紫花苜蓿种带细菌及其致病性
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摘要
紫花苜蓿(Medicago sativa)是我国最主要的豆科栽培牧草,健康的种子是优质高产的保证。本研究以美国、加拿大和中国甘肃等收获的53个紫花苜蓿种样为材料,研究了其种带细菌的多样性及其作用；明确紫花苜蓿芽期细菌性病害种类和病原；探讨了这些致病细菌的环境适应力,测定了病原菌的寄主范围以及侵染循环途径,旨在为紫花苜蓿种传病害的防治提供基础资料。所获主要结果如下：
1、紫花苜蓿种带细菌多样性。从供试种样中分离得到382株细菌分离物,其中19种细菌分离物为优势种群。结合常规表型特征、Biolog和16S rDNA鉴定方法,确定了该19种优势菌株分属于9个属18种细菌,其中革兰氏阴性菌5属10种,革兰氏阳性菌4属8种。其中11种为国际首次报道,分别为东方假单胞菌(Pseudomonas oriental is)2Ap1、简单芽孢杆菌(Bacillus simplex) ARI、萎蔫芽孢杆菌(Bacillus atrophaeus) BF、桃色欧文氏菌(Erwinia persicind)1Cp2、多粘类芽孢杆菌(Paenibacillus polymyxd)3Cp1、玫瑰色考克氏菌(Kocuria rosed)1fF6、中间苍白杆菌(Ochrobactrum intermedium) Gp1、短小芽孢杆菌(Bacillus pumilus)1gF2、成晶节杆菌(Arthrobacter crystallopoietes)HT7、根际苍白杆菌(Ochrobactrum rhizosphaerae) HT13和阴沟肠杆菌(Enterobacter cloacae) YZ-DS1。18种为国内首次报道,除以上11种外,另有7种分别是成团泛菌(Pantoea agglomerans)1Ap1、成团肠杆菌(Enterobacter agglomerans)1Bp1、荧光假单胞菌生物型I(Pseudomonas fluorescens Biovar Ⅰ)3Bp3、蜡状芽孢杆菌(Bacillus anthracis) Fp、阪崎肠杆菌(Enterobacter sakazakii)1gF3,荧光假单胞菌生物型Ⅳ(Pseudomonas fluorescens Biovar Ⅳ)2Hp1和枯草芽孢杆菌(Bacillus subtilis)HT17。
2、紫花苜蓿种带细菌的致病性。采用浸种接种法,以阿尔冈金品种的紫花苜蓿为材料,分别接种上述19种优势菌种。发现桃色欧文氏菌1Cp2、成团泛菌1Apl、成团肠杆菌1Bpl、阴沟肠杆菌YZ-DS1、中间苍白杆菌Gp1和根际苍白杆菌HT13等6个菌种有致病作用。与未接种的对照相比,发芽率降低4.61%-28.5%,种子萎蔫和腐烂率为25%-50%等。进一步的研究发现桃色欧文氏菌、成团泛菌和阴沟肠杆菌等三种病原分别引致苜蓿芽细菌性萎蔫病、苜蓿细菌性种子坏死病和苜蓿芽细菌性腐烂病。研究还发现多粘类芽孢杆菌3Cp1和枯草芽孢杆菌HT17对苜蓿生长有促进。
3、桃色欧文氏菌的生物学特性。研究发现,该菌的的生长温度为5～40℃,最适生长温度为28℃；在培养基中PEG6000含量0g/l～500g/l的范围内,其生长量随PEG6000的添加量而降低,但能耐受PEG6000含量400g/l的较低水势；桃色欧文氏菌在12h光暗交替条件下生长量最高(OD600=1.753),全日照条件下生长量次之(OD600=1.543),而全黑暗条件下生长最差(OD600=1.223).该菌耐受pH范围为4～10,pH7.0时生长量最高。0.5%NaCl有利其的生长,最高能耐受3%NaCl含量,不能在4%NaCl含量下生长。麦芽糖、葡萄糖、蔗糖、甘油和果糖等5种碳源可作为选择性培养基的碳源。大豆蛋白胨为的选择性培养基的氮源。而醋酸铵和磺胺分别是不利碳源和氮源。
4、桃色欧文氏菌的传播。本研究国际首次证实了在盆栽条件下桃色欧文氏菌能引起成株紫花苜蓿致病,发病率为100%。其可能通过水源、土壤和种子传播。通过悬浮液浸种或浇芽及喷雾接种,证实其可以通过自然孔口入侵紫花苜蓿,引起茎干、叶片、花朵和豆荚坏死以及种子呈现水渍状病变。一旦病原菌进入植株体内,其可运动到植株的根际、茎干、叶围、花朵最后至种子上得以保存。待条件合适将再次随播种种子侵染。
5、桃色欧文氏菌的寄主范围。人工接种条件下,该菌可侵染11种豆类作物和12种豆科牧草,包括紫花苜蓿、豌豆(Pisum sativum)、菜豆(Phaseolus vulgari)、大豆(Glycine max)、红豆(Vigna angularis),豇豆(V. unguiculata)、绿豆(V. radiate)、蚕豆(Vicia faba)、扁豆(Dolichos lablab)、小扁豆(Lens culinaris)、花生(Arachis hypogaea)、棉豆(Phaseolus lunatus)、红豆草(Onobrychis viciaefolia)、沙打旺(Astragalus adsurgens)、白三叶(Trifolium repens)、红三叶(T.pretense)、箭筈豌豆(Vicia sativa)、野豌豆(V. sepiuni)、歪头菜(V. unijuga)、胡枝子(Lespedeza bicolor)、百脉根(Lotus cornioulatus)、白花草木樨(Melilotus albus)和黄花草木樨(M.officinalis)。除前四种外,其余均为国际首次报道。
