高粱靶斑病菌致病机理及抗病基因分子标记定位
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摘要
高粱是重要的食用、酿酒、饲用和工业用原料，病菌侵染会导致粮食品质变劣、产量降低。同时，病原菌会产生真菌毒素，危害人类和动物健康。由生离蠕孢属病菌侵染所致高粱靶斑病（Bipolaris sorghicola）是高粱重要的叶部病害之一，病菌的致病机理以及遗传机制和分子水平的研究，在国内尚未见报道。本文围绕高粱靶斑病菌致病机理及品种抗靶斑病基因定位研究目标，在高粱靶斑病菌生物特性、病原菌种群多样性，病原菌粗毒素对种子萌发、根系生长及其对高粱胚根细胞膜透性和防御酶活性的影响，病原菌侵染后的高粱叶片光合指标和光系统Ⅱ的变化，高粱抗靶斑病特性遗传规律及抗病基因标记与定位，等诸多方面进行了较系统研究，获得了较理想结果，部分结果填补了我国该研究空白，为高粱抗靶斑病育种及病害治理等提供了科学依据。论文主要结果总结如下：
1.高粱靶斑病病原菌的生物学特性
生物学特性研究结果表明：病菌菌丝生长的适宜温度为25～30℃，分生孢子萌发的适宜温度为20～30℃。分生孢子萌发的适宜pH为3～6，偏酸性。大多糖类均可作为该病的碳源营养，如半乳糖和乳糖，而山梨糖对菌丝生长不太适宜。不同光照处理对病菌菌丝生长的影响不明显。
2.我国高粱靶斑病病原菌种类及种群多样性
从采自我国不同地区的感染高粱靶斑病病叶中分离出病原菌，通过形态学观察和rDNA-ITS序列分析进行多样性研究。结果表明，虽然这些发病地区病叶上的病斑差异很大，但病原菌在菌落形态和孢子形态方面却极为相似;通过这些菌株的rDNA-ITS序列聚类分析表明，它们与Bipolaris sorghicola相似度达99%,从而证实虽然不同地区病斑差异较大，但都是同一病原菌引起的病害。
3.高粱靶斑病菌粗毒素能降低细胞膜透性
采用测定不同浓度病原菌粗毒素处理的高粱种子L407B和Tx622B的胚根生长抑制率以及电导率和丙二醛(MDA)含量的方法，探讨高粱靶斑病菌粗毒素对高粱不同品种胚根生长及胚根细胞膜透性的影响。结果表明，病菌粗毒素对高粱胚根生长有明显的抑制作用。在同一毒素浓度下，处理后2d，处理组胚根的相对电导率和MDA含量均高于对照组；同一处理时间内，不同浓度毒素处理后胚根相对电导率和MDA含量均高于对照组。说明高粱靶斑病菌毒素能降低高粱胚根细胞膜透性，且不同高粱品种间差异显著。
4.高粱靶斑病菌粗毒素能抑制寄主保护酶活性
利用紫外可见分光光度计和比色法测定了不同浓度病原菌粗毒素处理的高粱种子L407B和Tx622B胚根保护酶（过氧化物酶、超氧化物歧化酶、多酚氧化酶和苯丙氨酸解氨酶）活性。结果表明，毒素处理后，高粱种子胚根过氧化物酶、超氧化物歧化酶、多酚氧化酶和苯丙氨酸解氨酶活性明显降低，说明毒素对寄主保护酶起抑制作用。
5.高粱靶斑病病原菌侵染会破坏高粱叶片PSII的结构和功能
以高粱品种Tx622B为试验材料,通过同时测定高粱叶片叶绿素含量、叶绿素荧光快速诱导动力学曲线，以及过氧化氢(H2O2)含量和丙二醛(MDA)含量的变化，来研究高粱靶斑病菌[Bipolaris sorghicola (Lefebvre&Sherwin) Alcorn]侵染高粱叶片后对光系统II(PSII)结构和功能的影响。研究结果表明，受病原菌侵染的高粱叶片,H2O2和MDA含量均升高,膜脂过氧化的程度加剧；快速叶绿素荧光诱导动力学曲线发生明显变化，进一步的JIP测试分析表明病原菌侵染严重伤害了PSII供体侧、阻断了受体侧QA到QB的电子传递，从而使反应中心的活性降低。
6.抗高粱靶斑病基因为隐性单基因
以高粱靶斑病抗病自交系L407B和感病自交系Tx622B的杂交后代构建的重组自交系群体为材料，利用数量遗传分析方法研究了L407B的抗病遗传规律。对高粱靶斑病田间性状调查结果进行分析，经卡方分析检验符合孟德尔单基因控制的遗传分离规律。F1代田间表现与亲本Tx622B完全相同，在F2中，抗病植株占F2代总植株的四分之一，初步证实自交系L407B的抗病性是受一对隐性基因控制的质量性状。
7.抗病基因在染色体上的位置
从覆盖高粱染色体组的132对以“Xtxp-”开头的SSR引物和230对以“sm-”开头的SSR引物中筛选出在抗感两亲本之间有差异且在抗性亲本和F2代抗池间无差异的引物。通过染色体步移方法，将高粱抗靶斑病基因定位于标记Xtmp303和sm05014之间，即该抗病基因位于第5条染色体短臂上5.73M—8.15M内。
Sorghum is important as food, wine, feed and industrial raw materials. Sorghum infectedby pathogens can lead to poor food quality and lower production.At the same time, pathogenscan produce mycotoxin to harm humans and animals. Target leaf spot which casued byBipolaris sorghicola is one of important foliar disease in sorghum varieties. At present, thereis no report on pathopoiesis mechanism as well as inheritance mechanism and moleculargenetics of this disease in China. This paper focuses on the pathogenesis and location ofresistance gene against target leaf spot in sorghum. There was a relatively systematic study inthe biological characteristics and population diversity of pathogen, the effect of pathogencrude toxins on seed germination and root growth as well as cell membrane