小麦—冰草杂交衍生系中优异基因的染色体定位与分子标记
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摘要
小麦野生近缘植物是小麦遗传改良的巨大基因库，如何从中发掘、转移并利用生产急需的抗逆、抗病和高产基因，既有重大的经济价值，又有长远的理论意义。冰草属(Agropyron Gaertn．，P Genome)植物具有普通小麦所没有的多种抗逆、抗病和优良产量结构基因。本研究室首次突破小麦-冰草远缘杂交，并进行了多年自交、回交，应用细胞学、原位杂交和SSR技术，获得了小麦-冰草异源二体附加系和部分易位系。本研究的任务是在此基础上，对冰草P染色体上携带的有益基因进行较系统的鉴定，明确其在小麦背景下的表达及其染色体定位；同时，通过构建抗白粉病小麦-冰草易位系(无蜡)与感病小麦品种(温麦6号)杂交F1代的DH群体，用SSR技术对来自冰草的抗白粉病基因进行遗传定位和分子标记。其主要结果如下：
1．采用小麦白粉病菌4个优势生理小种，对整套小麦-冰草二体附加系的苗期和成株期抗性鉴定表明，冰草的2P、7P染色体具有抗白粉病基因。此外，或许由于P基因组内染色体互换的结果，个别的3P和6P附加株系亦存在抗性基因，这些抗病基因在普通小麦基因组背景下能够充分表达。
2．用条锈病菌混合生理小种进行的田间诱发鉴定和生长箱控温鉴定表明，冰草6P染色体上具有抗条锈病基因，在小麦背景下表现中等抗性；用叶锈病菌混合生理小种在生长箱进行的鉴定表明，冰草2P、3P、7P染色体上具有对叶锈病免疫的基因，因此，冰草可作为普通小麦抗叶锈病的重要基因源。
3．利用反复干旱法对整套小麦-冰草二体附加系抗旱性的鉴定表明，冰草的抗旱性可能涉及较多的基因位点，单个附加系的抗旱性均弱于冰草，抗旱基因主要分布在2P、4P、6P和7P染色体上；对小麦-冰草二体附加系抗寒性的鉴定表明，多基因控制的抗寒基因主要分布于冰草的4P和6P染色体上。
4．对来源于冰草的重要农艺性状基因鉴定结果表明，以苗期植株匍匐或半匍匐、叶色深绿为标志的冬性或春化作用相关基因和叶片边缘卷曲或微卷的标记基因存在于2P、3P和5P染色体上；冰草的强分蘖基因位于2P和6P染色体上；冰草的以穗下颈长(占到其株高的1／2～2／3)，茎秆基部节间短，矮秆为特征的抗倒伏基因位于5P染色体上，并且与产量关系密切的多小穗数、多小花数、大穗基因亦存在于5P染色体；同时，5P也携有晚熟基因。此外，冰草的穗部无芒特征，主要与3P和6P染色体有关。基的电泳分析表明，在冰草Z559种子中未发现高分子谷蛋白亚基基因表达，而普通小麦亲本Fukuho具有2.2+12(D基因组)、2*(A基因组)和7+8(B基因组)5个HMWG亚基。二体附加系的谱带显示，冰草P染色体的存在能够不同程度抑制小麦D基因组编码的2.2和12亚基的表达。例如，只有一个2P附加株系显示了2.2+12亚基，3个6P附加株系仅显示了12亚基，其余附加系中2.2和12亚基完全不能表达。
6．对冰草及其小麦-冰草二体附加系种子中醇溶蛋白(Gli)亚基的电泳分析结果：冰草Z559种子中具有丰富的ω区和α、β区亚基，这些亚基与两个强筋小麦Marquis和McGuire有相似的带型。Z559包含了此二品种几乎所有的ω-亚基，以及α、β区的两个特异性亚基，冰草携带的ω-亚基、α和β亚基基因有可能作为小麦品质改良的基因源。但是16个小麦-冰草二体附加系中ω-亚基数少，并且与普通小麦亲本Fukuho相同，无冰草的ω-亚基特征。由于小麦醇溶蛋白的ω-亚基受第一同源群短臂上的基因位点控制，本研究中仅有第一同源群SSR标记的附加系Ⅱ-5-1中未见冰草的ω-亚基，所以说明有两种可能：尚未获得1P附加系或者冰草的ω-亚基基因在小麦背景下不能够表达。小麦的α、β醇溶蛋白基因位于第六同源群短臂，所有4个6P附加系在α和β区均有两条高强度的特异带，与冰草和两个强筋小麦相同，它们可能与好的品质有关，同时能够作为6P染色体的生化标记。此外在两个小麦-冰草衍生系无蜡和L012中发现ω和γ区亚基发生了变化。
7．小麦-冰草衍生系无蜡从Fukuho与冰草Z559杂交BC_3自交后代系选而来，对自粉病免疫。对其抗病基因推导结果，除了对E21表现中抗，对其余白粉病生理小种均为免疫。用集群分离法(BSA)对无蜡与感病亲本温麦6号杂交DH群体抗白粉病性的分子标记结果表明，DH群体中抗病与感病株的比例为1：1，抗病基因为单显性基因；获得抗病基因的3个SSR标记，将其定位在3D染色体的长臂上。抗病基因位于Xgwm341和Xgwm383之间，分别距离29.4 cM和26.6 cM，而另一个标记Xgwm645位于Xgwm341外侧26.4 cM处。根据以上结果，这一抗白粉病基因来源于冰草，已定位于普通小麦与冰草衍生系的3D染色体长臂，目前在3DL上尚未有定位的抗白粉病基因，因此认为该基因为新的抗白粉病基因，暂命名为Pm-Ac。
文章对来自冰草的这些重要的抗逆、抗病和丰产结构基因在小麦遗传改良中的可能应用进行了讨论，提出了建议和设想。
The wild relatives of common wheat (Triticum aestivum L.) are a potential gene reservoir for wheat improvement. It's very important economically and theoritically understanding that how to earthout, transfer and utilize some of desirable genes such as genes for biostress and abiostress resistance from the wild relatives of wheat. Agropyron cristatum Gaertn (P Genome) has been found to possess many sort of desirable traits distinguished by their high level of drought and cold tolerance, disease resistance and better plant-shape benefit to high yield, that are potentially valuable for wheat improvement.
