小麦成株抗条锈病的组织学和细胞学研究及小麦非寄主抗蚕豆锈病的机理研究
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摘要
由小麦条锈菌(Puccinia. striiformis f. sp. tritici)引发的小麦条锈病是威胁我国小麦生产的最重要病害之一。成株抗病性(Adult plant resistance APR)具有持久抗病性的特点，表现为苗期感病，成株期抗病，其在生产上具有巨大的应用潜力。从组织学和细胞学角度开展小麦成株抗病性互作特征研究，将为进一步解析成株抗性的分子机理奠定基础和提供依据，对选育持久抗病品种和实现小麦条锈病的可持续控制具有重要的理论和实际意义。本研究以小麦兴资9104和小麦条锈菌32号小种(CYR32)为研究对象，通过不同叶位的连续接种进行表型鉴定；对兴资从感病到抗病过程转变的关键生育期与锈菌的互作进行系统的组织学分析；通过组织化学的方法对互作过程中活性氧的积累进行检测；通过细胞学方法对成株抗性表达过程中植物与病菌的超微结构特征进行观测，从而阐明兴资9104成株抗小麦条锈病的组织学和细胞学特征。本研究的主要结果如下:1、通过对兴资9104不同叶位连续接种实验表明，兴资9104对CYR32的成株抗病性从分蘖期开始表达，随着生长发育逐渐增强，孕穗期后表现出最强的抗病性；2、光镜观测表明，三种组合中病菌均于接种后12h进入植物叶组织而形成气孔下囊，然而苗期、分蘖期和孕穗期组合中的最早的吸器形成时间分别为接种后12h、18h和24h，表明病菌的吸器分化和形成在成株期组合中明显的延迟和受抑。接种后48h，三种组合中，病菌在菌丝分枝数、菌丝长度和吸器母细胞的数量上差异并不明显，但苗期组合中，病菌从接种后48h开始，大量分化形成次生菌丝，从而促进其菌落的形成；而孕穗期组合中多数侵染点的的次生菌丝出现于接种后96h，表明病菌次生菌丝的发育明显的被抑制或延迟，并导致菌落形成的减少。病菌的侵染和吸器的形成在分蘖期和孕穗期组合中伴有植物细胞的坏死，这种典型的过敏性坏死在两种组合中分别出现于接种后18h和36h，随后含坏死细胞的侵染点比率和坏死的面积逐渐增加。接种后120h，分蘖期组合中80%的侵染点含有坏死细胞，而至接种后144h，孕穗期组合中所有的侵染点均有细胞坏死出现，部分侵染点的病菌结构完全被坏死细胞所包围，导致其并无次生菌丝的形成及败育。表明植物细胞坏死对病菌的发育具有明显的抑制作用。3、活性氧的组织化学检测结果表明，分蘖期和孕穗期组合中，于接种后24h和36h分别形成O_2迸发高峰，大量的侵染点出现NBT (Nitroblue tetrazolium)着色的时间点与吸器母细胞与植物细胞接触形成吸器的时间点一致，而NBT着色的细胞仍然保持生存状态。因此，O_2迸发早于病菌入侵细胞的坏死。而分蘖期和孕穗期组合中，被侵染的植物叶片中H_2O_2积累具有相似的双时相特征。在分蘖期中，第一个H_2O_2迸发高峰出现于接种后24h，而第二个H_2O_2迸发高峰出现于接种后72-96h；孕穗期的组合中第一个H_2O_2迸发高峰，出现于接种后48h，而第二个H_2O_2迸发高峰出现于接种后120h。H_2O_2第一个迸发时相与吸器的形成以及随后入侵细胞的坏死的时间点相一致，而其第二个迸发时相明显的与次生菌丝的形成以及随后的侵染细胞的坏死密切相关。而苗期组合中，大多数侵染点中，病菌的侵染并没有诱导O_2-和H_2O_2的产生。因此，O_2-和H_2O_2的产生和积累与寄主细胞过敏性坏死有着密切关系，在成株抗病性表达中可能起着重要作用。4、透射电镜观测表明，兴资对条锈菌成株抗病性的典型的结构防卫反应包括显著的细胞壁相关的抗性、入侵细胞的大量坏死以及吸器鞘的形成，这些亚细胞水平的防卫结构与抑制病菌的生长、加速病菌的衰老、导致侵染结构的细胞死亡密切相关。
非寄主抗病性(non-host resistance NHR)是整个植物物种，对特定的病菌物种的所有菌株所表现出的抗病性，是植物所表现出的最普遍的抗病形式，其具有持久抗性的特点，因此，在农业生产中具有广阔的应用前景。作为重要的农作物，小麦是柄锈菌属的寄主，长期遭到条锈菌、叶锈菌(P. triticina)和杆锈菌(P. graminis f. sp. tritici)的危害，但其对以与其亲缘关系较远的豆类植物作为寄主的单胞锈菌却保持抗病性，其中的非寄主抗病性的机制知之甚少。因此，本研究以小麦(Triticum aestivum)与蚕豆锈菌构成了一个新的研究体系，通过组织学和细胞学的研究，对病菌的发育进程和小麦的防卫反应进行观测；通过组织化学的方法检测互作过程中活性氧的迸发情况；通过实时定量PCR (quantitativeRT-PCR qRT-PCR)检测防卫相关基因和活性氧代谢相关基因的表达情况。本研究结果将揭示小麦对与其协同进化关系较远的豆类锈菌的非寄主抗病性的细胞学特征和分子基础，有助于对小麦抗锈性的全面理解。其主要研究结果如下：
1、光镜观测表明蚕豆锈菌在小麦叶片上能有效的萌发形成芽管，但蚕豆锈菌在小麦叶片上的气孔侵入率显著下降，超过98%的芽管(萌发体)不能准确的定位气孔并于其上方形成附着胞而进入叶组织。少数萌发体能进入气孔形成侵染点，从而分化形成胞间菌丝和吸器母细胞，但大多数的吸器母细胞未能侵入叶肉细胞而形成吸器。接种后96h，侵染点中吸器的形成率不到4%。
2、透射电镜观测表明病菌吸器母细胞的入侵诱导了植物细胞壁的加固和胞壁沉积物以及乳突的形成，入侵栓尽管能穿透细胞壁，但不能穿过乳突而于细胞内形成吸器。因此细胞壁相关的抗病性导致了病菌的败育，说明小麦对蚕豆锈菌的抗病性主要为吸器前抗性。尽管少数病菌的入侵栓能够穿透植物细胞的细胞壁而形成吸器，但光镜观测发现吸器被类似胼胝质的沉积物所包围，并发出强烈的自发荧光，这种吸器鞘状的结构可能限制吸器的发育和功能的发挥。同时吸器的形成并未激发植物过敏性坏死的形成，这与接种蚕豆锈菌后小麦叶片无任何症状的表型相一致。
3、活性氧的组织化学检测结果表明，蚕豆锈菌的侵染诱导了小麦细胞中H_2O_2的积累，但无O_2形成。H_2O_2主要分布于与病菌接触的细胞壁、正对吸器母细胞的乳突结构和吸器鞘部位，这些形成于防卫结构中的H_2O_2，可能有助于通过交联而加固病菌接触部位的细胞壁和乳突以及吸器鞘结构，并进而阻止了病菌在小麦组织中的发育。
