. The speed of germination in resting sporangia was extremely different. However the release mode of zoospores was the same with zoosporangium.
3. Occurences and Dynamic of Broad Bean Blister Disease
Outbreak and distribution of broad bean blister were limited in ecological zone above2400m altitude suitable for spring broad bean growth, Aba Tibetan Autonomous Prefecture, Northwestern Plateau of Sichuan province. Life cycle and clinical characteristics of this disease were as follows:The resting sporangia of O. viciae were lived with sick plant residue through summer and winter together, which was the agent for early infection. Zoospores liberated from resting sporangium would infect seedlings of broad bean in next spring, and then the symptoms of disease were found early in May. Zoosporangiums were constantly reproduced after the disease occurred in field, liberated zoospores with the presence of rain or dew for secondary infection. With highly repeated secondary infection, the disease spread quickly in the field and reached peak outbreak in flowering and pod formation stages, after that formed resting sporangium in following pod stage and then stopped gradually. This disease was able to spread in short distance by wind and rain. Continuous cropping and application of animal manure in which sick plant residue existed could cause this disease seriously. No disease-resistant breed is available in all the44breeds used for planting and disease-resistant identification. Pathogen in the sick plant residue from the same year did not infect autumn broad bean in basin zone according to preliminary epidemiological analysis.
4. Histopathology of Broad Bean Blister Disease
To understand the histopathology of broad bean blister caused by O. viciae, the anatomy of epidermal tissues and cross sections of leaf and stem of broad bean, and the growth of the fungus in host cells were observed with light microscope. The results showed that the pathogen infected and parasitized in the epidermal cells of leaf and stem. The whole thallus of the pathogen was converted into a zoosporangium or a resting sporangium. The host responded quickly to the invasion. Once the fungal body in the individual infected cells began to develop, the epidermal cells adjacent infected cell and differentiated cells (including mesophyll cells of leaf and cortex cells of stem) beneath the immediate area of invasion of the pathogen were stimulated to meristematic activity and converted into meristematic cells. The cells of normal basic tissues were replaced by the cells of largely undifferentiated parenchyma with capacity of division. These cells continued to dividing by periclinal or radial division resulting in hyperplasia at local infected site. This histopathological change of hyperplasia was the mechanism of inducing symptom of broad bean blister. The result also indicated the interaction of the pathogen and broad bean at cytological and histological level.
5. Pathological physiology of Broad Bean Blister Disease
Broad bean blister was characterized by strumae as major symptom in infected spots, caused brown stain and necrosis of infected cells. The abnormal pathological change indicated phytohormones changed in host tissues, the activity of antioxidant enzymes in disease tissues changed evidenced by brown stain and necrosis of infected tissues, and then hurt cell to death. In order to deep understand the histophysiology of broad bean blister, concentrations of phytohormones, activity of antioxidant enzyme and concentrations of malonaldehyde in diseased tissues were tested in this study. Results showed that concentrations of ZT and KT increased greatly, together with slight increase in IAA and GA3in early stage. They stimulated cells that were divided and proliferated in diseased tissue, which beginning make the spot verrucose in the early stage. At the same time, increased ABA hurt seedling of broad bean plants, caused them to dwarfing and dysmorphia, delayed their growth stage; damaged old plants and made their leaf pro-senescence and easy to abscission as results. Concentrations of GA3increased dramatically in the late-mid-period of infection, higher than that in health tissue by3times, slight increases were observed for both ZT and KT, as a result, cells what were divided and proliferated in early stage were swelled quickly, which making spot verrucose seriously. ABA increased to some degree in these stages, caused infected cells chlorisis and brown stain, Ultimately Death and leaf senescence. However, concentrations of IAA decreased in the late-mid-period, which was reflected the physiological response of suscept. In the early stage, the analysis of antioxidant enzymes activity in diseased tissues indicated that there are no significant differences of both SOD and POD in diseased tissue compared with health tissue as control; however both of them significantly increased in diseased tissues in late-mid-stages. Because active oxygen removing system was disrupted and active oxygen accumulated in the cells of spot. So, the increased activity of SOD and POD was the biochemical characteristic by infection. Activity of CAT in diseased tissues what in early or in late-mid-stage was lower than that in health tissues. It is well known CAT has high affinity to H2O2, so the decreased CAT led to accumulation of H2O2and then damaged the infected tissues deeply. The concentrations of MDA was higher significantly in diseased tissues in late-mid-stage indicated damages to cellular membrane were intensified, and the inreversible damages disrupted cell and ulcerated tissues finally. These studies uncovered the histophysiological mechanism of symptom caused by broad bean blister, and also discovered the change characteristics of major antioxidant enzymes in diseased tissues in the whole process of broad bean blister.
6. Molecular biology of O. viciae
The Internal Transcribed Spacer (ITS) was amplified with universal primers ITS1/ITS4using total DNA isolated from infected tissues, further identified by sequencing and deposited at NCBI GenBank under accession number HQ677595.1, which added new information of molecular biology for O. viciae. It was most closely related to Olpidium brassicae and Olpidium bornovanus=O radical, elucidated that taxonomic relationships of O. viciae and other Olpidiums at molecular biology level. The ITS sequences were amplified, compared and analyzed for polymorphism using broad bean blister samples collected from different regions, and the homology was reached above99%, which indicates the ITS sequence of O. viciae is highly conservative. Using designed primers of P1/P2, ITS of O. viciae was amplified specifically from diseased tissues with sensitivity of10pg rDNA after optimizing the amplification conditions, and it had practical value for application in early detection and pathogenic identification of broad bean blister when the characteristic symptom did not appear.
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