Based on the breakthrough of hybridization between common wheat (Fukuho) to A. cristatum (Z559) and following the F1 plant selfing and backcross to Fukuho, Many wheat-wheatgrass alien disomic addition lines and derivatives were produced by cytogenetical and Fluorescence In Situ Hybridization (FISH) identified methods. In this experiment, there is a systematical design to understand all probably desirable genes on each chromosome of P genome under typical test conditions with a full set of wheat-A. cristatum alien disomic addition lines. Meanwhile, an excellent powdery mildew resistance gene originated in A. cristatum (Z559) was mapped to wheat homologous group with SSR markers throughout construction and molecular marker analysis of a DH population of F1 hybrid between Wula, an A. cristatum derivative holding resistance gene, to Wenmai No. 6, a common wheat variety susceptible to powdery mildew. A new powdery mildew resistance gene was named combining data for response pattern and genetic location. The main successful results were shown as followings:With four Erysiphe graminis isolates, E09, E17, E21 and E26 presently each with low, moderate and high virulence to cropping wheat cultivars in China,This full et whest-A.cristatum disomic addition lines were used to artifical inoculation at adult-plant stage, and seedling stage by leaf segment identification method to evaluate their resistance response. It's shown that the powdery mildew resistance genes locate at 2P and 7P chromosomes, and maybe appear in 3P or 6P chromosome. All these resistant genes could express fine under common wheat genome background.
Yellow rust and leaf rust resistance identification were engaged in low-temperature growth chamber and field with mixture spore inoculated of 4 yellow rust isolates and 8 leaf rust isolates, respectively. The results indicated the gene resistant to yellow rust was located on 6P chromosome, with moderate degree resistance in wheat background. But genes for leaf rust resistance with immunity response were pointed on 3P, 7P and 2P chromosome. These genes could sufficiently express in wheat background, will being important gene reservoir in wheat breeding for leaf rust resistance.
The drought resistance identification of the full et of disomic addition lines with the method of "repeated drought periods" indicated that genes for drought resistance mainly scatter in the 2P, 4P, and 7P chromosomes. The resistant strength of each disomic addition line was lower than the A. cristatum parent. The genes for cold tolerance were located on 4P and 6P chromosomes. This cold tolerance response was weaker than the parent A. cristatum.
Identification of genes for some of the important agronomic traits originated in A.cristatum shown that: genes related to vernalization marked with creeping stem, curl or semi-curl and dark green leaf in seedling stage were located on 2P, 3P and 5P chromosomes; genes controlling tiller number were being 2P and 6P, and genes for potential high yield characterized by many spikelet number, many floscular number, longer peduncle length, and shorter plant height were located on 5P chromosomes but with an undesirable late maturity gene. Additionally, gene(s) for awnlessness appeared on 3P, 6P chromosomes.