4、qRT-PCR分析结果表明， TaPR1(病程相关蛋白1)、 TaPR2(-1,3-葡聚糖酶)、 TaPR3(几丁质酶)、 TaPR5(类甜蛋白)、 TaPAL (苯丙氨酸解氨酶), TaSOD (超氧化物歧化酶)、TaCAT (过氧化氢酶)和TaAPX (抗坏血酸-过氧化物酶)基因在互作中呈现出不同的表达特征，但所有基因于接种后12-48h之间呈现出一到多个上调表达的时间点，表明活性氧代谢相关基因和防卫相关的基因同样参与了小麦对蚕豆锈的非寄主抗病性，前者可能通过代谢调控活性氧的产生并避免活性氧的过量积累而激发细胞的坏死，后者参与小麦的非寄主抗病性则表明小麦对锈菌的非寄主抗病性和其基础抗性可能存在交叉和重叠。
Stripe rust of wheat, caused by Puccinia striiformis f. sp. tritici (Pst), is one of the mostdamaging diseases of wheat in China. Adult plant resistance (APR) to stripe rust is knownto be an important component of some sources of durable resistance in wheat, thus has greatpractical potentials in wheat production. Characterization of APR to stripe rust at histologicaland cytological level will set the foundation for further elucidation the mechanism of APR atmolecular level, therfore, will be of greatly important for appropriate utilization of resistantcultivars and sustainable control of the disease. Thus the present study was conducted toelucidate the phenotype of XZ9104(XZ) relative to leaf position and different plant growthstages by Pst(CYR32) inoculation. In addition, histopathological analysis was undertaken tocharacterize the cellular interaction between Pst and XZ at seedling (GS12), tillering(GS21-22) and boot stages (GS43-49), which represent the expression of resistance transitionfrom susceptibility to resistance. Also, the accumulation of ROS (reactive oxygen species)and variation in resistance in this interaction were studied systematically using histochemicalmethods. Finally, transmission electron microscopy (TEM) was used to describeultrastructural changes during XZ and Pst interactions. Our study will character the APR inXZ to Pst at histological and cytological level. The main results are as follows:1. To elucidate the correlation of APR expression and plant development, the leaves invarious positions at different growth stages were inoculated simultaneously with Pst raceCYR32in the greenhouse. The study result indicated that APR was initiated as early astillering stage, increased gradually as plants grew, and most effective at the boot stage.2. The microscopical observation revealed that the time points of the appearance ofhaustorium formation in the infection sites of plant leaves in the seedling, tillering and bootstages were12,18and24hai, respectively, although the pathogen entered plant leaves andformed a substomatal vesicle at12hai in all stages. It indicated that haustorial formation wasretarded or inhibited in adult plants. The hypha branching and length of hypha as well asnumber of haustorial mother cell (HMC) in all growth stages were similar until48hai andwithout obvious difference between all growth stages. In the seedling-stage plants, after48hai, massive formation of secondary hyphae as Pst proliferated rapidly which