化学诱导小麦条锈菌毒性突变研究
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摘要
小麦条锈病是我国小麦上最重要的病害之一，小麦条锈病菌的毒性变异是造成小麦
品种抗锈性丧失导致病害流行主要原因。关于锈菌的毒性变异机制已提出了多种途径，
普遍认为突变是产生新毒性基因的主要途径。由于在自然界中尚未发现小麦条锈菌的有
性态，对由于毒性基因的突变而产生的新小种很难进行深入的遗传分析，所以关于小麦
条锈菌的毒性变异研究较其它锈菌少，而关于毒性突变研究则更少。本研究应用化学诱
变剂甲基磺酸乙酯(EMS)对小麦条锈菌夏孢子进行诱变处理，对小麦条锈菌受诱变剂影
响的各项生物学效应和毒性突变的特点进行研究，主要取得了以下几项结果：
1. 建立了化学诱导小麦条锈菌毒性突变的优化体系：研究中发现诱变剂对处理当代
条锈菌夏孢子的各项生物学特性有明显的削弱性影响，使其夏孢子存活力降低，致病力
减弱和繁殖能力下降。不同处理时间和浓度的夏孢子萌发率经 CurveExpert(1.3)MMF 和
Harris 两个模型拟合的曲线为倒 S 型，表明小麦条锈菌夏孢子萌发率在 EMS 处理的浓
度曲线和时间曲线上均有一个最适拟合系数，且夏孢子对低剂量 EMS 十分敏感。据此
构建的 EMS 对小麦条锈菌毒性突变的优化体系为：室温下，pH＝7.0，C=0.03mol/L ,T=6～
8min。小麦条锈菌夏孢子在此处理环境下的致死率在 80%～90%之间，根据微生物诱变
筛选最佳诱变剂量的选择标准，上述参数可作为小麦条锈菌化学诱变的主要参量。
2. 筛选了具有不同毒性变异特点的小麦条锈菌 CY17 和 CY31 突变菌系：CY17 有 5
个毒性突变菌系17M1～17M5，17M1中9个突变菌株都对早洋发生了无毒性突变；17M2
对维尔发生了毒性突变；17M3 对南大 2419 发生了毒性突变；17M4 同时对南大 2419
和水源 11 都发生了毒性突变；17M5 对阿勃、水源 11 也同时为毒性突变。CY31 有 4
个毒性突变菌系 31M1～31M4，31M1 中 6 个菌株对尤二和 T.E 为毒性突变体，31F14-1
对丹麦 1 号只为中感，而 31MutS1是 CY32；31M2 对多个品种产生了无毒性突变；31M3
和 31M4 分别为两个无毒性突变体 31C7-1和 31C7-2，它们在繁殖一代时对其筛选品种中
四有致病能力，但在繁殖 3～4 代后，其致病力丧失，且对大部分鉴别寄主的反应型也
随着降低，表明了 EMS 的多位点诱变作用。聚类图表明各突变菌系与其原始菌株的相
关系数都在 85%以上，每个菌系中都有对不同品种发生多种微小突变的菌株，说明突变
菌系与原始菌株既有一定的同源关系，又有微小的毒性变异区别，是化学诱变剂 EMS
诱导单点突变和逐步突变后的结果。而且 CY17 以无毒性突变体居多，CY31 则以毒性
突变体居多, 表明诱导后的弱毒菌株多为无毒性突变，而强毒性小种则主要为毒性变异。
3. 研究了突变体与其原始菌株的寄生适合度：发现 CY17 各突变菌株之间的差异主
要显示在夏孢子的生活能力和夏孢子在小麦叶片上的侵染和扩展能力上。根据贡献值排
2 化学诱导小麦条锈菌毒性突变研究
列的顺序为：17F08-1＞17F09-1＞17Mut-1＞17F09-2＞17F15-1＞17F15-3＞17CK。无毒性突变
菌株的寄生适合度要高，原始菌株 CY17 的寄生适合度低于各突变菌株。CY31 突变株
的各寄生适合度则由夏孢子堆长度、潜伏期、产孢期和夏孢子萌发率决定，代表了各突
变菌系夏孢子的叶面扩张能力，引起发病的能力，夏孢子堆扩展后的衰老速度及夏孢子
的存活能力。其寄生能力的次序为：31F14-3＞31MutS-3＞31C13-2＞31MutH-1＞31F14-2＞
31F14-1＞31CK＞31C17-1＞31MutH-2＞31MutS-2＞31C7-1。毒性突变株 31F14-3和 31M utS-3
的生存能力最高，而无毒性突变株 31 C7-1则为最低，原始菌株 31CK 居中，肯定了强致
病力小种在国内的流行优势。
The studies here were conducted to determine if the chemical mutagen Ethyl Methane
Sulphonate (EMS) applied to urediospores could alter the virulence of Puccinia striiformis
and if mutants could detect on rust-resistant screening wheat cultivars. The results of the
studies were reported as follows:
1. EMS has extensive biological effects on the viability, pathogencity and reproduction
of stripe rust. Germination rate of radiated spores was remarkably reduced, its disposal time
and concentration curve have shown that urediospores of stripe rust were sensitive to lower
disposal concentration and to longer time. The dose-germination curve were followed MMF
and Harris mode in CurveExpert (1.3), they were inverse S shape. Implied its has a optimized
parameter among in the space. On primary leaf of susceptible cultivars inoculated mutated