By electrophoresis gel analysis of HMW glutenin subunits in seed of a set of wheat-A.cristatum addition lines and A.cristatum, it's shown that there are genes in A. cristatum (Z559) inhibiting the expression of HMW glutenin subunits under wheat background. The HMW glutenin genes of A. cristatum (Z559) no express, the female Fukuho possesses 5 subunits: 2．2+12 (encoded by D genome), 2* (A genome) and 7+8 (B genome), but the being of P chromosome had alternative inhibition effect to expression of 2．2 and 12 subunits encoded by D genome in different wheat-A.cristatum alien addition lines.
The expression pattern of genes for gliadin subunits of A. cristatum < L. > Gaertn (Z559) was identified. There were richω-gliadin subunits in Z559 but disappeared all these subunits in the 16 wheat-A. cristatum alien addition lines including a addition line with 1P marker. It's already clear thatω-gliadin is controlled by gene family of the first homologous group, so, perhaps there are two kind of situations: default 1P addition line or disable expression ofω-gliadin genes under wheat background. Two specificα-andβ-gliadin subunits were taken out in four 6P addition lines and Z559 resemble to two strong gluten wheat varieties, Marquis and McGuire. It's indicated that theseαandβ-gliadin genes could be not only resources of quality improvement for wheat but 锘縜lso biochemical markers of 6P chromosome. Otherwises, changes of蠅and纬-gliadin subunits was discovered in wheat-A. cristatum derivatives Wula and Lo12．鈽匒 new powdery mildew gene originated in A. cristatum was named based on data of molecular mapping and response pattern analysis. Wula, a wheat-A.cristatum derivative, selected from BC_2 of hybrid Fukuho脳Z559 by pedigree possesses gene immune to powdery mildew. Response pattern analysis indicated that this gene immune to almost all the powdery mildew isolates except to E21 with middle resistance. By establishing DH population of Wula脳Wenmai No.6(susceptible), genetic location and molecular mapping of the powdery mildw gene were carried out. The response pattern of DH families showed 1:1 ratio of resistance to susceptible individuals indicates that the resistance controlled by a single dominant gene. Bulked segregant analysis was used to identify microsatellite markers linked to the gene. Three SSR markers Xgwm341, Xgwm383 and Xgwm645 were mapped in coupling phase to the gene locus, thus, this resistance gene was located on long arm of 3D chromosome. The gene was mapped between Xgwm341 to Xgwm383 with 29．4cM and 26．6cM respectively. And Xgwm645 was mapped 26．4 cM out of Xgwm341．Because there hasn't been powdery mildew gene was located on 3DL until now, so, this is a new powdery mildew resistance gene originated in A. cristatum. It is designated Pm31 tentatively. The paper discussed the propably utilization of these important gene resources, given the suggestions and the perspects in the future.
1．采用小麦白粉病菌4个优势生理小种，对整套小麦-冰草二体附加系的苗期和成株期抗性鉴定表明，冰草的2P、7P染色体具有抗白粉病基因。此外，或许由于P基因组内染色体互换的结果，个别的3P和6P附加株系亦存在抗性基因，这些抗病基因在普通小麦基因组背景下能够充分表达。
2．用条锈病菌混合生理小种进行的田间诱发鉴定和生长箱控温鉴定表明，冰草6P染色体上具有抗条锈病基因，在小麦背景下表现中等抗性；用叶锈病菌混合生理小种在生长箱进行的鉴定表明，冰草2P、3P、7P染色体上具有对叶锈病免疫的基因，因此，冰草可作为普通小麦抗叶锈病的重要基因源。
3．利用反复干旱法对整套小麦-冰草二体附加系抗旱性的鉴定表明，冰草的抗旱性可能涉及较多的基因位点，单个附加系的抗旱性均弱于冰草，抗旱基因主要分布在2P、4P、6P和7P染色体上；对小麦-冰草二体附加系抗寒性的鉴定表明，多基因控制的抗寒基因主要分布于冰草的4P和6P染色体上。
