CIMMYT小麦品种Chapio条锈病成株抗性QTL定位及卷叶材料CMH83分子细胞学研究
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摘要
小麦是世界上分布最广、种植面积最大的粮食作物。由小麦条锈病菌(Puccinia striiformis f. sp. Tritici)引发的条锈病,是我国小麦最严重的病害之一,中国占有世界上最大的小麦条锈病流行区,约2.0千万公顷以上。选育抗病品种是防治小麦条锈病最为经济、安全、有效的措施,而成株抗性基因的合理利用是培育持久抗性品种的有效途径。因此,研究小麦条锈病成株抗性基因的遗传、发掘成株抗性基因及其紧密连锁的分子标记,能加快培育持久抗性品种,控制条锈病危害。小黑麦(X Triticosecale Wittmack)是人工杂交合成的新物种。小黑麦具有优良特性,穗大、粒多、抗逆性强、抗病性好等优点,是高产饲料作物,也是改良小麦品种的重要种质资源。植物叶片卷曲是叶片性状中的一个重要组成部分,是与植物抗旱相关的、对逆境刺激的一种反应,研究卷叶性状可以为抗旱机制及抗旱育种提供信息。
本研究对Avocet-YrA/Chapio的277个F6重组自交系(RILs)及亲本进行苗期及成株期条锈病抗性鉴定,田间表型鉴定放在4个试验点,再根据严重度进行抗性基因遗传分析,及利用完备区间作图法(ICIM)对Avocet-YrA/Chapio群体进行QTL定位及上位性分析；同时对小黑麦卷叶材料CMH83进行了分子细胞学特性研究、醇溶蛋白研究、叶片相对含水量(RWC)、失水率(WLR)及卷叶特性初步遗传研究。主要研究结果如下：
1、Chapio对条锈病表现为成株抗性,遗传分析表明其抗性由4对不完全显性基因控制。
供试材料Chapio苗期对条锈菌生理小种条中32(CYR32)表现为感病；在成株期,表现为高抗(HR)条锈病,严重度不大于5%；感病亲本Avocet-YrA在苗期和成株期感病；Avocet-YrA/Chapio的277个F6重组自交系(RILs)在苗期全部感病,成株期表现为抗感分离。对4个试验点间的相关性分析表明,在4个试验点间的条锈病严重度相关性达极显著水平(P<0.001)。次数分布图显示,严重度呈连续变异,但峰值略偏向成株抗性亲本一边呈偏正态分布,表明Chapio可能含有主效成株抗性基因。将RILs在墨西哥CIMMYT的严重度与在四川3个点锈病严重度的平均值相比较,发现有较多的品系差异较大,其中有22个品系的严重度差值超过了40%,最高达到了78.3%,说明Chapio有1个或几个成株抗性基因能鉴别墨西哥对锈菌小种YRMV09与中国小种CYR32。
根据4个试验点条锈病感病严重度调查数据,利用孟德尔遗传模型分别对4个试验点的277个F6重组自交系(RILs)分别进行遗传分析,经χ2测验表明,Chapio在每个试验点的条锈病成株抗性都表现为由4对基因控制(P>0.05)。结合Avocet-YrA x Chapio的正反交F1锈病严重度表现为40-50%,说明Chapio的成株抗性是由4对不完全显性基因控制。
2、发现4个加性成株抗性QTL和1对上位性QTL(QYr.saas-3BS x QYr.saas-7DS),其中QYr.saas-3DL是一个新QTL
在4个试验点上共检测到4个条锈病成株抗性QTL,都具有加性效应且都是由抗病亲本Chapio提供,它们位于3BS.3BL.5BL.7DS染色体上,分别命名为Qyr.saas-5BL、QYr.saas-3BS、Qyr.saas-7DS、Qyr.saas-3DL。QYr.saas-7DS最稳定地存在于所有试验点中,它介于SSR标记CSL V34-Xgwm295之间,且与这两个标记紧密连锁,平均加性效应为17.35,在各试验点可解释的表型变异分别为28.18-48.93%;QYr.saas-3BS、QYr.saas-3DL、QYr.saas-5BL分别介于SSR标记Xbarc147-Xwms533,Xgdm8-GDM128、Xgwm46-Xbarc267之间,连锁距离分别为2cM、10cM、38cM,平均加性效应分别为8.40、6.55、4.88,在各试验点可解释的表型变异分别为7.30-17.00%、3.84-12.44%和2.64-4.76%。其中QYr.saas-3DL是新QTL,其余还需进一步研究确定。该群体还检测到1对上位性QTL(QYr.saas-3BS x QYr.saas-7DS),并稳定地存在于3个环境及总平均值中,分别为可解释表现型变异的2.59-7.79%。
3、发现一个具有卷叶特性的次级小黑麦材料CMH83,其卷叶是显性性状。
卷叶现象是作物对干旱的一种反应。通过表型观察、细胞学和生理学研究,我们发现了一个卷叶六倍体次级小黑麦(X Triticosecale Wittmack)材料CMH83,它具有矮杆、卷叶以及叶片窄的表型特征。种子醇溶蛋白电泳研究表明,CMH83是一个遗传稳定材料。顺序Giemsa-C带和基因组原位杂交研究发现,CMH83具有12条黑麦染色体,其中两条黑麦染色体的异染色质端带丢失。在干旱胁迫下,CMH83的叶片相对含水量(RWC)和失水率(WLR)比小麦品种失水快,这种现象与卷叶特性相关。初步遗传研究表明,CMH83的卷叶特性由显性基因控制。次级小黑麦品系CMH83可被用于研究小黑麦的抗旱机理,以及干旱胁迫相关基因及其遗传机制。