urediospres, the latent period of rust infection was extended, rust incidence and severity were
reduced, reproduction of urediospores of stripe rust decreased, and the host-pathogen
interaction changed into incompatible with low-infection type. All of responses strengthened
with the increase of disposal dose. An optimized mutagenic system of wheat stripe rust
urediospore was designed，The results indicated that when the buffer PH value was7.0 in the
normal temperature conditions, the uredospore of tripe rust disposed 7-8 minutes with
0.03mol/L EMS solution the death rate has reached 80% to 90%. This number value adapted
to the selection criterion of the optimum mutagenic dose of the microbe chemical mutagenesis
experience value.
2.Urediospores of patents culture, predominant race CY17-5 and CY31-3 were treated
by EMS, induce mutant were screened on the Chinese differential cultivars and the
near-isogenic lines (NILs) of rust. 11 and 8 mutant isolations were obtained with estimated
mutation rates 10-5～10-6. 5 mutant groups isolate form CY17-5, code 17M1 to 17M5, they
have 86% similarity with its patents culture. And 4 mutant groups form CY31-3, code 31M1
to 31M4, using 92% similarity cut-off point. The virulence of the mutants to a set of
differentials used in Chinese race survey was different from all of the known races of stripe
rust. Most mutant isolates kept stable virulence such as 17M2 to Wirgilio, 17M3 to Mentana,
17M4 to Mentana、Suwon11 and 17M5 to Abbonanza、Suwon11, 31M1 were virulent on
Jubilejina 2 and Trigo Eureka. But races in 17M1 were avirlent on Early Premium, races of
31M2、31M3 and 31M4 were detected to avirlent on several differential cultivars. The
virulences of mutants were also surveyed to the near-isogenic lines of stripe rust. From the
4 化学诱导小麦条锈菌毒性突变研究
relative responses of Chinese cultivars and NILs, the resistant genes of former can be
conclusion.
3.The relative parasitic fitness of 16 mutants was evaluated on susceptible cultivars in
this study. The results showed that three primary element attributed to CY17 races adaptability,
which represent sporulation capacity, the speed of growth and senescence of colony
respectively. CY31 was decided by urediospore germination ability, sporulation speed and the
size of uredosporus. According to the principal components analysis, the sequence of CY17
fitness was 17F08-1＞17F09-1＞17Mut-1＞17F09-2＞17F15-1＞17F15-3＞17CK, the avirulent
mutants, 17F08-1 and 17F09-1 have higher vigor. Otherwise the order of CY31, 31F14-3＞
31mutS-3＞31C13-2＞31MutH-1＞31F14-2＞31F14-1＞31CK＞31C17-1＞31MutH-2＞31MutS-2
＞31C7-1, 31F14-3 and 31MutS-3 have more energy than the avirulent ones.
