小偃麦渗入系抗条锈基因分子标记与遗传定位
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摘要
小麦条锈病是由小麦条锈菌(Puccinia striiformis f. sp. tritici)引起的气传性真菌性病害,遍布于世界各大麦区,特别是高海拨的冷凉地区,是影响全世界小麦生产的三大重要病害之一。利用抗病基因历来是控制该病害的主要手段,但抗病性丧失是长期以来未能解决的大难题。科学表明,品种或抗源单一化而引起的毒性单化是造成抗病性丧失的重要原因。迄今国际上虽已正式命名了50多个抗条锈病基因(Yr1-Yr53),但除了Yr5、Yr10、Yr15等少数基因对我国目前流行的优势小种CYR32、 CYR33仍具有抗性外,绝大部分已成为无效基因。
从近缘植物种属导入新的抗病基因是实现抗源多样化的有效途径。中间偃麦草(Thinopyrum intermedium,2n=6x=42, JJsS)和彭提卡偃麦草(Th. ponticun,2n=10x=70, JJJJsJs)免疫或高抗条锈、白粉等多种小麦真菌病害,是普通小麦(Triticum aestivumL.)遗传改良的重要基因库。多年来,本实验室以八倍体小偃麦为桥梁,通过‘六×八’式杂交、回交,将中间偃麦草和彭提卡偃麦草的抗性基因逐步导入普通小麦,育成了一系列稳定高抗小麦条锈、白粉的小麦-异源渗入系。本研究中则采用遗传分析、细胞遗传学鉴定和SSR分子标记技术,分别对其中几个渗入系材料进行了抗条锈性遗传分析及其抗性基因的分子定位,旨在发掘新的抗条锈病基因,拓宽小麦抗条锈育种资源。其主要结果如下：
1、CH223是八倍体小偃麦新类型——TAI7047的一个渗入系。在苗期对其进行多小种抗性鉴定,结果表明CH223免疫CYR32、CYR33、v26等9个条锈菌生理小种,其抗性来自中间偃麦草。基因组原位杂交和染色体配对分析结果表明,CH223具有完整的21对染色体,且观察不到可见的外源DNA杂交信号,说明CH223是一个源于中间偃麦草的隐形异源渐渗系。将CH223与感病品种(系)‘台长29’和‘SY95-71’杂交,其F2、BC1代和F2：3家系的抗、感分离比均符合3：1、1：1和1：2：1,证明CH223对条锈菌系CYR32的抗性受1对显性基因控制,暂命名为YrCH223。利用211个来自台长29/CH223的F2代单株构建作图群体,发现5个与抗病基因连锁的多态性SSR标记,位置顺序为：Xgwm540-Xbarc1096-YrCH223-Xwmc47-Xwmc310-Xgpw7272,其遗传距离分别为21.9cM、8.0cM、7.2cM、12.5cM和11.3cM。最终根据小麦SSR遗传图谱和连锁标记在中国春缺体-四体、双端体的扩增结果,将该抗条锈病基因定位于4BL染色体上。由于这是第一个定位于小麦4BL的抗条锈病基因,因此国际小麦基因命名委员会正式将其定名为Yr50。
2、小麦-彭提卡偃麦草渗入系CH7102衍生于八倍体小偃麦‘小偃7430’。苗期抗性鉴定表明,CH7102免疫我国目前的优势小种CYR32、CYR33,其抗性反应型与‘小偃7430’及其野生亲本彭提卡偃麦草的抗性表现一致,而系谱中的所有小麦亲本均感病,因而推断CH7102的抗性来源为彭提卡偃麦草。对CH7102与感病小麦杂交的后代进行成株期抗性遗传分析鉴定,发现其对条锈小种CYR32的抗性受1对显性基因控制,暂命名为YrCH7102。随后使用集群分离分析法(BSA法)和SSR标记相结合,筛选到5个与抗性基因连锁的SSR标记,位置顺序为：Xbarcl24-Xgwm636-YrCH7102-Xgpw2204-Xgwm95-Xgwm296,遗传距离分别为5.0cM、8.6cM、8.4cM、2.7cM和14.4cM。根据小麦SSR遗传连锁图及利用中国春第2同源群缺体-四体、双端体材料对SSR标记的定位结果,推断YrCH7102位于2AS上。
3、CH7115是衍生于八倍体小偃麦‘小偃7430’并免疫当前优势条锈菌小种的另一个小偃麦渗入系。对CH7115的抗性鉴定和遗传分析结果表明,CH7115苗期免疫或近免疫流行条锈菌小种CYR32和CYR33,其抗性与抗性供体‘小偃7430’及野生供体彭提卡偃麦草相似。‘CH7115×台长29’杂交后代的成株期抗性鉴定显示,CH7115对条锈菌小种CYR32呈现1对显性基因的控制模式,暂命名为YrCH7115。随后将CH7115×台长29的F2代群体采用BSA法结合SSR标记技术进行分析,发现5个小麦SSR标记与抗病基因连锁,位置顺序为:Xgdm33-YrCH7115-Xgwm11-Xcfd65-Xgwm18-Xbarc137,遗传距离分别为10.5cM.7.1cM.0.4cM.2.7cM和1.2cM。通过中国春缺体-四体、双端体材料的进一步鉴定,这5个与抗病基因连锁的SSR标记被定位于1B染色体短臂上。据此推断YrCH7115位于1BS染色体上。
4、抗条锈病渗入系CH7359衍生于小麦×中间偃麦草的八倍体小偃麦TAI8335。苗期条锈菌多小种鉴定结果揭示,CH7359的条锈病抗性来自中间偃麦草。将CH7359与台长29和绵阳11的F1、F2、BC1、F2:3群体种植于大田并分析其抗感分离比,结果表明CH7359对条锈菌CYR32的抗性受1对显性基因控制,暂命名为YrCH7359。使用BSA法结合SSR标记技术对CH7359×绵阳11的F2群体进行分析,找到3个与抗病基因连锁的SSR标记,位置顺序为：Xgwm273-YrCH7359-Xgwm626-Xbarc24,遗传距离分别为8.6cM、5.9cM和3.2cM。同样应用中国春缺体-四体、双端体材料进行鉴定,这3个与抗条锈基因连锁的标记均位于6BL染色体上,因而最终将这个新基因位于6BL染色体。