subsequently confer the formation of small microcolony. In most of infection site, the secondary hyphaappeared at96hai in boot-stage plants. It indicated that the differentiation of secondaryhyphae was significantly inhibited, which decreased the development of microcolonies inplants of boot stage. One of the observable responses at the microscopic level was thenecrosis associated with the invaded plant cells coincided with the haustorial formation in thetillering and boot stages plants. The typical hypersensitive response (HR) was observed from18to36hai in leaves of plants in tillering and boot stages, respectively. With a longerincubation time, the percentage of infection sites with necrotic host cells and the necrotic areaincreased in inoculated leaves of plants in both tillering and boot stages, necrotic host cellswere observed in80%of the infection sites in the tillering plants by120hai and in100%ofthe infection sites of the boot-stage plants at144hai. At some infection site of adult plant,the infection unit were completely surrounded by the necrotic tissue which render the fungalabortion and lack of secondary hypha formation. It indicated that the death of host cellsmarkedly inhibited the development of hyphae.
3. The result of histochemical location of ROS revealed that a high frequency of NBT(Nitroblue tetrazolium) stained cells showed O_2accumulation was detected at24hai ontillering stage plants and at36hai on boot-stage plants, respectively. The high frequency ofNBT stained cells coincided with the infection stage when contacts were established betweenthe HMCs and the mesophyll cells and haustoria began to form, and while the host cells withNBT staining remained alive. It indicated that the production of O_2preceded HR of infectedcells. The H_2O_2accumulations in infected leaves of plants in tillering and booting stages weresimilar and both showed a biphasic process. On tillering plants, The first and second peak ofH_2O_2generation in invaded host cells appeared at24hai and72-96hai, respectively. Whileon boot plants these peaks appeared at48hai and120hai, respectively. The first burst phaseof H_2O_2coincided with the beginning of haustorium formation and the subsequent HR ininvaded host cells, while the second phase of H_2O_2generation coincided with the formation ofsecondary hyphae and the increasing number of necrotic host cells surrounding the attackedcells. In the infected leaves of seedlings, in contrast, at most infection sites, the pathogeninfection didn’t induce obvious H_2O_2and O_2generation. The result suggested that, thegeneration and accumulation of O_2-and H_2O_2are closely associated with the HR of host cells,and which may play an important role in the expression of APR.