permeability anddefense enzyme activity in sorghum radicle, the changes of sorghum leaf photosyntheticindexes and photosystem Ⅱa fter pathogen infection, the genetic law of resistance to thedisease and the marker and location of resistance gene and many other aspects. The studyobtained ideal results, parts of them filled in the blank about this research in China. Theresults provided scientific basis for breeding of resistant varieties and disease management.
The main results were summarized as follows:
1. Biological characteristics of pathogen which caused target leaf spot on sorghum
The result indicated that optimum temperature for pathogen mycelia growth was25-30℃, and that for conidiagermination was20-30℃.The optimum pH for conidiagermination was-6, leaning to acidity. The pathogen could use many kind of sugar as acarbon nutrient，eg.lactose and galactose. However, sorbose was not fit for pathogen myceliagrowth. Different light condition for mycelium growth was not obvious.
2. The category and population diversity of pathogen in China
In this diversity research, we isolated pathogens from infected target leaf spot diseaseleaves of different regions of China by morphological observation and rDNA-ITS sequenceanalysis.The results show that although these lesions have significant differences, the colonymorphology and spore morphology of pathogens are very similar. The rDNA-ITS sequence cluster analysis showed that the similarity of these pathogens with Bipolaris sorghicola reachto99%.All of these confirmed this disease caused by the same pathogen,although theselesions have significant differences.
3. The Bipolaris sorghicola toxins can reduce the permeability of cell membrane
The seeds of L407B and Tx622B were treated by different concentrations of Bipolarissorghicola crude toxins.And the radicel growth inhibition rate as well as the electricalconductivity and malondialdehyde(MDA) content were determined to studied the effect ofcrude toxins of Bipolaris sorghicola on growth and embrance permeability of differentvarieties of sorghum radicle. The results showed that crude toxin could obviously inhibitgrowth of sorghum radicel. The electrical conductivity and MDA content were all higher inthe group which was treated by Bipolaris sorghicola crude toxins with same concentrationtwo days before than in control group. The electrical conductivity and MDA content werealso higher in the group which was treated by Bipolaris sorghicola crude toxins with differentconcentration on the same treated time. This demonstrates that Bipolaris sorghicola crudetoxins can reduce membrane permeability of sorghum radicle and there is a significantdifference among the different varieties of sorghum.