5、抗条锈病渗入系CH7203也衍生于八倍体小偃麦TAI8335。苗期经多个条锈菌小种鉴定并结合系谱分析,推断出CH7203的条锈病抗性也来自中间偃麦草。将CH7203与绵阳11的各杂交、回交群体种植于大田并分析其抗感分离比,结果表明CH7203对条锈菌CYR32的抗性也受1对显性基因控制。同样也使用BSA法结合SSR标记技术对CH7203×绵阳11的F2群体进行分析,发现2个小麦SSR标记与抗病基因连锁,位置顺序为：YrCH7203-Xbarc170-Xwmc161,遗传距离分别为4.1cM和6.3cM。两个连锁标记经中国春缺体-四体、双端体材料的鉴定,被定位在4AL染色体上,但由于连锁的标记位于目标基因同侧,所以该抗条锈基因的准确定位还有待更多连锁标记的筛选。
综上所述,本研究以5个小偃麦渐渗系(CH223、CH7102、CH7115、CH7359、 CH7203)为试材,应用遗传学、细胞遗传学和分子标记等技术,详尽分析了这5个小麦抗条锈新品系的抗性遗传机制和抗性位点的染色体定位。通过对这5个品系的F1、F2、BC1和F2：3群体,进行苗期和成株期的抗条锈性鉴定。抗性表型的遗传分析证明这5个品系的抗条锈性分别均由一个显性基因控制,且抗性位点来源于中间偃麦草或彭提卡偃麦草,是一类小麦条锈病的新抗源。以各品系相应的F2为作图群体,应用SSR标记技术分别构建了这5个新抗性位点的分子连锁图谱。进一步应用中国春缺体-四体、双端体材料鉴定与这5个抗性位点紧密连锁分子标记的染色体位置,初步将各自所携带的抗条锈基因分别定位于4BL、2AS、1BS、6BL和4A。其中定位于4BL染色体上的抗条锈基因已被国际命名委员会正式命名为Yr50。这5个新抗条锈位点的定位及分子标记的建立为下一步抗性基因的图位克隆及其功能鉴定奠定了基础。基因组原位杂交结果显示这些新品系都属于隐形异源渐渗系,这种抗性材料的抗性基因易于稳定遗传和导入新品系以培育优良抗性品种。这些新基因的发现不仅有助于小麦抗条锈分子标记聚合育种,而且对于实现抗源多样化及我国小麦条锈病的可持续控制具有重要的理论及应用价值。
Wheat stripe (yellow) rust, caused by Puccinia striiformis f. sp. tritici (Pst), is a major disease that can cause tremendous wheat production losses worldwide, especially in the cooler and wetter environments. Since the appearance of PST race CYR32and CYR33which are the most widely virulent and predominant pathotypes in China, a lot of wheat cultivars have become susceptible, resulting in the epidemic of stripe rust. Frequent changes in the pathogen population and the homology of stripe rust resistance background in commercial wheat cultivars are the main reasons of the epidemic. Therefore, using resistant cultivars is the most economical and safty method to reduce damage caused by this disease.
Alien resistant gene transfer is a valuable means of increasing the amount of resistance diversity. The Thinopyrum species, Th. intermedium (Host)(2n=6x=42, JJSS) and Th. ponticum (Podp.)(2n=10x=70,JJJJSJS) had many excellent resistance gene for wheat disease, it is a valuable source of wheat resistance breeding. At present, more and more stripe rust resistance varieties derived from the two Thinopyrum species have been cultivated and used for wheat breeding. Recently, several new Thinopyrum-derived multi-resistance lines have been developed, and they are highly resistant to stripe rust. In this study, our aims were to determine the inheritance, chromosome location and linkage to molecular markers of these new lines. The main results are as follows:
