落叶型棉花黄萎病菌在华中棉区广泛分布及其原因分析
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摘要
近年来,华中棉区棉花黄萎病(Verticillium Wilt)发生越来越严重,特别是田间广泛出现了落叶型棉花黄萎病症状。对病原菌遗传多样性的认识是有效防治棉花黄萎病的关键。但是目前对华中棉区棉花黄萎病菌(Verticillium dahliae)遗传多样性的认识十分有限。鉴此,本研究在分析华中棉区棉花黄萎病菌遗传多样性的基础上进一步分析落叶型棉花黄萎病菌广泛分布的原因。
针对第一个问题,本研究从湖北省、湖南省、安徽省、江西省、江苏省共分离和收集到341个棉花黄萎病菌菌株。从3个层面揭示棉花黄萎病菌的遗传多样性。其中有来源于华中棉区30个县(市)52个取样点129个菌株,来源于湖北省7块不同发病率棉田86个菌株,来源于湖北省6个典型黄萎病株的126个菌株。本研究分别对3个群体进行了营养亲和性(Vegetative compatibility grouping,简称VCG)分析和分子标记分析。
在第一层面的棉黄萎病菌菌株中,VCG1A型占94.6%,VCG2型占5.4%。在第二层面的棉黄萎病菌菌株中,源于5块棉田的病菌均为VCG1A型。在源于另外两块棉田的菌株中,VCG1A(?)型分别占94%和90.9%,而VCG2型分别占6.0%和9.1%。在第三层面的棉黄萎病菌菌株中,来源于6个典型病株的菌株全部属于VCG1A型。这些结果说明VCG1A型菌株已在华中棉区广泛分布。
在陆地棉品种鄂棉24和银瑞361上测定了VCG1A型和VCG2型代表性菌株的致病力。结果表明：接种14天后,VCG1A型和VCG2型菌株均能引起棉苗发病,VCG1A型菌株引起典型的落叶症状(defoliation,简称D),因而其致病型属于D型。而VCG2型菌株则没有引起落叶症状(non-defoliation,简称ND),因而其致病型属于ND型。这些结果说明：VCG1A型和VCG2型菌株的致病力存在明显分化。
针对第二个问题,我们评价了湖北省及其周边棉区的主栽品种对棉花黄萎病菌的抗感性以及棉花黄萎病菌对温度的反应。结果表明：供试的12个棉花品种对VCG1A/D型菌株4TM6-15极其感病,而6个棉花品种(C111、湘杂棉3号F2、富抗杂3号F1、湘杂棉2号F2、盛杂棉9号F1、华丰杂1号F1)对VCG2/ND型菌株1HN-1感病,3个棉花品种(鄂杂棉19F1、富抗杂518F1、鄂杂棉4号)对菌株1HN-1中感,另外3个棉花品种(鄂杂棉3号、福棉2号、丰棉03F1)对菌株1HN-1抵抗。结果还表明：在株高、根重、叶重和茎重等4个指标上,VCG1A/D型菌株接种的棉苗显著(P<0.05)低于VCG2型菌株接种的棉苗。这些结果表明：目前在湖北省及其周边棉区推广的棉花品种对VCG1A/D型菌株高感。
在棉花黄萎病菌对温度反应方面,VCG1A/D型和VCG2/ND型菌株菌丝生长的最适温度均为25℃。在308下,VCG1A/D型菌株在PDA和CDA上生长速度显著(P<0.05)快于非落叶型菌株。在10～33℃条件下,VCG1A/D型菌株分生孢子萌发率显著(P<0.05)高于VCG2/ND型菌株。在25/25℃、27/27℃、30/25℃(昼/夜温度)条件下,VCG1A/D型菌株导致棉花植株落叶,而VCG2/ND型菌株也导致棉花植株发病,但不引起棉花落叶。VCG1A/D型菌株引起的病害进程曲线下面积值(Area under disease progress curve,简称AUDPC)和维管束变色指数值(Vascular discoloration index,简称VDI)显著(P<0.05)高于VCG2/ND型菌株引起的AUDPC值和VDI值。在30/30℃、33/27℃条件下,在鄂棉24上,VCG1A/D型菌株和VCG2/ND型菌株引起轻微病害症状,VCG1A/D型菌株引起的VDI值显著(P<0.05)高于VCG2/ND型菌株。在银瑞361上,VCG1A/D型和VCG2/ND型菌株均没有引起病害。在33/33℃下,两类菌株接种棉苗后,棉苗叶片和维管束都没有显现任何病害症状。这些结果表明：VCG1A/D型菌株对高温(30℃)的忍耐性较VCG2/ND型菌株强,可能更适应湖北省的气候条件。
研究结果说明：VCG1A/D型菌株在湖北省已广泛流行。棉花品种对VCG1A/D型菌株的高度敏感性以及VCG1A/D型菌株对湖北省气候条件的适应性可能是造成VCG1A/D型菌株广泛流行的原因。
Verticillium wilt (VW) of cotton occurred more and more severe in central China, paticularly the defoliating symptoms appeared in the fields. Knowledge on genetic and virulence divercity in Verticillium dahliae is the key for efficient management of VW. A total of341strains of V. dahliae were collected from cotton in central China for studying the genetic diversity of V. dahliae at three different levels. The first level included129strains from52districts in30counties/cities in central China. The second level included86strains of V. dahliae from7fields with different incidence of VW. The third level included126strains of V. dahliae from6plants. Vegetative compatibility grouping (VCG) assessment and PCR-based molecular pathotyping were used to evaluate the genetic diversity of V. dahliae.