Lucerne(Medicago sativa) is the main cultivated forage legume in China and high-quality seeds guarantee its high yield. In this study,53seed samples from the United States, Canada, and Gansu province of China were collected as research materials. The objectives of this dissertation were (i) to understand the diversity of lucerne seed-borne bacteria and their functions,(ii) to investigate the patterns and pathogen of bacterial diseases of lucerne, and (iii) to probe the environmental adaptability of pathogenic microbes and to identify some specific pathogen and their hosts, as well as the infection way of the pathogen, so that we can provide basic information and theoretical evidence for controlling bacterial diseases of lucerne.
1. Diversity of lucerne seed-borne bacteria. From the tested lucerne seeds,382bacterial strains were isolated and among which19strains were the predominant bacterial populations. These19bacterial strains were identified to be9genera and18species using the methods of phenotype characteristics, Biolog and16S rDNA In these trains,10species of five genera were Gram negative bacteria, and eight species of four genera were Gram positive bacteria. Eleven species of bacteria were reported worldwide for the first time and they were Pseudomonas orientalis2Ap1, Bacillus simplex ARI, Bacillus atrophaeus BF, Erwinia persicina1Cp2, Paenibacillus polymyxa3Cp1, Kocuria rosea fF6, Ochrobactrum intermedium Gp1, Bacillus pumilus1gF2, Arthrobacter crystallopoietes HT7, Ochrobactrum rhizosphaerae HT13, and Enterobacter cloacae YZ-DS1. Seven species of bacteria first reported in China were Pantoea agglomerans1Ap1, Enterobacter agglomerans1Bp1, Pseudomonas fluorescens Biovar Ⅰ3Bp3, Bacillus cereus Fp, Enterobacter sakazakii1gF3, Pseudomonas fluorescens Biovar Ⅳ2Hp1, and Bacillus subtilis HT17.
2. Pathogenicity of lucerne seed-borne bacteria. Algonquin lucerne seeds were soaked in a dominant bacterial suspension and several varieties such as E. persicina1Cp2, P. agglomerans1Ap1, E. agglomerans1Bp1, E. cloacae YZ-DS1, O. intermedium GPl and O. rhizosphaerae HT13were found to be pathogenic to lucerne by hindering the plant growth and affecting the yield. In fact, only three bacterial diseases of lucerne were found in previous germination trials and they were bacterial sprout wilt caused by E. persicina, bacterial sprout decay caused by E. cloacae, and bacterial seed necrosis caused by P. agglomerans. In the study, two beneficial bacteria, P. polymyxa3Cp1and B. subtil is HT17, were identified.