1. CH223is a Th. intermedium-derived wheat introgression line and resistant to stripe rust. To investigate the inheritance of stripe rust resistance introgressed from Th. intermedium, CH223was crossed to susceptible cultivars Taichung29and SY95-71to yield segregating populations. The F2, F3and BC1were tested for segregation of stripe rust resistance. Genetic analysis showed that resistance of CH223was controlled by a single dominant gene. An F2segregating population from CH223/Taichung29was further used for microsatellite screening and gene mapping. The resistance gene was linked to five co-dominant genomic SSR markers, Xgwm540, Xbarc1096, Xwmc310, Xgpw7272and Xwmc47, their most likely order was Xgwm540-Xbarc1096-YrCH223-Xwmc310-Xgpw7272-Xwmc47at21.9,8.0,7.2,12.5and11.3cM, respectively. Through the Chinese Spring nullisomic-tetrasomic and ditelosomic lines detection, the polymorphic markers and the resistance gene were assigned to chromosome4BL. As no stripe rust resistance gene was previously assigned to chromosome4BL, this new resistance gene was designated Yr50. Yr50, together with the identified closely linked markers, could be useful in marker-assisted selection for pyramiding stripe rust resistance genes.
2. CH7102, the resistance to stripe rust was introgressed into common wheat (Triticum aestivum L.) from Thinopyrum ponticum, using a resistant partial amphiploid as a bridging parent in crosses with susceptible wheat lines. The evaluation of resistance reactions in this study demonstrated that the wheat line CH7102has exhibited a high level of resistance to Chinese Pst races CYR32and CYR33, the most widely virulent and predominant pathotypes in China. The resistance responses of CH7102to the races tested was similar to those of its donor parent Xiaoyan7430as well as the wild parent, whereas all the wheat parents involved were susceptible, indicating that the resistance conferred by CH7102was possibly derived from T. ponticum. The F1,F2, F3and BC1populations from a cross of CH7102with susceptible line were tested for segregation of stripe rust resistance. Genetic analysis all showed that resistance of CH7102was controlled by a single dominant gene. Through microsatellite screening five co-dominant genomic SSR markers were found linking to resistance gene, Xbarcl24, Xgwm636, Xgpw2204, Xgwm95and Xgwm296. Their most likely order was Xbarc124-Xgwm636-YrCH7102-Xgpw2204-Xgwm95-Xgwm296at5.0cM,8.6cM,8.4cM,2.7cM and14.4cM, respectively. Through the nullisomic-tetrasomic and ditelosomic lines detection, the polymorphic markers and the resistance gene were assigned to chromosome2AS.