The results indicated that in the first level, VCG1A and VCG2isolates accounted for94.6and5.4%, respectively; In the second level, VCG1A isolates prevailed (100%) in5cotton fields, in the remaining2fields, two VCGs were identified with the VCG1A isolates accounting for90.9and94%, respectively. In the third level, the VCG1A isolates accounted for100%.
Pathogenicity of19isolates of V. dahliae, including10VCG1A isolates and9VCG2isolates, was tested on seedlings of two upland cotton cultivars E Mian24and Yin Rui361. The isolates from two VCGs were virulent on the two cotton cultivars. After two weeks of post-inoculation (25℃), the VCG1A isolates caused defoliation symptoms, whereas the VCG2isolates caused wilt disease, but did not cause any defoliation symptoms. Therefore, the VCG1A is the severe pathotype, whereas the VCG2is the mild pathotype.
A study was conducted to determine the level of resistance of12commercial cotton cultivars to infection by the both pathotypes of V. dahliae under growth chamber conditions.(ⅰ) All the12cotton cultivars were susceptible to the defoliating pathotype (D-pathotype) of V. dahliae. All the12cultivars were more susceptible to the D-pathotype isolates than to non-defoliating pathotype (ND-pathotype) isolates. Six of evaluated cultivars (including C111, Xiang Za Mian3, Fu Kang Za3, Xiang Za Mian2, Sheng Za Mian9, Hua Feng Za1) were susceptible to the ND-pathotype isolates of V. dahliae at different levels. Three of evaluated cultivars (E Za Mian19, Fu Kang Za518, E Za Mian4) were moderately susceptible to ND-pathotype isolates. Three of evaluated cultivars (E Za Mian3, Fu Mian2, Feng Mian.03) were resistant to the ND-pathotype isolates;(ⅱ) A resistant related phenotypic reaction to the disease was quantified by using four growth parameters (plant height, leaf weight, stem weight, total shoot weight) measured27days after inoculation. The four growth parameters of plants inoculated D pathotype isolates were lower than that of plants inoculated ND pathotype isolates. The percentage of decrease in leaf weight was found to be the best parameter to indicate cotton resistance.
Effect of temperature on conidial germination, mycelial growth and aggressiveness in two VCGs of V. dahliae was studied,(ⅰ) VCG1A/D isolates and VCG2/ND isolates had the same optimum temperature of25℃for mycelial growth on PDA and CDA media. VCG2/ND isolates grew faster than VCG1A/D isolates at20to25℃, but VCG1A/D isolates grew faster than VCG2/ND isolates at30℃;(ⅱ) Conidia of VCG1A/D isolates germinated more rapidly than conidia of VCG2/ND isolates at temperatures of10-33℃;(ⅲ) The VCG1A/D isolates induced defoliation at25/25℃,27/27℃,30/25℃(day/night temperatures) on two cotton cultivars, whereas VCG2/ND pathotype isolates did not induce any defoliation symptoms on the cotton cultivars. Under30/30℃or33/27℃, the leaf symptoms were not visible. However, the VCG1A/D isolates caused higher vascular discoloration index (VDI) on E Mian24than VCG2/ND isolates. All the D-and ND-pathotypes isolates were not pathogenic on Yin Rui361at33/33℃.