Wheat(Triticum spp.) was the largest area crop in the world. Stripe rust, caused by the fungus Puccinia striiformis tritici is a widespread disease of wheat that caused significant yield losses annually in many wheat-growing regions of the world. It also was one of the most important wheat diseases in China, its epidemic area can reach above 2.0×107 hm2. Breeding resistant varieties are more effective, economic and safe method to control it, particularly for cultivars with adult-plant resistance (APR). Therefore, one of the aims of this study were to genetic analysis and identify the quantitative trait loci(QTLs) for APR to stripe rust in wheat, and to assess the stability of the QTL effects detected across 4 environments. This knowledge could lead to a more efficient strategy in wheat breeding for durable resistance against stripe rust. Triticale (X Triticosecale Wittmack) is a man-made species with a short history of just over 100 years since the first attempt was made to produce an artificial hybrid. Triticale have good agronomic traits such as large head, more grains, resistance to biotic and abiotic stresses. Leaf rolling may be associated with improved grain yield in some drought situations. Identification of different kinds of rolling leaf types provides a theoretical basis and technical support for leaf shape improvement and also for crop drought tolerance.
277 RILs F6 developed from Avocet-YrA/Chapio were used to genetic analysis and QTL mapping with Inclusive Composite Interval Mapping Method (IciMapping, ICIM, V3.0)according to rust severity at adult plant stage at 4 environments. At same time, Rolling leaf material CMH83 from CIMMYT was studied by phenotypic, cytological and physiological analysis. The results are as follow:
1. Chapio were controlled by four pairs incomplete dominance adult-plant resistance (APR) genes against stripe rust
Chapio was susceptible to stripe rust race (CYR32) at seedling stage, and high resistance at adult-plant resistance(APR), all of 277 RILs F6 developed from Avocet-YrA/Chapio were susceptible to stripe rust race (CYR32), and segregated at adult plant stage. Correlation analysis showed that correlation coefficients were significant among 4 environments(P<0.001). Frequency distribution was a littler skew distribution which suggested that Chapio probably contained larger effect gene for APR. Stripe rust severity in some lines showed different value between CIMMYT and China, particularly for 22 lines which difference reached 40-78.3%. It showed that one or several genes in Chapio can identify the races between Chinese race CYR32 and Mexican race YRMV09.
Genetic analysis suggested that Chapio contained 4 APR genes by Chi-squared analyses (P>0.05). Rust severity were 40-50% in Avocet-TrA x Chapio F1 and Chapio x Avocet-YrA F1. Therefore, those result suggested that there were four pairs incomplete dominance adult-plant resistance(APR) genes against stripe rust in CIMMYT variety Chapio.