4. The Bipolaris sorghicola toxins of pathogen can inhibit host protective enzyme activity
The method of chromometry and Ultraviolet-visible spectrophotometer were used todetect the effecte of Bipolaris sorghicola toxins on the activity of protective enzymes such asperoxidase, superoxide dismutase, phenylalnine ammonialyase and polyphenoloxidase insorghum radicle. The results showed that the activities of peroxidase, superoxide dismutase,phenylalnine ammonialyase and polyphenoloxidase were significantly reduced in thesorghum radicle after Bipolaris sorghicola toxins treatmemt. The results indicated that theBipolaris sorghicola toxins inhibited the activities of protective enzymes.
5. The structure and function of PSII in sorghum leave will be damaged when pathogeninfection
Using sorghum variety Tx622B as materials, the effects of the infection of Bipolarissorghicola on photosystem II (PSII) structure and function of sorghum leaves were studiedby determining the chlorophyll content, chlorophyll fluorescence transient, hydrogen peroxide(H2O2)content and malondialdehyde (MDA) content in leaves of sorghum.The resultsshowed that the infected by B. sorghicola induced increase in MDA content of sorghumLeaves along with enhanced lipid peroxidation and chlorophyll fluorescence transient inLeaves of sorghum appeared with some difference. Further JIP-test analysis showed that thepathogen infection seriously hurt the donor side of PSII, blocking the QAto QBelectrontransfer, reducing the activity of reaction center.
6. The resistant gene to target spot leaf was controlled by recessive monogene
The recombinat inbred lines population which was constructed by the descendent ofL407B (immune inbred line) and Tx622B (susceptible inbred line),as materials accompaniedby the method of quantitative heredity,was used to study the heredity of resistance to thedisease.According to the resistant analysis of the field investigation and X2examination,proved that the genetic mechanism was in accordant with Mendelian monogenesegregation law.Appearances in F1families was consistent with Tx622B. In F2families,resistant plants is one quarter of all plants.To verify the resistant character of L407B wascontrolled by recessive monogene preliminarily.
7. The site of resistant gene on chromosome
Screened out of SSR markers which132"Xtxp-" and230"sm-" at the beginningdistribute on the whole sorghum chromosome.These markers showed polymorphic betweenresistant and susceptible parents and consistency between resistant parent and resistant-pool ofF2generation.By chromosome walking, one resistant gene to target spot leaf was mappedbetween Xtxp303and sm05014on the short arm of chromosome5within5.73M-8.15M.
1.高粱靶斑病病原菌的生物学特性
生物学特性研究结果表明：病菌菌丝生长的适宜温度为25～30℃，分生孢子萌发的适宜温度为20～30℃。分生孢子萌发的适宜pH为3～6，偏酸性。大多糖类均可作为该病的碳源营养，如半乳糖和乳糖，而山梨糖对菌丝生长不太适宜。不同光照处理对病菌菌丝生长的影响不明显。
2.我国高粱靶斑病病原菌种类及种群多样性
从采自我国不同地区的感染高粱靶斑病病叶中分离出病原菌，通过形态学观察和rDNA-ITS序列分析进行多样性研究。结果表明，虽然这些发病地区病叶上的病斑差异很大，但病原菌在菌落形态和孢子形态方面却极为相似;通过这些菌株的rDNA-ITS序列聚类分析表明，它们与Bipolaris sorghicola相似度达99%,从而证实虽然不同地区病斑差异较大，但都是同一病原菌引起的病害。