This study suggests that the VCG1A/D pathotype has become prevalent on cotton in Hubei Province and the surrounding areas. High susceptibility of the cotton cultivars and high aggressiveness of the VCG1A/D pathotype of V. dahliae on cotton under25-33℃might be responsible for the rapid spread and the prevalence of this pathotype in Hubei.
针对第一个问题,本研究从湖北省、湖南省、安徽省、江西省、江苏省共分离和收集到341个棉花黄萎病菌菌株。从3个层面揭示棉花黄萎病菌的遗传多样性。其中有来源于华中棉区30个县(市)52个取样点129个菌株,来源于湖北省7块不同发病率棉田86个菌株,来源于湖北省6个典型黄萎病株的126个菌株。本研究分别对3个群体进行了营养亲和性(Vegetative compatibility grouping,简称VCG)分析和分子标记分析。
在第一层面的棉黄萎病菌菌株中,VCG1A型占94.6%,VCG2型占5.4%。在第二层面的棉黄萎病菌菌株中,源于5块棉田的病菌均为VCG1A型。在源于另外两块棉田的菌株中,VCG1A(?)型分别占94%和90.9%,而VCG2型分别占6.0%和9.1%。在第三层面的棉黄萎病菌菌株中,来源于6个典型病株的菌株全部属于VCG1A型。这些结果说明VCG1A型菌株已在华中棉区广泛分布。
在陆地棉品种鄂棉24和银瑞361上测定了VCG1A型和VCG2型代表性菌株的致病力。结果表明：接种14天后,VCG1A型和VCG2型菌株均能引起棉苗发病,VCG1A型菌株引起典型的落叶症状(defoliation,简称D),因而其致病型属于D型。而VCG2型菌株则没有引起落叶症状(non-defoliation,简称ND),因而其致病型属于ND型。这些结果说明：VCG1A型和VCG2型菌株的致病力存在明显分化。
针对第二个问题,我们评价了湖北省及其周边棉区的主栽品种对棉花黄萎病菌的抗感性以及棉花黄萎病菌对温度的反应。结果表明：供试的12个棉花品种对VCG1A/D型菌株4TM6-15极其感病,而6个棉花品种(C111、湘杂棉3号F2、富抗杂3号F1、湘杂棉2号F2、盛杂棉9号F1、华丰杂1号F1)对VCG2/ND型菌株1HN-1感病,3个棉花品种(鄂杂棉19F1、富抗杂518F1、鄂杂棉4号)对菌株1HN-1中感,另外3个棉花品种(鄂杂棉3号、福棉2号、丰棉03F1)对菌株1HN-1抵抗。结果还表明：在株高、根重、叶重和茎重等4个指标上,VCG1A/D型菌株接种的棉苗显著(P<0.05)低于VCG2型菌株接种的棉苗。这些结果表明：目前在湖北省及其周边棉区推广的棉花品种对VCG1A/D型菌株高感。
在棉花黄萎病菌对温度反应方面,VCG1A/D型和VCG2/ND型菌株菌丝生长的最适温度均为25℃。在308下,VCG1A/D型菌株在PDA和CDA上生长速度显著(P<0.05)快于非落叶型菌株。在10～33℃条件下,VCG1A/D型菌株分生孢子萌发率显著(P<0.05)高于VCG2/ND型菌株。在25/25℃、27/27℃、30/25℃(昼/夜温度)条件下,VCG1A/D型菌株导致棉花植株落叶,而VCG2/ND型菌株也导致棉花植株发病,但不引起棉花落叶。VCG1A/D型菌株引起的病害进程曲线下面积值(Area under disease progress curve,简称AUDPC)和维管束变色指数值(Vascular discoloration index,简称VDI)显著(P<0.05)高于VCG2/ND型菌株引起的AUDPC值和VDI值。在30/30℃、33/27℃条件下,在鄂棉24上,VCG1A/D型菌株和VCG2/ND型菌株引起轻微病害症状,VCG1A/D型菌株引起的VDI值显著(P<0.05)高于VCG2/ND型菌株。在银瑞361上,VCG1A/D型和VCG2/ND型菌株均没有引起病害。在33/33℃下,两类菌株接种棉苗后,棉苗叶片和维管束都没有显现任何病害症状。这些结果表明：VCG1A/D型菌株对高温(30℃)的忍耐性较VCG2/ND型菌株强,可能更适应湖北省的气候条件。
研究结果说明：VCG1A/D型菌株在湖北省已广泛流行。棉花品种对VCG1A/D型菌株的高度敏感性以及VCG1A/D型菌株对湖北省气候条件的适应性可能是造成VCG1A/D型菌株广泛流行的原因。
Verticillium wilt (VW) of cotton occurred more and more severe in central China, paticularly the defoliating symptoms appeared in the fields. Knowledge on genetic and virulence divercity in Verticillium dahliae is the key for efficient management of VW. A total of341strains of V. dahliae were collected from cotton in central China for studying the genetic diversity of V. dahliae at three different levels. The first level included129strains from52districts in30counties/cities in central China. The second level included86strains of V. dahliae from7fields with different incidence of VW. The third level included126strains of V. dahliae from6plants. Vegetative compatibility grouping (VCG) assessment and PCR-based molecular pathotyping were used to evaluate the genetic diversity of V. dahliae.