4. The result of TEM study revealed that the typical APR reactions of XZ at the subcellularlevel included significant cell wall-related resistance, necrosis of penetrated host cells, and theformation of a haustorial sheath. These structural defense responses are believed tosignificantly inhibit the growth of Pst by accelerating the senescence of hyphae and cell death of infection structures.
Non-host resistance (NHR), which is defined as resistance shown by all genotypes of aplant species against all genotypes of a given pathogen species, is the most common form ofdisease resistance exhibited by plants against the majority of potentially pathogenicmicroorganisms. NHR to cereal rusts has proved to be durable and appears to have greatpotential for engineering plant immunity. Little is known about the nature of this effectivedefense mechanism to the pathogen of remotely related plant species in important cropspecies, wheat, which can be damaged by several Puccinia species, P. striiformis f. sp. tritici,P. triticina and P. graminis f. sp. tritici, but is immune to all Uromyces species. Here weestablished a novel non-adapted pathosystem based on wheat (Triticum aestivum) and thebroad bean rust pathogen Uromyces fabae (Uf). We first investigated the reaction of a wheatcultivar to the infection by the broad bean rust pathogen using light and electron microscopes.Histochemical experiments were conducted to detect ROS generation in the non-hostpathosystem. Expression profiles of candidate genes related to basal resistance and oxidativestress in wheat leaves inoculated with Uf were assayed by quantitative RT-PCR (qRT-PCR).Our study will revealed the cytological and molecular basis of NHR in wheat against rustpathogen of remotely related plant species and lead to a comprehensive understanding ofresistance in wheat against rust fungi.
1. In the wheat-Uf interaction, microscopic observations showed that urediospores germinatedefficiently on wheat leaves. However, over98%of the germ tubes failed to form appressoriaover stomata and developed substomatal vesicles inside wheat leaves. Therefore, stomapenetration was significantly reduced in Uf infection of wheat. For the few that invadedthrough stomata and formed the intercellular hypha and haustorial mother cell, the majority ofHMC failed to penetrate wheat mesophyll cells. At96hours after inoculation, less than4%ofthe Uf infection units that had entered the mesophyll tissue formed haustoria.
2. The result of TEM study revealed that the Attempted penetration by haustorium mothercells induced the thickening of cell wall and the formation of papillae in plant cells, thepenetration peg emerging from the HMC appeared to puncture through the plant cell wall butfailed to penetrate through the papilla and form haustoria in wheat cells. Thus, it indicatedthat cell wall-related resistance arrested the development or growth of Uf penetration pegs andNHR of wheat to Uf is pre-haustorial. Although a few penetration pegs could breach the cellwall and periphery deposits and formed haustoria, the light microscopic observation revealedthat these haustoria formed by Uf in wheat cells were encased in callose-like deposits that hadstrong auto-fluorescence. Moreover, the formation of haustoria did not trigger thehypersensitive response in penetrated wheat cells, which was consistent with the macroscopic phenotype.
2. The result of histochemical location of AOS revealed that in responses to Uf infection,accumulation of H_2O_2, but not O_2, was detected in wheat cells. H_2O_2was mainly detected incell wall in close contact with IH or HMCs, papillae subjacent to HMC, and depositsresponsible for haustorium encasement. The cross-link fueled by H_2O_2likely reinforces thecell wall at the sites in contact with fungal structures and strengthening papillae as well as theencasement of haustoria and results in the arrest of Uf growth in wheat.
3. The putative TaPR1(pathogenesis-related protein1), TaPR2(-1,3-glucanase), TaPR3(chitinase), TaPR5(thaumatin-like), TaPAL (phenylalanine ammonia lyase), TaSOD(superoxide dismutase), TaCAT (catalase), and TaAPX (ascorbate-peroxidase) genes wereselected as candidate defense-related genes for assaying their expression levels during Ufinfection by qRT-PCR. Although they had different expression profiles, all of these eightgenes were up-regulated at one or more time-points between12-48hai. It revealed that thesedefense related genes also may contribute to preventing Uf infection of wheat, genes involvedin oxidative stress may regulate the ROS generation and avoid over-accumulation of ROS andeliciting HR, similar PR genes have been implicated in wheat resistance against both adaptedand non-adapted rust fungi may indicate that NHR and basal resistance to adapted pathogensmay involve the same or similar mechanisms in wheat.