3.CH7115is another stripe rust resistance gene which was also introgressed from Thinopyrum ponticum. The resistance responses of CH7115to the races tested was similar to those of its donor parent Xiaoyan7430as well as the wild parent, which indicated the resistance of CH7115was derived from Thinopyrum ponticum. The F1, F2, F3and BC1populations from a cross of CH7115with susceptible line were tested for segregation of stripe rust resistance. Genetic analysis all showed that resistance of CH7115was controlled by only one dominant gene.Through microsatellite screening five co-dominant genomic SSR markers were found linking to resistance gene, Xgdm33, Xgwmll, Xcfd65, Xgwm18and Xbarc137. Their most likely order was Xgdm33-YrCH7115-Xgwm11-Xcfd65-Xgwml8-Xbarcl37at10.5cM,7.1cM,0.4cM,2.7cM and1.2cM, respectively. Through the nullisomic-tetrasomic and ditelosomic lines detection, the polymorphic markers and the resistance gene were assigned to chromosome IBS.
4. TAI8335was a new partial amphiploid derived from Thinopyrum intermedium. CH7359was a homogeneous BC2F4-derived wheat lines using TAI8335as its donor parent. Through races test, CH7359showed similar phenotype to its donor parent and the wild parent, which indicated the resistance of Line CH7359was derived from Thinopyrum-intermedium. The F1, F2, F3and BC1populations from a cross of CH7359with susceptible line were tested for segregation of stripe rust resistance. Genetic analysis all showed that resistance of CH7359was controlled by only one dominant gene. After microsatellite screening three co-dominant genomic SSR markers was found linking to resistance gene. Using Mapmaker3.0software to map their linkage, we got their most likely order was Xgwm273-YrCH7359-Xgwm626-Xbarc24at8.6cM,5.9cM and3.2cM, respectively. Finally the polymorphic markers and the resistance gene were assigned to chromosome6BL.
5. CH7203is another stripe rust resistance gene which was also introgressed from Thinopyrum Intermedium, using the resistant partial amphiploid TAI8335as a bridging parent crossed with susceptible wheat lines MianYang11. The resistance response of CH7203to the tested races was similar to those of its donor parent as well as the wild parent, which indicated the resistance of CH7203derived from Thinopyrum Intermedium. Genetic analysis of the F1, F2, F3and BC1populations from the cross of CH7203with susceptible line Mian Yang11showed that resistance of CH7203was controlled by only one dominant gene. Microsatellite screening results showed two co-dominant genomic SSR markers were linked to resistance gene, Xbarc170and Xwmcl61, their most likely order was YrCH7203-Xbarc170-Xwmc161at4.1cM,6.3cM, respectively. Through nullisomic-tetrasomic and ditelosomic lines detection, the polymorphic markers were assigned to chromosome4AL. But the two linking SSR makers were located the same side to resistance gene, the target gene was only assigned to chromosome4A. We need to find more tight linking markers to locate this new resistance gene exactly.
In conclusion, through conventional hybrid analysis, we found stripe rust resistances of five wheat-Thinopyrum germplasm lines (CH223, CH7102, CH7115, CH7359, CH7203) were all controlled by a single dominant gene and the resistance was derived from Th. intermedium or Th. poticum. Resistance genes carried by the five lines were respectively assigned to4BL2AS, IBS,6BL and4A, using SSR molecular marker technique to analyze F2derivations. Both the resource and chromosome of these genes are different from other known stripe rust genes. Among them, a new resistance gene carried by CH223was formally designated Yr50. Identification and analysis of these new stripe rust resistance lines will be beneficial for map-based gene cloning and gene function characterization. Cytological analyses using GISH also detected no chromosomal segments from alien species, so the resistance of these new lines can be stably inherited. It will be available for wheat molecular marker-assisted resistance breeding and increasing the diversity of wheat resistance resource.