The results indicated that in the first level, VCG1A and VCG2isolates accounted for94.6and5.4%, respectively; In the second level, VCG1A isolates prevailed (100%) in5cotton fields, in the remaining2fields, two VCGs were identified with the VCG1A isolates accounting for90.9and94%, respectively. In the third level, the VCG1A isolates accounted for100%.
Pathogenicity of19isolates of V. dahliae, including10VCG1A isolates and9VCG2isolates, was tested on seedlings of two upland cotton cultivars E Mian24and Yin Rui361. The isolates from two VCGs were virulent on the two cotton cultivars. After two weeks of post-inoculation (25℃), the VCG1A isolates caused defoliation symptoms, whereas the VCG2isolates caused wilt disease, but did not cause any defoliation symptoms. Therefore, the VCG1A is the severe pathotype, whereas the VCG2is the mild pathotype.
A study was conducted to determine the level of resistance of12commercial cotton cultivars to infection by the both pathotypes of V. dahliae under growth chamber conditions.(ⅰ) All the12cotton cultivars were susceptible to the defoliating pathotype (D-pathotype) of V. dahliae. All the12cultivars were more susceptible to the D-pathotype isolates than to non-defoliating pathotype (ND-pathotype) isolates. Six of evaluated cultivars (including C111, Xiang Za Mian3, Fu Kang Za3, Xiang Za Mian2, Sheng Za Mian9, Hua Feng Za1) were susceptible to the ND-pathotype isolates of V. dahliae at different levels. Three of evaluated cultivars (E Za Mian19, Fu Kang Za518, E Za Mian4) were moderately susceptible to ND-pathotype isolates. Three of evaluated cultivars (E Za Mian3, Fu Mian2, Feng Mian.03) were resistant to the ND-pathotype isolates;(ⅱ) A resistant related phenotypic reaction to the disease was quantified by using four growth parameters (plant height, leaf weight, stem weight, total shoot weight) measured27days after inoculation. The four growth parameters of plants inoculated D pathotype isolates were lower than that of plants inoculated ND pathotype isolates. The percentage of decrease in leaf weight was found to be the best parameter to indicate cotton resistance.
Effect of temperature on conidial germination, mycelial growth and aggressiveness in two VCGs of V. dahliae was studied,(ⅰ) VCG1A/D isolates and VCG2/ND isolates had the same optimum temperature of25℃for mycelial growth on PDA and CDA media. VCG2/ND isolates grew faster than VCG1A/D isolates at20to25℃, but VCG1A/D isolates grew faster than VCG2/ND isolates at30℃;(ⅱ) Conidia of VCG1A/D isolates germinated more rapidly than conidia of VCG2/ND isolates at temperatures of10-33℃;(ⅲ) The VCG1A/D isolates induced defoliation at25/25℃,27/27℃,30/25℃(day/night temperatures) on two cotton cultivars, whereas VCG2/ND pathotype isolates did not induce any defoliation symptoms on the cotton cultivars. Under30/30℃or33/27℃, the leaf symptoms were not visible. However, the VCG1A/D isolates caused higher vascular discoloration index (VDI) on E Mian24than VCG2/ND isolates. All the D-and ND-pathotypes isolates were not pathogenic on Yin Rui361at33/33℃.
This study suggests that the VCG1A/D pathotype has become prevalent on cotton in Hubei Province and the surrounding areas. High susceptibility of the cotton cultivars and high aggressiveness of the VCG1A/D pathotype of V. dahliae on cotton under25-33℃might be responsible for the rapid spread and the prevalence of this pathotype in Hubei.
引文
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