2. Four additive APR QTLs and one pair epistatic interactions QTL had been detected, QYr.saas-3DL should be a new QTL
Four QTLs for APR against stripe rust have been detected by Inclusive Composite Interval Mapping Method on chromosomes 3BS,3DL,5BL and 7DS in 4 environments, designated QYr.saas-3BS, QYr,saas-3DL, QYr.saas-5BL and QYr.saas-7DS, respectively, explaining from 2.64 to 48.93% of the phenotypic variance. All QTLs were from Chapio. The QTL QYr.saas-3DL, different from all QTLs reported previously, was flanked by SSR markers Xgdm8 and GDM128, with a genetic distance of 10 cM, which accounted for 3.84-12.44% of phenotypic variance. The QTL QYr.caas-7DS flanked by markers csLV34 and Xgwm295, which were closely linked to QYr.caas-7DS which showed the largest effect for resistance to stripe rust and accounted for 28.18-48.93% of phenotypic variance. The QTLs QYr.saas-3BS and QYr.saas-5BL flanked by markers Xbarc147 and Xwms533, Xgwm46 and Xbarc267, with a genetic distance of 2.0 cM and 38 cM, which accounted for 7.30-17.00% and 2.64-4.76% of phenotypic variance, respectively. QYr.saas-3DL should be a new QTL, others needed to further confirm. One pair epistatic interactions QTLs (QYr.saas-3BS x QYr.saas-7DS) had been detected, which accounted for 2.59-7.79% of the phenotypic variance in 3 environments and average value. QTLs QYr.saas-3BS, QYr.saas-3DL, and QYr.saas-7DS could be used in marker-assisted selection for APR to striper rust in wheat breeding programs.
3. Leaf rolling occurs in some cereal genotypes in response to drought. In the present study, we identified a rolling leaf genotype CMH83 of hexaploid Triticale (X Triticosecale Wittmack), that exhibited reduced plant height, rolling and narrow leaves, by phenotypic, cytological and physiological analysis. Gliadin electrophoresis of seeds protein showed that CMH83 was genetically stable. The sequential Giemsa-C banding and genomic in situ hybridization showed that the CMH83 contained 12 rye chromosomes, of which two pairs of chromosomes had reduced telomeric heterochromatin bands. The test of leaf relative water content (RWC) and water loss rate (WLR) of CMH83 compared with those of wheat cultivars indicated that rapid water loss after drought stress in CMH83 could be associated with the leaf rolling phenotypes. Leaf rolling phenotype of CMH83 was a dominant trait in our preliminary inheritance studies. The triticale line CMH83 can be used to study the mechanism of triticale for drought resistances.
本研究对Avocet-YrA/Chapio的277个F6重组自交系(RILs)及亲本进行苗期及成株期条锈病抗性鉴定,田间表型鉴定放在4个试验点,再根据严重度进行抗性基因遗传分析,及利用完备区间作图法(ICIM)对Avocet-YrA/Chapio群体进行QTL定位及上位性分析；同时对小黑麦卷叶材料CMH83进行了分子细胞学特性研究、醇溶蛋白研究、叶片相对含水量(RWC)、失水率(WLR)及卷叶特性初步遗传研究。主要研究结果如下：
1、Chapio对条锈病表现为成株抗性,遗传分析表明其抗性由4对不完全显性基因控制。