从近缘植物种属导入新的抗病基因是实现抗源多样化的有效途径。中间偃麦草(Thinopyrum intermedium,2n=6x=42, JJsS)和彭提卡偃麦草(Th. ponticun,2n=10x=70, JJJJsJs)免疫或高抗条锈、白粉等多种小麦真菌病害,是普通小麦(Triticum aestivumL.)遗传改良的重要基因库。多年来,本实验室以八倍体小偃麦为桥梁,通过‘六×八’式杂交、回交,将中间偃麦草和彭提卡偃麦草的抗性基因逐步导入普通小麦,育成了一系列稳定高抗小麦条锈、白粉的小麦-异源渗入系。本研究中则采用遗传分析、细胞遗传学鉴定和SSR分子标记技术,分别对其中几个渗入系材料进行了抗条锈性遗传分析及其抗性基因的分子定位,旨在发掘新的抗条锈病基因,拓宽小麦抗条锈育种资源。其主要结果如下：
1、CH223是八倍体小偃麦新类型——TAI7047的一个渗入系。在苗期对其进行多小种抗性鉴定,结果表明CH223免疫CYR32、CYR33、v26等9个条锈菌生理小种,其抗性来自中间偃麦草。基因组原位杂交和染色体配对分析结果表明,CH223具有完整的21对染色体,且观察不到可见的外源DNA杂交信号,说明CH223是一个源于中间偃麦草的隐形异源渐渗系。将CH223与感病品种(系)‘台长29’和‘SY95-71’杂交,其F2、BC1代和F2：3家系的抗、感分离比均符合3：1、1：1和1：2：1,证明CH223对条锈菌系CYR32的抗性受1对显性基因控制,暂命名为YrCH223。利用211个来自台长29/CH223的F2代单株构建作图群体,发现5个与抗病基因连锁的多态性SSR标记,位置顺序为：Xgwm540-Xbarc1096-YrCH223-Xwmc47-Xwmc310-Xgpw7272,其遗传距离分别为21.9cM、8.0cM、7.2cM、12.5cM和11.3cM。最终根据小麦SSR遗传图谱和连锁标记在中国春缺体-四体、双端体的扩增结果,将该抗条锈病基因定位于4BL染色体上。由于这是第一个定位于小麦4BL的抗条锈病基因,因此国际小麦基因命名委员会正式将其定名为Yr50。
2、小麦-彭提卡偃麦草渗入系CH7102衍生于八倍体小偃麦‘小偃7430’。苗期抗性鉴定表明,CH7102免疫我国目前的优势小种CYR32、CYR33,其抗性反应型与‘小偃7430’及其野生亲本彭提卡偃麦草的抗性表现一致,而系谱中的所有小麦亲本均感病,因而推断CH7102的抗性来源为彭提卡偃麦草。对CH7102与感病小麦杂交的后代进行成株期抗性遗传分析鉴定,发现其对条锈小种CYR32的抗性受1对显性基因控制,暂命名为YrCH7102。随后使用集群分离分析法(BSA法)和SSR标记相结合,筛选到5个与抗性基因连锁的SSR标记,位置顺序为：Xbarcl24-Xgwm636-YrCH7102-Xgpw2204-Xgwm95-Xgwm296,遗传距离分别为5.0cM、8.6cM、8.4cM、2.7cM和14.4cM。根据小麦SSR遗传连锁图及利用中国春第2同源群缺体-四体、双端体材料对SSR标记的定位结果,推断YrCH7102位于2AS上。
3、CH7115是衍生于八倍体小偃麦‘小偃7430’并免疫当前优势条锈菌小种的另一个小偃麦渗入系。对CH7115的抗性鉴定和遗传分析结果表明,CH7115苗期免疫或近免疫流行条锈菌小种CYR32和CYR33,其抗性与抗性供体‘小偃7430’及野生供体彭提卡偃麦草相似。‘CH7115×台长29’杂交后代的成株期抗性鉴定显示,CH7115对条锈菌小种CYR32呈现1对显性基因的控制模式,暂命名为YrCH7115。随后将CH7115×台长29的F2代群体采用BSA法结合SSR标记技术进行分析,发现5个小麦SSR标记与抗病基因连锁,位置顺序为:Xgdm33-YrCH7115-Xgwm11-Xcfd65-Xgwm18-Xbarc137,遗传距离分别为10.5cM.7.1cM.0.4cM.2.7cM和1.2cM。通过中国春缺体-四体、双端体材料的进一步鉴定,这5个与抗病基因连锁的SSR标记被定位于1B染色体短臂上。据此推断YrCH7115位于1BS染色体上。