The results suggested that the technique mutation of wheat stripe rust is feasibility and
credibility. Mutation is important mechanism of virulence variation in asexual population of
rust fungi.
品种抗锈性丧失导致病害流行主要原因。关于锈菌的毒性变异机制已提出了多种途径，
普遍认为突变是产生新毒性基因的主要途径。由于在自然界中尚未发现小麦条锈菌的有
性态，对由于毒性基因的突变而产生的新小种很难进行深入的遗传分析，所以关于小麦
条锈菌的毒性变异研究较其它锈菌少，而关于毒性突变研究则更少。本研究应用化学诱
变剂甲基磺酸乙酯(EMS)对小麦条锈菌夏孢子进行诱变处理，对小麦条锈菌受诱变剂影
响的各项生物学效应和毒性突变的特点进行研究，主要取得了以下几项结果：
1. 建立了化学诱导小麦条锈菌毒性突变的优化体系：研究中发现诱变剂对处理当代
条锈菌夏孢子的各项生物学特性有明显的削弱性影响，使其夏孢子存活力降低，致病力
减弱和繁殖能力下降。不同处理时间和浓度的夏孢子萌发率经 CurveExpert(1.3)MMF 和
Harris 两个模型拟合的曲线为倒 S 型，表明小麦条锈菌夏孢子萌发率在 EMS 处理的浓
度曲线和时间曲线上均有一个最适拟合系数，且夏孢子对低剂量 EMS 十分敏感。据此
构建的 EMS 对小麦条锈菌毒性突变的优化体系为：室温下，pH＝7.0，C=0.03mol/L ,T=6～
8min。小麦条锈菌夏孢子在此处理环境下的致死率在 80%～90%之间，根据微生物诱变
筛选最佳诱变剂量的选择标准，上述参数可作为小麦条锈菌化学诱变的主要参量。
2. 筛选了具有不同毒性变异特点的小麦条锈菌 CY17 和 CY31 突变菌系：CY17 有 5
个毒性突变菌系17M1～17M5，17M1中9个突变菌株都对早洋发生了无毒性突变；17M2
对维尔发生了毒性突变；17M3 对南大 2419 发生了毒性突变；17M4 同时对南大 2419
和水源 11 都发生了毒性突变；17M5 对阿勃、水源 11 也同时为毒性突变。CY31 有 4
个毒性突变菌系 31M1～31M4，31M1 中 6 个菌株对尤二和 T.E 为毒性突变体，31F14-1
对丹麦 1 号只为中感，而 31MutS1是 CY32；31M2 对多个品种产生了无毒性突变；31M3
和 31M4 分别为两个无毒性突变体 31C7-1和 31C7-2，它们在繁殖一代时对其筛选品种中
四有致病能力，但在繁殖 3～4 代后，其致病力丧失，且对大部分鉴别寄主的反应型也
随着降低，表明了 EMS 的多位点诱变作用。聚类图表明各突变菌系与其原始菌株的相
关系数都在 85%以上，每个菌系中都有对不同品种发生多种微小突变的菌株，说明突变
菌系与原始菌株既有一定的同源关系，又有微小的毒性变异区别，是化学诱变剂 EMS
诱导单点突变和逐步突变后的结果。而且 CY17 以无毒性突变体居多，CY31 则以毒性
突变体居多, 表明诱导后的弱毒菌株多为无毒性突变，而强毒性小种则主要为毒性变异。
3. 研究了突变体与其原始菌株的寄生适合度：发现 CY17 各突变菌株之间的差异主
要显示在夏孢子的生活能力和夏孢子在小麦叶片上的侵染和扩展能力上。根据贡献值排
2 化学诱导小麦条锈菌毒性突变研究
列的顺序为：17F08-1＞17F09-1＞17Mut-1＞17F09-2＞17F15-1＞17F15-3＞17CK。无毒性突变
菌株的寄生适合度要高，原始菌株 CY17 的寄生适合度低于各突变菌株。CY31 突变株
的各寄生适合度则由夏孢子堆长度、潜伏期、产孢期和夏孢子萌发率决定，代表了各突
变菌系夏孢子的叶面扩张能力，引起发病的能力，夏孢子堆扩展后的衰老速度及夏孢子
的存活能力。其寄生能力的次序为：31F14-3＞31MutS-3＞31C13-2＞31MutH-1＞31F14-2＞
31F14-1＞31CK＞31C17-1＞31MutH-2＞31MutS-2＞31C7-1。毒性突变株 31F14-3和 31M utS-3
的生存能力最高，而无毒性突变株 31 C7-1则为最低，原始菌株 31CK 居中，肯定了强致
病力小种在国内的流行优势。
The studies here were conducted to determine if the chemical mutagen Ethyl Methane
Sulphonate (EMS) applied to urediospores could alter the virulence of Puccinia striiformis
and if mutants could detect on rust-resistant screening wheat cultivars. The results of the
studies were reported as follows:
1. EMS has extensive biological effects on the viability, pathogencity and reproduction
of stripe rust. Germination rate of radiated spores was remarkably reduced, its disposal time
and concentration curve have shown that urediospores of stripe rust were sensitive to lower
disposal concentration and to longer time. The dose-germination curve were followed MMF
and Harris mode in CurveExpert (1.3), they were inverse S shape. Implied its has a optimized
parameter among in the space. On primary leaf of susceptible cultivars inoculated mutated
urediospres, the latent period of rust infection was extended, rust incidence and severity were