供试材料Chapio苗期对条锈菌生理小种条中32(CYR32)表现为感病；在成株期,表现为高抗(HR)条锈病,严重度不大于5%；感病亲本Avocet-YrA在苗期和成株期感病；Avocet-YrA/Chapio的277个F6重组自交系(RILs)在苗期全部感病,成株期表现为抗感分离。对4个试验点间的相关性分析表明,在4个试验点间的条锈病严重度相关性达极显著水平(P<0.001)。次数分布图显示,严重度呈连续变异,但峰值略偏向成株抗性亲本一边呈偏正态分布,表明Chapio可能含有主效成株抗性基因。将RILs在墨西哥CIMMYT的严重度与在四川3个点锈病严重度的平均值相比较,发现有较多的品系差异较大,其中有22个品系的严重度差值超过了40%,最高达到了78.3%,说明Chapio有1个或几个成株抗性基因能鉴别墨西哥对锈菌小种YRMV09与中国小种CYR32。
根据4个试验点条锈病感病严重度调查数据,利用孟德尔遗传模型分别对4个试验点的277个F6重组自交系(RILs)分别进行遗传分析,经χ2测验表明,Chapio在每个试验点的条锈病成株抗性都表现为由4对基因控制(P>0.05)。结合Avocet-YrA x Chapio的正反交F1锈病严重度表现为40-50%,说明Chapio的成株抗性是由4对不完全显性基因控制。
2、发现4个加性成株抗性QTL和1对上位性QTL(QYr.saas-3BS x QYr.saas-7DS),其中QYr.saas-3DL是一个新QTL
在4个试验点上共检测到4个条锈病成株抗性QTL,都具有加性效应且都是由抗病亲本Chapio提供,它们位于3BS.3BL.5BL.7DS染色体上,分别命名为Qyr.saas-5BL、QYr.saas-3BS、Qyr.saas-7DS、Qyr.saas-3DL。QYr.saas-7DS最稳定地存在于所有试验点中,它介于SSR标记CSL V34-Xgwm295之间,且与这两个标记紧密连锁,平均加性效应为17.35,在各试验点可解释的表型变异分别为28.18-48.93%;QYr.saas-3BS、QYr.saas-3DL、QYr.saas-5BL分别介于SSR标记Xbarc147-Xwms533,Xgdm8-GDM128、Xgwm46-Xbarc267之间,连锁距离分别为2cM、10cM、38cM,平均加性效应分别为8.40、6.55、4.88,在各试验点可解释的表型变异分别为7.30-17.00%、3.84-12.44%和2.64-4.76%。其中QYr.saas-3DL是新QTL,其余还需进一步研究确定。该群体还检测到1对上位性QTL(QYr.saas-3BS x QYr.saas-7DS),并稳定地存在于3个环境及总平均值中,分别为可解释表现型变异的2.59-7.79%。
3、发现一个具有卷叶特性的次级小黑麦材料CMH83,其卷叶是显性性状。
卷叶现象是作物对干旱的一种反应。通过表型观察、细胞学和生理学研究,我们发现了一个卷叶六倍体次级小黑麦(X Triticosecale Wittmack)材料CMH83,它具有矮杆、卷叶以及叶片窄的表型特征。种子醇溶蛋白电泳研究表明,CMH83是一个遗传稳定材料。顺序Giemsa-C带和基因组原位杂交研究发现,CMH83具有12条黑麦染色体,其中两条黑麦染色体的异染色质端带丢失。在干旱胁迫下,CMH83的叶片相对含水量(RWC)和失水率(WLR)比小麦品种失水快,这种现象与卷叶特性相关。初步遗传研究表明,CMH83的卷叶特性由显性基因控制。次级小黑麦品系CMH83可被用于研究小黑麦的抗旱机理,以及干旱胁迫相关基因及其遗传机制。
Wheat(Triticum spp.) was the largest area crop in the world. Stripe rust, caused by the fungus Puccinia striiformis tritici is a widespread disease of wheat that caused significant yield losses annually in many wheat-growing regions of the world. It also was one of the most important wheat diseases in China, its epidemic area can reach above 2.0×107 hm2. Breeding resistant varieties are more effective, economic and safe method to control it, particularly for cultivars with adult-plant resistance (APR). Therefore, one of the aims of this study were to genetic analysis and identify the quantitative trait loci(QTLs) for APR to stripe rust in wheat, and to assess the stability of the QTL effects detected across 4 environments. This knowledge could lead to a more efficient strategy in wheat breeding for durable resistance against stripe rust. Triticale (X Triticosecale Wittmack) is a man-made species with a short history of just over 100 years since the first attempt was made to produce an artificial hybrid. Triticale have good agronomic traits such as large head, more grains, resistance to biotic and abiotic stresses. Leaf rolling may be associated with improved grain yield in some drought situations. Identification of different kinds of rolling leaf types provides a theoretical basis and technical support for leaf shape improvement and also for crop drought tolerance.