3. Biological characteristics of Erwinia persicina. The growth temperature for E. persicina was5～40℃and the optimal temperature was28℃. The growth rate of E. persicina was reduced as water content decreased in the culture media, but E. persicina could tolerate low water content. The best light treatment condition for E. persicina growth was the12h light/dark photoperiod. E. persicina showed the lowest growth rate when incubated in complete darkness and a medium growth rate under full light conditions. Although E. persicina could tolerate pH values between4and10, the highest growth rate was observed at pH7. Salt concentration at0.5%NaCl was beneficial to the growth of E. persicina. The bacteria could tolerate3%NaCl; however, they could not grow in4%NaCl. The results showed that maltose, glucose, sucrose, glycerol and fucose could be used as carbon sources, soy peptone as the nitrogen source, and ammonium acetate and sulfanilamide as the adverse carbon and nitrogen source, respectively, in selective media for E. persicina
4. Transmission of Erwinia persicina. It was verified that E. persicina could infect mature lucerne and the incidence rate was100%. The transmission might be achieved through water source, soil and seeds. Using the above inoculation method, we found that E. persicina could invade the tested lucerne through natural openings, which could result in the necrosis of stems, leaves, flowers and bean pods as well as the water-soaked lesion of seeds. Once E. persicina enter lucerne, they could move with the help of flagella to rhizosphere, stems, leaves, flowers and subsequently to seeds for preservation. Once the environment is suitable for the growth of E. persicina, they would infect new plants.
5. Hosts of Erwinia persicina. It was proved that E. persicina could infect23species in15genera of legumes and all organs in each plant including stems, leaves, flowers, bean pods and seeds. Among11leguminous crops and12leguminous forages used in this experiment, it was known that lucerne, pea, kidney bean and soybean could be infected by E. persicina, while the other species were newly identified hosts, which include grain legumes such as Vigna angularis, V. unguiculata, V. radiate, Vicia faba, Dolichos lablab, Lens culinaris, Arachis hypogaea, and Phaseolus lunatus, as well as forage legumes such as Onobrychis viciaefolia, Astragalus adsurgens, Trifolium repens, T. pretense, Vicia sativa, V. sepium, V. unijuga, Lespedeza bicolor, Lotus cornioulatus, Melilotus albus, and M. officinalis.
