外源茉莉酸影响马铃薯对晚疫病抗性的初步研究
摘要
马铃薯晚疫病是危害马铃薯生产的重要病害,其病原菌(Phytophthora infestans (Mont.) De Bary)变异快且容易发生流行,给病害研究和防治技术研发带来困难。目前就其病原菌及其毒素的致病机理研究较少,缺乏对现有栽培品种抗性诱导相关的研究工作。本研究以云南省目前主要栽培的米拉、合作88号和滇薯6号等马铃薯品种及来自云南的ZY15、LSX18和XH05-5-4晚疫病菌为试验材料,较为系统地研究了马铃薯晚疫病病原菌致病作用因子及马铃薯—病原菌互作中外源JA对马铃薯抗性的影响,得到以下结果：
1.用液体培养基培养P. infestan 30 d,过滤获得粗毒素,硫酸铵沉淀获得精毒素,进行葡聚糖G-75分离和SDS聚丙烯酰胺凝胶电泳,并进行酶活性和生物活性测定、叶片超微结构分析,试验结果表明：(1)P. infestans及其毒素都可以毒害马铃薯,其毒素是由多种蛋白质组成的群体,分子量在20.709kDa-71.763kDa之间,等电点4.32-9.10之间,对寄主有非专化性毒害作用,稀释200倍也能引起毒害症状发生。毒素的毒力由P. infestans生理小种所决定,即小种不同产生的毒素其毒力也不同。马铃薯对毒素的抵抗能力因品种抗性不同而异,具有高抗晚疫病的地上部分植株或块茎对毒素的抵抗能力高于水平抗性品种和感病品种。(2) P.infestans及其毒素影响马铃薯叶片的光合作用,并破坏马铃薯叶片细胞膜和叶绿体,导致叶片坏死。
2.系统进行了茉莉酸(JA)对P. infestans抑制作用的测定、JA处理马铃薯植株的抗性评价、抗性相关酶(PAL、CAT和POD)活性测定等研究,试验结果表明：(1)JA存在于马铃薯叶片中,含量大小因品种而异,滇薯6号叶片中JA含量高于合作88号和米拉；外源JA引起马铃薯植株内源JA的改变,但变化幅度随马铃薯品种和外源JA的浓度而异。同时,外源茉莉酸对晚疫病菌有直接的抑制作用。(2)外源JA能够诱导马铃薯对晚疫病菌产生抵抗作用,不同品种、同一品种地上部和块茎对外源JA的浓度反应也不同,滇薯6号地上部经100μmol·L-1 JA处理抗性最好,块茎经200μmol·L-1 JA处理抗性最好；米拉地上部经150μmol·L-1 JA处理抗性最好,块茎是200μmol·L-1 JA处理抗性最好；合作88号地上部和块茎都是经150μmol·L-1 JA处理抗性最好。JA浓度的高低与马铃薯植株的抗性不呈相关性,地上部表现出了抗性的提高,而块茎的抗性未必提高。因此,可以利用不同浓度JA诱导不同品种的抗性。在大田将JA做为诱导剂,在马铃薯的不同生长时期喷施,提高马铃薯对晚疫病的抗性,从而达到防治病害的目的。(3)外源JA能够增强马铃薯叶片中抗性相关酶的活性。马铃薯品种不同,酶活性变化不同,酶活性强度与病害发生程度呈负相关关系。外源JA可以使米拉叶片中PAL活性显著增加,使合作88号和滇薯6号叶片中的POD活性显著增强。推测JA作为信号分子启动了植株防御酶活性来抵抗病原菌的入侵。(4) P.infestans在侵染马铃薯时将受到寄主不同程度的抵抗,马铃薯品种抗性不同其抵抗的能力不同。供试的3个品种中,滇薯6号具有较好的抵抗晚疫病菌侵染的能力,其气孔数量少,减少了病原菌入侵的几率,从而抑制了入侵能力和入侵速度。合作88号次之,而米拉气孔数量多,有利于病原菌的入侵。外源喷施JA后在马铃薯的细胞超微结构方面未检测到相应抗性结构的形成,但细胞壁明显加厚,叶绿体中出现空洞状结构。
3.2-D PAGE电泳技术可以用于分离马铃薯叶片的蛋白质,外源JA的使用改变了马铃薯叶片中部分蛋白质的组成,米拉叶片蛋白点遗失3个,新增特异蛋白点5个,上调在2倍以上蛋白点8个,下调在2倍以上蛋白点3个。合作88号叶片出现特异蛋白点1个,上调2倍以上蛋白点13个,下调2倍以下蛋白点8个。滇薯6号未出现特异蛋白点,上调蛋白点10个,下调蛋白点10个,遗失蛋白点10个。进一步对米拉叶片JA诱导蛋白质进行质谱鉴定,结果表明：(1)特异蛋白点5个中有3个与半胱氨酸蛋白酶抑制剂同源；1个是未命名蛋白；另1个与苏氨酸脱氨酶同源。(2)下调点3个中2个与核酮糖-1,5-二磷酸羧化酶/加氧酶(Rubisco)大亚基同源。(3)上调点8个中有3个为假设蛋白,匹配得分分别是47、70、56,均未达到有效得分；1个与钙结合蛋白匹配得分为72；1个与Rubisco小亚基3匹配得分是79；1个与液泡H+-ATPase B亚基匹配得分170,其中有39%氨基酸序列匹配,可能参与Ca2+信号转导过程,进而参与植物生长发育和响应逆境胁迫过程。(4)遗失蛋白3个,分别与组氨酸激酶、乙醇脱氢酶和硫氧还蛋白过氧化物酶同源。
总之,外源茉莉酸对马铃薯抗晚疫病的影响表现在以下两个方面。首先,茉莉酸对晚疫病菌有直接的抑制作用。其次,外源茉莉酸作为信号分子启动了抗性相关酶的合成,如诱导产生半胱氨酸蛋白酶抑制剂和苏氨酸脱氨酶和一些与光合作用相关的钙结合蛋白。因此,外源茉莉酸通过抑制病原菌和提高马铃薯自身的生命活力来表现抗病性。
Potato late blight is one of the most harmful diseases in the world. The P.infestans variation and disease epidemic occur frequently, bringing difficulty to its research and the development of integrated control technique. At present, less attention is focused on pathogenesis mechanism of P.infestans and its toxins, especially the induced resistance of current potato varieties. In this paper, three potato varieties(Mira, S88 and PB06) and three isolates(ZY15,LSX18 and XH05-5-4)of P.infestans from Yunnan province were employed as materials, the pathogenic factor of P.infestans and the effects of exogenous jasmonic acid on potato resistance to late blight have been systematically studied. The results are as follows:
1. The P.infestans was cultured in the liquid rye medium for 30 d, crude toxin was obtained after filtration, refined toxin was obtained by (NH4)2SO4 precipitation. The refined toxin was further purified by glucan G-75 chromatography column and SDS polyacrylamide gel electrophoresis. And then, enzyme activity and bioactivity were tested, and the ultrastructure of potato leaf was observed. The results indicate:(1) Both P.infestans and its toxins can be toxic to damage potato. The toxins are some enzymes, its molecular weights is from 20.709kDa to 71.763kDa. It has non-specific poisoning effect on the host, the symptoms still appeared by toxin even after diluted for 200 times. The toxicity is determined by the physiological race of P.infestans, indicating different physiological race may have different toxicity. Different potato varieties have different resistance to toxins, vertical resistance variety is more resistant than horizontal resistant variety and susceptible variety. (2) The photosynthesis and respiration of potato leaf were affected by P.infestans and its toxins, membrane and chloroplast of potato leaf were destroyed, causing eventual leaf death.