4．对来源于冰草的重要农艺性状基因鉴定结果表明，以苗期植株匍匐或半匍匐、叶色深绿为标志的冬性或春化作用相关基因和叶片边缘卷曲或微卷的标记基因存在于2P、3P和5P染色体上；冰草的强分蘖基因位于2P和6P染色体上；冰草的以穗下颈长(占到其株高的1／2～2／3)，茎秆基部节间短，矮秆为特征的抗倒伏基因位于5P染色体上，并且与产量关系密切的多小穗数、多小花数、大穗基因亦存在于5P染色体；同时，5P也携有晚熟基因。此外，冰草的穗部无芒特征，主要与3P和6P染色体有关。基的电泳分析表明，在冰草Z559种子中未发现高分子谷蛋白亚基基因表达，而普通小麦亲本Fukuho具有2.2+12(D基因组)、2*(A基因组)和7+8(B基因组)5个HMWG亚基。二体附加系的谱带显示，冰草P染色体的存在能够不同程度抑制小麦D基因组编码的2.2和12亚基的表达。例如，只有一个2P附加株系显示了2.2+12亚基，3个6P附加株系仅显示了12亚基，其余附加系中2.2和12亚基完全不能表达。
6．对冰草及其小麦-冰草二体附加系种子中醇溶蛋白(Gli)亚基的电泳分析结果：冰草Z559种子中具有丰富的ω区和α、β区亚基，这些亚基与两个强筋小麦Marquis和McGuire有相似的带型。Z559包含了此二品种几乎所有的ω-亚基，以及α、β区的两个特异性亚基，冰草携带的ω-亚基、α和β亚基基因有可能作为小麦品质改良的基因源。但是16个小麦-冰草二体附加系中ω-亚基数少，并且与普通小麦亲本Fukuho相同，无冰草的ω-亚基特征。由于小麦醇溶蛋白的ω-亚基受第一同源群短臂上的基因位点控制，本研究中仅有第一同源群SSR标记的附加系Ⅱ-5-1中未见冰草的ω-亚基，所以说明有两种可能：尚未获得1P附加系或者冰草的ω-亚基基因在小麦背景下不能够表达。小麦的α、β醇溶蛋白基因位于第六同源群短臂，所有4个6P附加系在α和β区均有两条高强度的特异带，与冰草和两个强筋小麦相同，它们可能与好的品质有关，同时能够作为6P染色体的生化标记。此外在两个小麦-冰草衍生系无蜡和L012中发现ω和γ区亚基发生了变化。
7．小麦-冰草衍生系无蜡从Fukuho与冰草Z559杂交BC_3自交后代系选而来，对自粉病免疫。对其抗病基因推导结果，除了对E21表现中抗，对其余白粉病生理小种均为免疫。用集群分离法(BSA)对无蜡与感病亲本温麦6号杂交DH群体抗白粉病性的分子标记结果表明，DH群体中抗病与感病株的比例为1：1，抗病基因为单显性基因；获得抗病基因的3个SSR标记，将其定位在3D染色体的长臂上。抗病基因位于Xgwm341和Xgwm383之间，分别距离29.4 cM和26.6 cM，而另一个标记Xgwm645位于Xgwm341外侧26.4 cM处。根据以上结果，这一抗白粉病基因来源于冰草，已定位于普通小麦与冰草衍生系的3D染色体长臂，目前在3DL上尚未有定位的抗白粉病基因，因此认为该基因为新的抗白粉病基因，暂命名为Pm-Ac。
文章对来自冰草的这些重要的抗逆、抗病和丰产结构基因在小麦遗传改良中的可能应用进行了讨论，提出了建议和设想。
The wild relatives of common wheat (Triticum aestivum L.) are a potential gene reservoir for wheat improvement. It's very important economically and theoritically understanding that how to earthout, transfer and utilize some of desirable genes such as genes for biostress and abiostress resistance from the wild relatives of wheat. Agropyron cristatum
Based on the breakthrough of hybridization between common wheat (Fukuho) to A. cristatum (Z559) and following the F1 plant selfing and backcross to Fukuho, Many wheat-wheatgrass alien disomic addition lines and derivatives were produced by cytogenetical and Fluorescence In Situ Hybridization (FISH) identified methods. In this experiment, there is a systematical design to understand all probably desirable genes on each chromosome of P genome under typical test conditions with a full set of wheat-A. cristatum alien disomic addition lines. Meanwhile, an excellent powdery mildew resistance gene originated in A. cristatum (Z559) was mapped to wheat homologous group with SSR markers throughout construction and molecular marker analysis of a DH population of F1 hybrid between Wula, an A. cristatum derivative holding resistance gene, to Wenmai No. 6, a common wheat variety susceptible to powdery mildew. A new powdery mildew resistance gene was named combining data for response pattern and genetic location. The main successful results were shown as followings:With four Erysiphe graminis isolates, E09, E17, E21 and E26 presently each with low, moderate and high virulence to cropping wheat cultivars in China,This full et whest-A.cristatum disomic addition lines were used to artifical inoculation at adult-plant stage, and seedling stage by leaf segment identification method to evaluate their resistance response. It's shown that the powdery mildew resistance genes locate at 2P and 7P chromosomes, and maybe appear in 3P or 6P chromosome. All these resistant genes could express fine under common wheat genome background.