1、紫花苜蓿种带细菌多样性。从供试种样中分离得到382株细菌分离物,其中19种细菌分离物为优势种群。结合常规表型特征、Biolog和16S rDNA鉴定方法,确定了该19种优势菌株分属于9个属18种细菌,其中革兰氏阴性菌5属10种,革兰氏阳性菌4属8种。其中11种为国际首次报道,分别为东方假单胞菌(Pseudomonas oriental is)2Ap1、简单芽孢杆菌(Bacillus simplex) ARI、萎蔫芽孢杆菌(Bacillus atrophaeus) BF、桃色欧文氏菌(Erwinia persicind)1Cp2、多粘类芽孢杆菌(Paenibacillus polymyxd)3Cp1、玫瑰色考克氏菌(Kocuria rosed)1fF6、中间苍白杆菌(Ochrobactrum intermedium) Gp1、短小芽孢杆菌(Bacillus pumilus)1gF2、成晶节杆菌(Arthrobacter crystallopoietes)HT7、根际苍白杆菌(Ochrobactrum rhizosphaerae) HT13和阴沟肠杆菌(Enterobacter cloacae) YZ-DS1。18种为国内首次报道,除以上11种外,另有7种分别是成团泛菌(Pantoea agglomerans)1Ap1、成团肠杆菌(Enterobacter agglomerans)1Bp1、荧光假单胞菌生物型I(Pseudomonas fluorescens Biovar Ⅰ)3Bp3、蜡状芽孢杆菌(Bacillus anthracis) Fp、阪崎肠杆菌(Enterobacter sakazakii)1gF3,荧光假单胞菌生物型Ⅳ(Pseudomonas fluorescens Biovar Ⅳ)2Hp1和枯草芽孢杆菌(Bacillus subtilis)HT17。
2、紫花苜蓿种带细菌的致病性。采用浸种接种法,以阿尔冈金品种的紫花苜蓿为材料,分别接种上述19种优势菌种。发现桃色欧文氏菌1Cp2、成团泛菌1Apl、成团肠杆菌1Bpl、阴沟肠杆菌YZ-DS1、中间苍白杆菌Gp1和根际苍白杆菌HT13等6个菌种有致病作用。与未接种的对照相比,发芽率降低4.61%-28.5%,种子萎蔫和腐烂率为25%-50%等。进一步的研究发现桃色欧文氏菌、成团泛菌和阴沟肠杆菌等三种病原分别引致苜蓿芽细菌性萎蔫病、苜蓿细菌性种子坏死病和苜蓿芽细菌性腐烂病。研究还发现多粘类芽孢杆菌3Cp1和枯草芽孢杆菌HT17对苜蓿生长有促进。
3、桃色欧文氏菌的生物学特性。研究发现,该菌的的生长温度为5～40℃,最适生长温度为28℃；在培养基中PEG6000含量0g/l～500g/l的范围内,其生长量随PEG6000的添加量而降低,但能耐受PEG6000含量400g/l的较低水势；桃色欧文氏菌在12h光暗交替条件下生长量最高(OD600=1.753),全日照条件下生长量次之(OD600=1.543),而全黑暗条件下生长最差(OD600=1.223).该菌耐受pH范围为4～10,pH7.0时生长量最高。0.5%NaCl有利其的生长,最高能耐受3%NaCl含量,不能在4%NaCl含量下生长。麦芽糖、葡萄糖、蔗糖、甘油和果糖等5种碳源可作为选择性培养基的碳源。大豆蛋白胨为的选择性培养基的氮源。而醋酸铵和磺胺分别是不利碳源和氮源。
4、桃色欧文氏菌的传播。本研究国际首次证实了在盆栽条件下桃色欧文氏菌能引起成株紫花苜蓿致病,发病率为100%。其可能通过水源、土壤和种子传播。通过悬浮液浸种或浇芽及喷雾接种,证实其可以通过自然孔口入侵紫花苜蓿,引起茎干、叶片、花朵和豆荚坏死以及种子呈现水渍状病变。一旦病原菌进入植株体内,其可运动到植株的根际、茎干、叶围、花朵最后至种子上得以保存。待条件合适将再次随播种种子侵染。
5、桃色欧文氏菌的寄主范围。人工接种条件下,该菌可侵染11种豆类作物和12种豆科牧草,包括紫花苜蓿、豌豆(Pisum sativum)、菜豆(Phaseolus vulgari)、大豆(Glycine max)、红豆(Vigna angularis),豇豆(V. unguiculata)、绿豆(V. radiate)、蚕豆(Vicia faba)、扁豆(Dolichos lablab)、小扁豆(Lens culinaris)、花生(Arachis hypogaea)、棉豆(Phaseolus lunatus)、红豆草(Onobrychis viciaefolia)、沙打旺(Astragalus adsurgens)、白三叶(Trifolium repens)、红三叶(T.pretense)、箭筈豌豆(Vicia sativa)、野豌豆(V. sepiuni)、歪头菜(V. unijuga)、胡枝子(Lespedeza bicolor)、百脉根(Lotus cornioulatus)、白花草木樨(Melilotus albus)和黄花草木樨(M.officinalis)。除前四种外,其余均为国际首次报道。
Lucerne(Medicago sativa) is the main cultivated forage legume in China and high-quality seeds guarantee its high yield. In this study,53seed samples from the United States, Canada, and Gansu province of China were collected as research materials. The objectives of this dissertation were (i) to understand the diversity of lucerne seed-borne bacteria and their functions,(ii) to investigate the patterns and pathogen of bacterial diseases of lucerne, and (iii) to probe the environmental adaptability of pathogenic microbes and to identify some specific pathogen and their hosts, as well as the infection way of the pathogen, so that we can provide basic information and theoretical evidence for controlling bacterial diseases of lucerne.