3.高粱靶斑病菌粗毒素能降低细胞膜透性
采用测定不同浓度病原菌粗毒素处理的高粱种子L407B和Tx622B的胚根生长抑制率以及电导率和丙二醛(MDA)含量的方法，探讨高粱靶斑病菌粗毒素对高粱不同品种胚根生长及胚根细胞膜透性的影响。结果表明，病菌粗毒素对高粱胚根生长有明显的抑制作用。在同一毒素浓度下，处理后2d，处理组胚根的相对电导率和MDA含量均高于对照组；同一处理时间内，不同浓度毒素处理后胚根相对电导率和MDA含量均高于对照组。说明高粱靶斑病菌毒素能降低高粱胚根细胞膜透性，且不同高粱品种间差异显著。
4.高粱靶斑病菌粗毒素能抑制寄主保护酶活性
利用紫外可见分光光度计和比色法测定了不同浓度病原菌粗毒素处理的高粱种子L407B和Tx622B胚根保护酶（过氧化物酶、超氧化物歧化酶、多酚氧化酶和苯丙氨酸解氨酶）活性。结果表明，毒素处理后，高粱种子胚根过氧化物酶、超氧化物歧化酶、多酚氧化酶和苯丙氨酸解氨酶活性明显降低，说明毒素对寄主保护酶起抑制作用。
5.高粱靶斑病病原菌侵染会破坏高粱叶片PSII的结构和功能
以高粱品种Tx622B为试验材料,通过同时测定高粱叶片叶绿素含量、叶绿素荧光快速诱导动力学曲线，以及过氧化氢(H2O2)含量和丙二醛(MDA)含量的变化，来研究高粱靶斑病菌[Bipolaris sorghicola (Lefebvre&Sherwin) Alcorn]侵染高粱叶片后对光系统II(PSII)结构和功能的影响。研究结果表明，受病原菌侵染的高粱叶片,H2O2和MDA含量均升高,膜脂过氧化的程度加剧；快速叶绿素荧光诱导动力学曲线发生明显变化，进一步的JIP测试分析表明病原菌侵染严重伤害了PSII供体侧、阻断了受体侧QA到QB的电子传递，从而使反应中心的活性降低。
6.抗高粱靶斑病基因为隐性单基因
以高粱靶斑病抗病自交系L407B和感病自交系Tx622B的杂交后代构建的重组自交系群体为材料，利用数量遗传分析方法研究了L407B的抗病遗传规律。对高粱靶斑病田间性状调查结果进行分析，经卡方分析检验符合孟德尔单基因控制的遗传分离规律。F1代田间表现与亲本Tx622B完全相同，在F2中，抗病植株占F2代总植株的四分之一，初步证实自交系L407B的抗病性是受一对隐性基因控制的质量性状。
7.抗病基因在染色体上的位置
从覆盖高粱染色体组的132对以“Xtxp-”开头的SSR引物和230对以“sm-”开头的SSR引物中筛选出在抗感两亲本之间有差异且在抗性亲本和F2代抗池间无差异的引物。通过染色体步移方法，将高粱抗靶斑病基因定位于标记Xtmp303和sm05014之间，即该抗病基因位于第5条染色体短臂上5.73M—8.15M内。
Sorghum is important as food, wine, feed and industrial raw materials. Sorghum infectedby pathogens can lead to poor food quality and lower production.At the same time, pathogenscan produce mycotoxin to harm humans and animals. Target leaf spot which casued byBipolaris sorghicola is one of important foliar disease in sorghum varieties. At present, thereis no report on pathopoiesis mechanism as well as inheritance mechanism and moleculargenetics of this disease in China. This paper focuses on the pathogenesis and location ofresistance gene against target leaf spot in sorghum. There was a relatively systematic study inthe biological characteristics and population diversity of pathogen, the effect of pathogencrude toxins on seed germination and root growth as well as cell membrane permeability anddefense enzyme activity in sorghum radicle, the changes of sorghum leaf photosyntheticindexes and photosystem Ⅱa fter pathogen infection, the genetic law of resistance to thedisease and the marker and location of resistance gene and many other aspects. The studyobtained ideal results, parts of them filled in the blank about this research in China. Theresults provided scientific basis for breeding of resistant varieties and disease management.
The main results were summarized as follows:
1. Biological characteristics of pathogen which caused target leaf spot on sorghum
The result indicated that optimum temperature for pathogen mycelia growth was25-30℃, and that for conidiagermination was20-30℃.The optimum pH for conidiagermination was-6, leaning to acidity. The pathogen could use many kind of sugar as acarbon nutrient，eg.lactose and galactose. However, sorbose was not fit for pathogen myceliagrowth. Different light condition for mycelium growth was not obvious.