Yellow rust and leaf rust resistance identification were engaged in low-temperature growth chamber and field with mixture spore inoculated of 4 yellow rust isolates and 8 leaf rust isolates, respectively. The results indicated the gene resistant to yellow rust was located on 6P chromosome, with moderate degree resistance in wheat background. But genes for leaf rust resistance with immunity response were pointed on 3P, 7P and 2P chromosome. These genes could sufficiently express in wheat background, will being important gene reservoir in wheat breeding for leaf rust resistance.
The drought resistance identification of the full et of disomic addition lines with the method of "repeated drought periods" indicated that genes for drought resistance mainly scatter in the 2P, 4P, and 7P chromosomes. The resistant strength of each disomic addition line was lower than the A. cristatum parent. The genes for cold tolerance were located on 4P and 6P chromosomes. This cold tolerance response was weaker than the parent A. cristatum.
Identification of genes for some of the important agronomic traits originated in A.cristatum shown that: genes related to vernalization marked with creeping stem, curl or semi-curl and dark green leaf in seedling stage were located on 2P, 3P and 5P chromosomes; genes controlling tiller number were being 2P and 6P, and genes for potential high yield characterized by many spikelet number, many floscular number, longer peduncle length, and shorter plant height were located on 5P chromosomes but with an undesirable late maturity gene. Additionally, gene(s) for awnlessness appeared on 3P, 6P chromosomes.
By electrophoresis gel analysis of HMW glutenin subunits in seed of a set of wheat-A.cristatum addition lines and A.cristatum, it's shown that there are genes in A. cristatum (Z559) inhibiting the expression of HMW glutenin subunits under wheat background. The HMW glutenin genes of A. cristatum (Z559) no express, the female Fukuho possesses 5 subunits: 2．2+12 (encoded by D genome), 2* (A genome) and 7+8 (B genome), but the being of P chromosome had alternative inhibition effect to expression of 2．2 and 12 subunits encoded by D genome in different wheat-A.cristatum alien addition lines.
The expression pattern of genes for gliadin subunits of A. cristatum < L. > Gaertn (Z559) was identified. There were richω-gliadin subunits in Z559 but disappeared all these subunits in the 16 wheat-A. cristatum alien addition lines including a addition line with 1P marker. It's already clear thatω-gliadin is controlled by gene family of the first homologous group, so, perhaps there are two kind of situations: default 1P addition line or disable expression ofω-gliadin genes under wheat background. Two specificα-andβ-gliadin subunits were taken out in four 6P addition lines and Z559 resemble to two strong gluten wheat varieties, Marquis and McGuire. It's indicated that theseαandβ-gliadin genes could be not only resources of quality improvement for wheat but 锘縜lso biochemical markers of 6P chromosome. Otherwises, changes of蠅and纬-gliadin subunits was discovered in wheat-A. cristatum derivatives Wula and Lo12．鈽匒 new powdery mildew gene originated in A. cristatum was named based on data of molecular mapping and response pattern analysis. Wula, a wheat-A.cristatum derivative, selected from BC_2 of hybrid Fukuho脳Z559 by pedigree possesses gene immune to powdery mildew. Response pattern analysis indicated that this gene immune to almost all the powdery mildew isolates except to E21 with middle resistance. By establishing DH population of Wula脳Wenmai No.6(susceptible), genetic location and molecular mapping of the powdery mildw gene were carried out. The response pattern of DH families showed 1:1 ratio of resistance to susceptible individuals indicates that the resistance controlled by a single dominant gene. Bulked segregant analysis was used to identify microsatellite markers linked to the gene. Three SSR markers Xgwm341, Xgwm383 and Xgwm645 were mapped in coupling phase to the gene locus, thus, this resistance gene was located on long arm of 3D chromosome. The gene was mapped between Xgwm341 to Xgwm383 with 29．4cM and 26．6cM respectively. And Xgwm645 was mapped 26．4 cM out of Xgwm341．Because there hasn't been powdery mildew gene was located on 3DL until now, so, this is a new powdery mildew resistance gene originated in A. cristatum. It is designated Pm31 tentatively. The paper discussed the propably utilization of these important gene resources, given the suggestions and the perspects in the future.
引文
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