4、抗条锈病渗入系CH7359衍生于小麦×中间偃麦草的八倍体小偃麦TAI8335。苗期条锈菌多小种鉴定结果揭示,CH7359的条锈病抗性来自中间偃麦草。将CH7359与台长29和绵阳11的F1、F2、BC1、F2:3群体种植于大田并分析其抗感分离比,结果表明CH7359对条锈菌CYR32的抗性受1对显性基因控制,暂命名为YrCH7359。使用BSA法结合SSR标记技术对CH7359×绵阳11的F2群体进行分析,找到3个与抗病基因连锁的SSR标记,位置顺序为：Xgwm273-YrCH7359-Xgwm626-Xbarc24,遗传距离分别为8.6cM、5.9cM和3.2cM。同样应用中国春缺体-四体、双端体材料进行鉴定,这3个与抗条锈基因连锁的标记均位于6BL染色体上,因而最终将这个新基因位于6BL染色体。
5、抗条锈病渗入系CH7203也衍生于八倍体小偃麦TAI8335。苗期经多个条锈菌小种鉴定并结合系谱分析,推断出CH7203的条锈病抗性也来自中间偃麦草。将CH7203与绵阳11的各杂交、回交群体种植于大田并分析其抗感分离比,结果表明CH7203对条锈菌CYR32的抗性也受1对显性基因控制。同样也使用BSA法结合SSR标记技术对CH7203×绵阳11的F2群体进行分析,发现2个小麦SSR标记与抗病基因连锁,位置顺序为：YrCH7203-Xbarc170-Xwmc161,遗传距离分别为4.1cM和6.3cM。两个连锁标记经中国春缺体-四体、双端体材料的鉴定,被定位在4AL染色体上,但由于连锁的标记位于目标基因同侧,所以该抗条锈基因的准确定位还有待更多连锁标记的筛选。
综上所述,本研究以5个小偃麦渐渗系(CH223、CH7102、CH7115、CH7359、 CH7203)为试材,应用遗传学、细胞遗传学和分子标记等技术,详尽分析了这5个小麦抗条锈新品系的抗性遗传机制和抗性位点的染色体定位。通过对这5个品系的F1、F2、BC1和F2：3群体,进行苗期和成株期的抗条锈性鉴定。抗性表型的遗传分析证明这5个品系的抗条锈性分别均由一个显性基因控制,且抗性位点来源于中间偃麦草或彭提卡偃麦草,是一类小麦条锈病的新抗源。以各品系相应的F2为作图群体,应用SSR标记技术分别构建了这5个新抗性位点的分子连锁图谱。进一步应用中国春缺体-四体、双端体材料鉴定与这5个抗性位点紧密连锁分子标记的染色体位置,初步将各自所携带的抗条锈基因分别定位于4BL、2AS、1BS、6BL和4A。其中定位于4BL染色体上的抗条锈基因已被国际命名委员会正式命名为Yr50。这5个新抗条锈位点的定位及分子标记的建立为下一步抗性基因的图位克隆及其功能鉴定奠定了基础。基因组原位杂交结果显示这些新品系都属于隐形异源渐渗系,这种抗性材料的抗性基因易于稳定遗传和导入新品系以培育优良抗性品种。这些新基因的发现不仅有助于小麦抗条锈分子标记聚合育种,而且对于实现抗源多样化及我国小麦条锈病的可持续控制具有重要的理论及应用价值。
Wheat stripe (yellow) rust, caused by Puccinia striiformis f. sp. tritici (Pst), is a major disease that can cause tremendous wheat production losses worldwide, especially in the cooler and wetter environments. Since the appearance of PST race CYR32and CYR33which are the most widely virulent and predominant pathotypes in China, a lot of wheat cultivars have become susceptible, resulting in the epidemic of stripe rust. Frequent changes in the pathogen population and the homology of stripe rust resistance background in commercial wheat cultivars are the main reasons of the epidemic. Therefore, using resistant cultivars is the most economical and safty method to reduce damage caused by this disease.