1. Diversity of lucerne seed-borne bacteria. From the tested lucerne seeds,382bacterial strains were isolated and among which19strains were the predominant bacterial populations. These19bacterial strains were identified to be9genera and18species using the methods of phenotype characteristics, Biolog and16S rDNA In these trains,10species of five genera were Gram negative bacteria, and eight species of four genera were Gram positive bacteria. Eleven species of bacteria were reported worldwide for the first time and they were Pseudomonas orientalis2Ap1, Bacillus simplex ARI, Bacillus atrophaeus BF, Erwinia persicina1Cp2, Paenibacillus polymyxa3Cp1, Kocuria rosea fF6, Ochrobactrum intermedium Gp1, Bacillus pumilus1gF2, Arthrobacter crystallopoietes HT7, Ochrobactrum rhizosphaerae HT13, and Enterobacter cloacae YZ-DS1. Seven species of bacteria first reported in China were Pantoea agglomerans1Ap1, Enterobacter agglomerans1Bp1, Pseudomonas fluorescens Biovar Ⅰ3Bp3, Bacillus cereus Fp, Enterobacter sakazakii1gF3, Pseudomonas fluorescens Biovar Ⅳ2Hp1, and Bacillus subtilis HT17.
2. Pathogenicity of lucerne seed-borne bacteria. Algonquin lucerne seeds were soaked in a dominant bacterial suspension and several varieties such as E. persicina1Cp2, P. agglomerans1Ap1, E. agglomerans1Bp1, E. cloacae YZ-DS1, O. intermedium GPl and O. rhizosphaerae HT13were found to be pathogenic to lucerne by hindering the plant growth and affecting the yield. In fact, only three bacterial diseases of lucerne were found in previous germination trials and they were bacterial sprout wilt caused by E. persicina, bacterial sprout decay caused by E. cloacae, and bacterial seed necrosis caused by P. agglomerans. In the study, two beneficial bacteria, P. polymyxa3Cp1and B. subtil is HT17, were identified.
3. Biological characteristics of Erwinia persicina. The growth temperature for E. persicina was5～40℃and the optimal temperature was28℃. The growth rate of E. persicina was reduced as water content decreased in the culture media, but E. persicina could tolerate low water content. The best light treatment condition for E. persicina growth was the12h light/dark photoperiod. E. persicina showed the lowest growth rate when incubated in complete darkness and a medium growth rate under full light conditions. Although E. persicina could tolerate pH values between4and10, the highest growth rate was observed at pH7. Salt concentration at0.5%NaCl was beneficial to the growth of E. persicina. The bacteria could tolerate3%NaCl; however, they could not grow in4%NaCl. The results showed that maltose, glucose, sucrose, glycerol and fucose could be used as carbon sources, soy peptone as the nitrogen source, and ammonium acetate and sulfanilamide as the adverse carbon and nitrogen source, respectively, in selective media for E. persicina
4. Transmission of Erwinia persicina. It was verified that E. persicina could infect mature lucerne and the incidence rate was100%. The transmission might be achieved through water source, soil and seeds. Using the above inoculation method, we found that E. persicina could invade the tested lucerne through natural openings, which could result in the necrosis of stems, leaves, flowers and bean pods as well as the water-soaked lesion of seeds. Once E. persicina enter lucerne, they could move with the help of flagella to rhizosphere, stems, leaves, flowers and subsequently to seeds for preservation. Once the environment is suitable for the growth of E. persicina, they would infect new plants.
5. Hosts of Erwinia persicina. It was proved that E. persicina could infect23species in15genera of legumes and all organs in each plant including stems, leaves, flowers, bean pods and seeds. Among11leguminous crops and12leguminous forages used in this experiment, it was known that lucerne, pea, kidney bean and soybean could be infected by E. persicina, while the other species were newly identified hosts, which include grain legumes such as Vigna angularis, V. unguiculata, V. radiate, Vicia faba, Dolichos lablab, Lens culinaris, Arachis hypogaea, and Phaseolus lunatus, as well as forage legumes such as Onobrychis viciaefolia, Astragalus adsurgens, Trifolium repens, T. pretense, Vicia sativa, V. sepium, V. unijuga, Lespedeza bicolor, Lotus cornioulatus, Melilotus albus, and M. officinalis.
引文
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