277 RILs F6 developed from Avocet-YrA/Chapio were used to genetic analysis and QTL mapping with Inclusive Composite Interval Mapping Method (IciMapping, ICIM, V3.0)according to rust severity at adult plant stage at 4 environments. At same time, Rolling leaf material CMH83 from CIMMYT was studied by phenotypic, cytological and physiological analysis. The results are as follow:
1. Chapio were controlled by four pairs incomplete dominance adult-plant resistance (APR) genes against stripe rust
Chapio was susceptible to stripe rust race (CYR32) at seedling stage, and high resistance at adult-plant resistance(APR), all of 277 RILs F6 developed from Avocet-YrA/Chapio were susceptible to stripe rust race (CYR32), and segregated at adult plant stage. Correlation analysis showed that correlation coefficients were significant among 4 environments(P<0.001). Frequency distribution was a littler skew distribution which suggested that Chapio probably contained larger effect gene for APR. Stripe rust severity in some lines showed different value between CIMMYT and China, particularly for 22 lines which difference reached 40-78.3%. It showed that one or several genes in Chapio can identify the races between Chinese race CYR32 and Mexican race YRMV09.
Genetic analysis suggested that Chapio contained 4 APR genes by Chi-squared analyses (P>0.05). Rust severity were 40-50% in Avocet-TrA x Chapio F1 and Chapio x Avocet-YrA F1. Therefore, those result suggested that there were four pairs incomplete dominance adult-plant resistance(APR) genes against stripe rust in CIMMYT variety Chapio.
2. Four additive APR QTLs and one pair epistatic interactions QTL had been detected, QYr.saas-3DL should be a new QTL
Four QTLs for APR against stripe rust have been detected by Inclusive Composite Interval Mapping Method on chromosomes 3BS,3DL,5BL and 7DS in 4 environments, designated QYr.saas-3BS, QYr,saas-3DL, QYr.saas-5BL and QYr.saas-7DS, respectively, explaining from 2.64 to 48.93% of the phenotypic variance. All QTLs were from Chapio. The QTL QYr.saas-3DL, different from all QTLs reported previously, was flanked by SSR markers Xgdm8 and GDM128, with a genetic distance of 10 cM, which accounted for 3.84-12.44% of phenotypic variance. The QTL QYr.caas-7DS flanked by markers csLV34 and Xgwm295, which were closely linked to QYr.caas-7DS which showed the largest effect for resistance to stripe rust and accounted for 28.18-48.93% of phenotypic variance. The QTLs QYr.saas-3BS and QYr.saas-5BL flanked by markers Xbarc147 and Xwms533, Xgwm46 and Xbarc267, with a genetic distance of 2.0 cM and 38 cM, which accounted for 7.30-17.00% and 2.64-4.76% of phenotypic variance, respectively. QYr.saas-3DL should be a new QTL, others needed to further confirm. One pair epistatic interactions QTLs (QYr.saas-3BS x QYr.saas-7DS) had been detected, which accounted for 2.59-7.79% of the phenotypic variance in 3 environments and average value. QTLs QYr.saas-3BS, QYr.saas-3DL, and QYr.saas-7DS could be used in marker-assisted selection for APR to striper rust in wheat breeding programs.
3. Leaf rolling occurs in some cereal genotypes in response to drought. In the present study, we identified a rolling leaf genotype CMH83 of hexaploid Triticale (X Triticosecale Wittmack), that exhibited reduced plant height, rolling and narrow leaves, by phenotypic, cytological and physiological analysis. Gliadin electrophoresis of seeds protein showed that CMH83 was genetically stable. The sequential Giemsa-C banding and genomic in situ hybridization showed that the CMH83 contained 12 rye chromosomes, of which two pairs of chromosomes had reduced telomeric heterochromatin bands. The test of leaf relative water content (RWC) and water loss rate (WLR) of CMH83 compared with those of wheat cultivars indicated that rapid water loss after drought stress in CMH83 could be associated with the leaf rolling phenotypes. Leaf rolling phenotype of CMH83 was a dominant trait in our preliminary inheritance studies. The triticale line CMH83 can be used to study the mechanism of triticale for drought resistances.
引文
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