Alien resistant gene transfer is a valuable means of increasing the amount of resistance diversity. The Thinopyrum species, Th. intermedium (Host)(2n=6x=42, JJSS) and Th. ponticum (Podp.)(2n=10x=70,JJJJSJS) had many excellent resistance gene for wheat disease, it is a valuable source of wheat resistance breeding. At present, more and more stripe rust resistance varieties derived from the two Thinopyrum species have been cultivated and used for wheat breeding. Recently, several new Thinopyrum-derived multi-resistance lines have been developed, and they are highly resistant to stripe rust. In this study, our aims were to determine the inheritance, chromosome location and linkage to molecular markers of these new lines. The main results are as follows:
1. CH223is a Th. intermedium-derived wheat introgression line and resistant to stripe rust. To investigate the inheritance of stripe rust resistance introgressed from Th. intermedium, CH223was crossed to susceptible cultivars Taichung29and SY95-71to yield segregating populations. The F2, F3and BC1were tested for segregation of stripe rust resistance. Genetic analysis showed that resistance of CH223was controlled by a single dominant gene. An F2segregating population from CH223/Taichung29was further used for microsatellite screening and gene mapping. The resistance gene was linked to five co-dominant genomic SSR markers, Xgwm540, Xbarc1096, Xwmc310, Xgpw7272and Xwmc47, their most likely order was Xgwm540-Xbarc1096-YrCH223-Xwmc310-Xgpw7272-Xwmc47at21.9,8.0,7.2,12.5and11.3cM, respectively. Through the Chinese Spring nullisomic-tetrasomic and ditelosomic lines detection, the polymorphic markers and the resistance gene were assigned to chromosome4BL. As no stripe rust resistance gene was previously assigned to chromosome4BL, this new resistance gene was designated Yr50. Yr50, together with the identified closely linked markers, could be useful in marker-assisted selection for pyramiding stripe rust resistance genes.
2. CH7102, the resistance to stripe rust was introgressed into common wheat (Triticum aestivum L.) from Thinopyrum ponticum, using a resistant partial amphiploid as a bridging parent in crosses with susceptible wheat lines. The evaluation of resistance reactions in this study demonstrated that the wheat line CH7102has exhibited a high level of resistance to Chinese Pst races CYR32and CYR33, the most widely virulent and predominant pathotypes in China. The resistance responses of CH7102to the races tested was similar to those of its donor parent Xiaoyan7430as well as the wild parent, whereas all the wheat parents involved were susceptible, indicating that the resistance conferred by CH7102was possibly derived from T. ponticum. The F1,F2, F3and BC1populations from a cross of CH7102with susceptible line were tested for segregation of stripe rust resistance. Genetic analysis all showed that resistance of CH7102was controlled by a single dominant gene. Through microsatellite screening five co-dominant genomic SSR markers were found linking to resistance gene, Xbarcl24, Xgwm636, Xgpw2204, Xgwm95and Xgwm296. Their most likely order was Xbarc124-Xgwm636-YrCH7102-Xgpw2204-Xgwm95-Xgwm296at5.0cM,8.6cM,8.4cM,2.7cM and14.4cM, respectively. Through the nullisomic-tetrasomic and ditelosomic lines detection, the polymorphic markers and the resistance gene were assigned to chromosome2AS.