非寄主抗病性(non-host resistance NHR)是整个植物物种，对特定的病菌物种的所有菌株所表现出的抗病性，是植物所表现出的最普遍的抗病形式，其具有持久抗性的特点，因此，在农业生产中具有广阔的应用前景。作为重要的农作物，小麦是柄锈菌属的寄主，长期遭到条锈菌、叶锈菌(P. triticina)和杆锈菌(P. graminis f. sp. tritici)的危害，但其对以与其亲缘关系较远的豆类植物作为寄主的单胞锈菌却保持抗病性，其中的非寄主抗病性的机制知之甚少。因此，本研究以小麦(Triticum aestivum)与蚕豆锈菌构成了一个新的研究体系，通过组织学和细胞学的研究，对病菌的发育进程和小麦的防卫反应进行观测；通过组织化学的方法检测互作过程中活性氧的迸发情况；通过实时定量PCR (quantitativeRT-PCR qRT-PCR)检测防卫相关基因和活性氧代谢相关基因的表达情况。本研究结果将揭示小麦对与其协同进化关系较远的豆类锈菌的非寄主抗病性的细胞学特征和分子基础，有助于对小麦抗锈性的全面理解。其主要研究结果如下：
1、光镜观测表明蚕豆锈菌在小麦叶片上能有效的萌发形成芽管，但蚕豆锈菌在小麦叶片上的气孔侵入率显著下降，超过98%的芽管(萌发体)不能准确的定位气孔并于其上方形成附着胞而进入叶组织。少数萌发体能进入气孔形成侵染点，从而分化形成胞间菌丝和吸器母细胞，但大多数的吸器母细胞未能侵入叶肉细胞而形成吸器。接种后96h，侵染点中吸器的形成率不到4%。
2、透射电镜观测表明病菌吸器母细胞的入侵诱导了植物细胞壁的加固和胞壁沉积物以及乳突的形成，入侵栓尽管能穿透细胞壁，但不能穿过乳突而于细胞内形成吸器。因此细胞壁相关的抗病性导致了病菌的败育，说明小麦对蚕豆锈菌的抗病性主要为吸器前抗性。尽管少数病菌的入侵栓能够穿透植物细胞的细胞壁而形成吸器，但光镜观测发现吸器被类似胼胝质的沉积物所包围，并发出强烈的自发荧光，这种吸器鞘状的结构可能限制吸器的发育和功能的发挥。同时吸器的形成并未激发植物过敏性坏死的形成，这与接种蚕豆锈菌后小麦叶片无任何症状的表型相一致。
3、活性氧的组织化学检测结果表明，蚕豆锈菌的侵染诱导了小麦细胞中H_2O_2的积累，但无O_2形成。H_2O_2主要分布于与病菌接触的细胞壁、正对吸器母细胞的乳突结构和吸器鞘部位，这些形成于防卫结构中的H_2O_2，可能有助于通过交联而加固病菌接触部位的细胞壁和乳突以及吸器鞘结构，并进而阻止了病菌在小麦组织中的发育。
4、qRT-PCR分析结果表明， TaPR1(病程相关蛋白1)、 TaPR2(-1,3-葡聚糖酶)、 TaPR3(几丁质酶)、 TaPR5(类甜蛋白)、 TaPAL (苯丙氨酸解氨酶), TaSOD (超氧化物歧化酶)、TaCAT (过氧化氢酶)和TaAPX (抗坏血酸-过氧化物酶)基因在互作中呈现出不同的表达特征，但所有基因于接种后12-48h之间呈现出一到多个上调表达的时间点，表明活性氧代谢相关基因和防卫相关的基因同样参与了小麦对蚕豆锈的非寄主抗病性，前者可能通过代谢调控活性氧的产生并避免活性氧的过量积累而激发细胞的坏死，后者参与小麦的非寄主抗病性则表明小麦对锈菌的非寄主抗病性和其基础抗性可能存在交叉和重叠。
Stripe rust of wheat, caused by Puccinia striiformis f. sp. tritici (Pst), is one of the mostdamaging diseases of wheat in China. Adult plant resistance (APR) to stripe rust is knownto be an important component of some sources of durable resistance in wheat, thus has greatpractical potentials in wheat production. Characterization of APR to stripe rust at histologicaland cytological level will set the foundation for further elucidation the mechanism of APR atmolecular level, therfore, will be of greatly important for appropriate utilization of resistantcultivars and sustainable control of the disease. Thus the present study was conducted toelucidate the phenotype of XZ9104(XZ) relative to leaf position and different plant growthstages by Pst(CYR32) inoculation. In addition, histopathological analysis was undertaken tocharacterize the cellular interaction between Pst and XZ at seedling (GS12), tillering(GS21-22) and boot stages (GS43-49), which represent the expression of resistance transitionfrom susceptibility to resistance. Also, the accumulation of ROS (reactive oxygen species)and variation in resistance in this interaction were studied systematically using histochemicalmethods. Finally, transmission electron microscopy (TEM) was used to describeultrastructural changes during XZ and Pst interactions. Our study will character the APR inXZ to Pst at histological and cytological level. The main results are as follows:1. To elucidate the correlation of APR expression and plant development, the leaves invarious positions at different growth stages were inoculated simultaneously with Pst raceCYR32in the greenhouse. The study result indicated that APR was initiated as early astillering stage, increased gradually as plants grew, and most effective at the boot stage.2. The microscopical observation revealed that the time points of the appearance ofhaustorium formation in the infection sites of plant leaves