reduced, reproduction of urediospores of stripe rust decreased, and the host-pathogen
interaction changed into incompatible with low-infection type. All of responses strengthened
with the increase of disposal dose. An optimized mutagenic system of wheat stripe rust
urediospore was designed，The results indicated that when the buffer PH value was7.0 in the
normal temperature conditions, the uredospore of tripe rust disposed 7-8 minutes with
0.03mol/L EMS solution the death rate has reached 80% to 90%. This number value adapted
to the selection criterion of the optimum mutagenic dose of the microbe chemical mutagenesis
experience value.
2.Urediospores of patents culture, predominant race CY17-5 and CY31-3 were treated
by EMS, induce mutant were screened on the Chinese differential cultivars and the
near-isogenic lines (NILs) of rust. 11 and 8 mutant isolations were obtained with estimated
mutation rates 10-5～10-6. 5 mutant groups isolate form CY17-5, code 17M1 to 17M5, they
have 86% similarity with its patents culture. And 4 mutant groups form CY31-3, code 31M1
to 31M4, using 92% similarity cut-off point. The virulence of the mutants to a set of
differentials used in Chinese race survey was different from all of the known races of stripe
rust. Most mutant isolates kept stable virulence such as 17M2 to Wirgilio, 17M3 to Mentana,
17M4 to Mentana、Suwon11 and 17M5 to Abbonanza、Suwon11, 31M1 were virulent on
Jubilejina 2 and Trigo Eureka. But races in 17M1 were avirlent on Early Premium, races of
31M2、31M3 and 31M4 were detected to avirlent on several differential cultivars. The
virulences of mutants were also surveyed to the near-isogenic lines of stripe rust. From the
4 化学诱导小麦条锈菌毒性突变研究
relative responses of Chinese cultivars and NILs, the resistant genes of former can be
conclusion.
3.The relative parasitic fitness of 16 mutants was evaluated on susceptible cultivars in
this study. The results showed that three primary element attributed to CY17 races adaptability,
which represent sporulation capacity, the speed of growth and senescence of colony
respectively. CY31 was decided by urediospore germination ability, sporulation speed and the
size of uredosporus. According to the principal components analysis, the sequence of CY17
fitness was 17F08-1＞17F09-1＞17Mut-1＞17F09-2＞17F15-1＞17F15-3＞17CK, the avirulent
mutants, 17F08-1 and 17F09-1 have higher vigor. Otherwise the order of CY31, 31F14-3＞
31mutS-3＞31C13-2＞31MutH-1＞31F14-2＞31F14-1＞31CK＞31C17-1＞31MutH-2＞31MutS-2
＞31C7-1, 31F14-3 and 31MutS-3 have more energy than the avirulent ones.
The results suggested that the technique mutation of wheat stripe rust is feasibility and
credibility. Mutation is important mechanism of virulence variation in asexual population of
rust fungi.
引文
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