2. The category and population diversity of pathogen in China
In this diversity research, we isolated pathogens from infected target leaf spot diseaseleaves of different regions of China by morphological observation and rDNA-ITS sequenceanalysis.The results show that although these lesions have significant differences, the colonymorphology and spore morphology of pathogens are very similar. The rDNA-ITS sequence cluster analysis showed that the similarity of these pathogens with Bipolaris sorghicola reachto99%.All of these confirmed this disease caused by the same pathogen,although theselesions have significant differences.
3. The Bipolaris sorghicola toxins can reduce the permeability of cell membrane
The seeds of L407B and Tx622B were treated by different concentrations of Bipolarissorghicola crude toxins.And the radicel growth inhibition rate as well as the electricalconductivity and malondialdehyde(MDA) content were determined to studied the effect ofcrude toxins of Bipolaris sorghicola on growth and embrance permeability of differentvarieties of sorghum radicle. The results showed that crude toxin could obviously inhibitgrowth of sorghum radicel. The electrical conductivity and MDA content were all higher inthe group which was treated by Bipolaris sorghicola crude toxins with same concentrationtwo days before than in control group. The electrical conductivity and MDA content werealso higher in the group which was treated by Bipolaris sorghicola crude toxins with differentconcentration on the same treated time. This demonstrates that Bipolaris sorghicola crudetoxins can reduce membrane permeability of sorghum radicle and there is a significantdifference among the different varieties of sorghum.
4. The Bipolaris sorghicola toxins of pathogen can inhibit host protective enzyme activity
The method of chromometry and Ultraviolet-visible spectrophotometer were used todetect the effecte of Bipolaris sorghicola toxins on the activity of protective enzymes such asperoxidase, superoxide dismutase, phenylalnine ammonialyase and polyphenoloxidase insorghum radicle. The results showed that the activities of peroxidase, superoxide dismutase,phenylalnine ammonialyase and polyphenoloxidase were significantly reduced in thesorghum radicle after Bipolaris sorghicola toxins treatmemt. The results indicated that theBipolaris sorghicola toxins inhibited the activities of protective enzymes.
5. The structure and function of PSII in sorghum leave will be damaged when pathogeninfection
Using sorghum variety Tx622B as materials, the effects of the infection of Bipolarissorghicola on photosystem II (PSII) structure and function of sorghum leaves were studiedby determining the chlorophyll content, chlorophyll fluorescence transient, hydrogen peroxide(H2O2)content and malondialdehyde (MDA) content in leaves of sorghum.The resultsshowed that the infected by B. sorghicola induced increase in MDA content of sorghumLeaves along with enhanced lipid peroxidation and chlorophyll fluorescence transient inLeaves of sorghum appeared with some difference. Further JIP-test analysis showed that thepathogen infection seriously hurt the donor side of PSII, blocking the QAto QBelectrontransfer, reducing the activity of reaction center.
6. The resistant gene to target spot leaf was controlled by recessive monogene
The recombinat inbred lines population which was constructed by the descendent ofL407B (immune inbred line) and Tx622B (susceptible inbred line),as materials accompaniedby the method of quantitative heredity,was used to study the heredity of resistance to thedisease.According to the resistant analysis of the field investigation and X2examination,proved that the genetic mechanism was in accordant with Mendelian monogenesegregation law.Appearances in F1families was consistent with Tx622B. In F2families,resistant plants is one quarter of all plants.To verify the resistant character of L407B wascontrolled by recessive monogene preliminarily.
7. The site of resistant gene on chromosome
Screened out of SSR markers which132"Xtxp-" and230"sm-" at the beginningdistribute on the whole sorghum chromosome.These markers showed polymorphic betweenresistant and susceptible parents and consistency between resistant parent and resistant-pool ofF2generation.By chromosome walking, one resistant gene to target spot leaf was mappedbetween Xtxp303and sm05014on the short arm of chromosome5within5.73M-8.15M.
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