in the seedling, tillering and bootstages were12,18and24hai, respectively, although the pathogen entered plant leaves andformed a substomatal vesicle at12hai in all stages. It indicated that haustorial formation wasretarded or inhibited in adult plants. The hypha branching and length of hypha as well asnumber of haustorial mother cell (HMC) in all growth stages were similar until48hai andwithout obvious difference between all growth stages. In the seedling-stage plants, after48hai, massive formation of secondary hyphae as Pst proliferated rapidly which subsequently confer the formation of small microcolony. In most of infection site, the secondary hyphaappeared at96hai in boot-stage plants. It indicated that the differentiation of secondaryhyphae was significantly inhibited, which decreased the development of microcolonies inplants of boot stage. One of the observable responses at the microscopic level was thenecrosis associated with the invaded plant cells coincided with the haustorial formation in thetillering and boot stages plants. The typical hypersensitive response (HR) was observed from18to36hai in leaves of plants in tillering and boot stages, respectively. With a longerincubation time, the percentage of infection sites with necrotic host cells and the necrotic areaincreased in inoculated leaves of plants in both tillering and boot stages, necrotic host cellswere observed in80%of the infection sites in the tillering plants by120hai and in100%ofthe infection sites of the boot-stage plants at144hai. At some infection site of adult plant,the infection unit were completely surrounded by the necrotic tissue which render the fungalabortion and lack of secondary hypha formation. It indicated that the death of host cellsmarkedly inhibited the development of hyphae.
3. The result of histochemical location of ROS revealed that a high frequency of NBT(Nitroblue tetrazolium) stained cells showed O_2accumulation was detected at24hai ontillering stage plants and at36hai on boot-stage plants, respectively. The high frequency ofNBT stained cells coincided with the infection stage when contacts were established betweenthe HMCs and the mesophyll cells and haustoria began to form, and while the host cells withNBT staining remained alive. It indicated that the production of O_2preceded HR of infectedcells. The H_2O_2accumulations in infected leaves of plants in tillering and booting stages weresimilar and both showed a biphasic process. On tillering plants, The first and second peak ofH_2O_2generation in invaded host cells appeared at24hai and72-96hai, respectively. Whileon boot plants these peaks appeared at48hai and120hai, respectively. The first burst phaseof H_2O_2coincided with the beginning of haustorium formation and the subsequent HR ininvaded host cells, while the second phase of H_2O_2generation coincided with the formation ofsecondary hyphae and the increasing number of necrotic host cells surrounding the attackedcells. In the infected leaves of seedlings, in contrast, at most infection sites, the pathogeninfection didn’t induce obvious H_2O_2and O_2generation. The result suggested that, thegeneration and accumulation of O_2-and H_2O_2are closely associated with the HR of host cells,and which may play an important role in the expression of APR.