3.CH7115is another stripe rust resistance gene which was also introgressed from Thinopyrum ponticum. The resistance responses of CH7115to the races tested was similar to those of its donor parent Xiaoyan7430as well as the wild parent, which indicated the resistance of CH7115was derived from Thinopyrum ponticum. The F1, F2, F3and BC1populations from a cross of CH7115with susceptible line were tested for segregation of stripe rust resistance. Genetic analysis all showed that resistance of CH7115was controlled by only one dominant gene.Through microsatellite screening five co-dominant genomic SSR markers were found linking to resistance gene, Xgdm33, Xgwmll, Xcfd65, Xgwm18and Xbarc137. Their most likely order was Xgdm33-YrCH7115-Xgwm11-Xcfd65-Xgwml8-Xbarcl37at10.5cM,7.1cM,0.4cM,2.7cM and1.2cM, respectively. Through the nullisomic-tetrasomic and ditelosomic lines detection, the polymorphic markers and the resistance gene were assigned to chromosome IBS.
4. TAI8335was a new partial amphiploid derived from Thinopyrum intermedium. CH7359was a homogeneous BC2F4-derived wheat lines using TAI8335as its donor parent. Through races test, CH7359showed similar phenotype to its donor parent and the wild parent, which indicated the resistance of Line CH7359was derived from Thinopyrum-intermedium. The F1, F2, F3and BC1populations from a cross of CH7359with susceptible line were tested for segregation of stripe rust resistance. Genetic analysis all showed that resistance of CH7359was controlled by only one dominant gene. After microsatellite screening three co-dominant genomic SSR markers was found linking to resistance gene. Using Mapmaker3.0software to map their linkage, we got their most likely order was Xgwm273-YrCH7359-Xgwm626-Xbarc24at8.6cM,5.9cM and3.2cM, respectively. Finally the polymorphic markers and the resistance gene were assigned to chromosome6BL.
5. CH7203is another stripe rust resistance gene which was also introgressed from Thinopyrum Intermedium, using the resistant partial amphiploid TAI8335as a bridging parent crossed with susceptible wheat lines MianYang11. The resistance response of CH7203to the tested races was similar to those of its donor parent as well as the wild parent, which indicated the resistance of CH7203derived from Thinopyrum Intermedium. Genetic analysis of the F1, F2, F3and BC1populations from the cross of CH7203with susceptible line Mian Yang11showed that resistance of CH7203was controlled by only one dominant gene. Microsatellite screening results showed two co-dominant genomic SSR markers were linked to resistance gene, Xbarc170and Xwmcl61, their most likely order was YrCH7203-Xbarc170-Xwmc161at4.1cM,6.3cM, respectively. Through nullisomic-tetrasomic and ditelosomic lines detection, the polymorphic markers were assigned to chromosome4AL. But the two linking SSR makers were located the same side to resistance gene, the target gene was only assigned to chromosome4A. We need to find more tight linking markers to locate this new resistance gene exactly.
In conclusion, through conventional hybrid analysis, we found stripe rust resistances of five wheat-Thinopyrum germplasm lines (CH223, CH7102, CH7115, CH7359, CH7203) were all controlled by a single dominant gene and the resistance was derived from Th. intermedium or Th. poticum. Resistance genes carried by the five lines were respectively assigned to4BL2AS, IBS,6BL and4A, using SSR molecular marker technique to analyze F2derivations. Both the resource and chromosome of these genes are different from other known stripe rust genes. Among them, a new resistance gene carried by CH223was formally designated Yr50. Identification and analysis of these new stripe rust resistance lines will be beneficial for map-based gene cloning and gene function characterization. Cytological analyses using GISH also detected no chromosomal segments from alien species, so the resistance of these new lines can be stably inherited. It will be available for wheat molecular marker-assisted resistance breeding and increasing the diversity of wheat resistance resource.
引文
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