4. The result of TEM study revealed that the typical APR reactions of XZ at the subcellularlevel included significant cell wall-related resistance, necrosis of penetrated host cells, and theformation of a haustorial sheath. These structural defense responses are believed tosignificantly inhibit the growth of Pst by accelerating the senescence of hyphae and cell death of infection structures.
Non-host resistance (NHR), which is defined as resistance shown by all genotypes of aplant species against all genotypes of a given pathogen species, is the most common form ofdisease resistance exhibited by plants against the majority of potentially pathogenicmicroorganisms. NHR to cereal rusts has proved to be durable and appears to have greatpotential for engineering plant immunity. Little is known about the nature of this effectivedefense mechanism to the pathogen of remotely related plant species in important cropspecies, wheat, which can be damaged by several Puccinia species, P. striiformis f. sp. tritici,P. triticina and P. graminis f. sp. tritici, but is immune to all Uromyces species. Here weestablished a novel non-adapted pathosystem based on wheat (Triticum aestivum) and thebroad bean rust pathogen Uromyces fabae (Uf). We first investigated the reaction of a wheatcultivar to the infection by the broad bean rust pathogen using light and electron microscopes.Histochemical experiments were conducted to detect ROS generation in the non-hostpathosystem. Expression profiles of candidate genes related to basal resistance and oxidativestress in wheat leaves inoculated with Uf were assayed by quantitative RT-PCR (qRT-PCR).Our study will revealed the cytological and molecular basis of NHR in wheat against rustpathogen of remotely related plant species and lead to a comprehensive understanding ofresistance in wheat against rust fungi.
1. In the wheat-Uf interaction, microscopic observations showed that urediospores germinatedefficiently on wheat leaves. However, over98%of the germ tubes failed to form appressoriaover stomata and developed substomatal vesicles inside wheat leaves. Therefore, stomapenetration was significantly reduced in Uf infection of wheat. For the few that invadedthrough stomata and formed the intercellular hypha and haustorial mother cell, the majority ofHMC failed to penetrate wheat mesophyll cells. At96hours after inoculation, less than4%ofthe Uf infection units that had entered the mesophyll tissue formed haustoria.
2. The result of TEM study revealed that the Attempted penetration by haustorium mothercells induced the thickening of cell wall and the formation of papillae in plant cells, thepenetration peg emerging from the HMC appeared to puncture through the plant cell wall butfailed to penetrate through the papilla and form haustoria in wheat cells. Thus, it indicatedthat cell wall-related resistance arrested the development or growth of Uf penetration pegs andNHR of wheat to Uf is pre-haustorial. Although a few penetration pegs could breach the cellwall and periphery deposits and formed haustoria, the light microscopic observation revealedthat these haustoria formed by Uf in wheat cells were encased in callose-like deposits that hadstrong auto-fluorescence. Moreover, the formation of haustoria did not trigger thehypersensitive response in penetrated wheat cells, which was consistent with the macroscopic phenotype.
2. The result of histochemical location of AOS revealed that in responses to Uf infection,accumulation of H_2O_2, but not O_2, was detected in wheat cells. H_2O_2was mainly detected incell wall in close contact with IH or HMCs, papillae subjacent to HMC, and depositsresponsible for haustorium encasement. The cross-link fueled by H_2O_2likely reinforces thecell wall at the sites in contact with fungal structures and strengthening papillae as well as theencasement of haustoria and results in the arrest of Uf growth in wheat.
3. The putative TaPR1(pathogenesis-related protein1), TaPR2(-1,3-glucanase), TaPR3(chitinase), TaPR5(thaumatin-like), TaPAL (phenylalanine ammonia lyase), TaSOD(superoxide dismutase), TaCAT (catalase), and TaAPX (ascorbate-peroxidase) genes wereselected as candidate defense-related genes for assaying their expression levels during Ufinfection by qRT-PCR. Although they had different expression profiles, all of these eightgenes were up-regulated at one or more time-points between12-48hai. It revealed that thesedefense related genes also may contribute to preventing Uf infection of wheat, genes involvedin oxidative stress may regulate the ROS generation and avoid over-accumulation of ROS andeliciting HR, similar PR genes have been implicated in wheat resistance against both adaptedand non-adapted rust fungi may indicate that NHR and basal resistance to adapted pathogensmay involve the same or similar mechanisms in wheat.
引文
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