2. The inhibitory effect of JA to P.infestans was tested, including the resistance evaluation of potato under JA treatment, the activity measurement of PAL, CAT and POD. The results show:(1) Naturally occurring JA was found in potato leaf, the JA content in leaf were different in different potato varieties. The JA content of PB06 was higher than S88 or Mira. While the exogenous JA changed the JA content in potato leaf, but the change range varied with potato varieties and JA treatment concentrations. Meanwhile, JA was found to have immediate inhibitory effect on P.infestans. (2) Resistance of potato to late blight was induced by exogenous JA, while the resistant ability was affected by JA concentrations, potato variety and the organ of the plants. In PB06, JA treatment at 100μmol·L-1 and 200μmol·L-1 were found to be the most effective concentrations for upper part and the tuber respectively. In Mira, the most effective concentrations for upper part and the tuber were 150μmol·L-1 and 200μmol·L-1 respectively. The optimal JA treatment concentration for both the upper part and the tuber of S88 is 150μmol·L-1. The JA content was not correlated to resistance of potato, only the upper part of plant showed increased resistance while the tuber did not show any resistance improvement. Therefore, different JA content will be used to induce resistance for different potato varieties. JA can be used as an inducer when sprayed JA on potato plants in field, inducing resistance of potato to late blight. (3) Activities of resistance related enzymes in potato leaf were enhanced by exogenous JA. The change in enzyme activity varied with potato varieties. There was a negative correlation between enzyme activity and degree of the late blight occurrence. The PAL activity of Mira leaf was enhanced by exogenous JA, both the POD activities of PB06 and S88 were also increased by JA. It was speculated that exogenous JA served as a signal molecule activating the defense enzymes against P.infestans invasion. (4) P.infestansw faced resistance somehow when it invaded the host, depending on potato resistance ability. The resistance ability of PB06 was better than S88 and Mira, because it has less stomatic numbers. No special resistance structure in potato leaf was observed after exogenous JA treatment, however the cell wall was found to be thickened and some small cavities in the chloroplast were discovered under electronic microscope.
3. The proteins in potato leaf were separated by 2-D PAGE, protein composition of potato leaf were partly changed by exogenous JA. Three proteins of Mira were lost, while five new specific proteins,8 up-regulated proteins and 3 down-regulated proteins were found in Mira leaf. One new specific protein,13 up-regulated proteins and 8 down-regulated proteins were found in S88 leaf. No new specific protein was found in PB06 leaf, but 10 up-regulated proteins and 10 down-regulated proteins were found, and ten proteins were lost. The proteins induced by exogenous JA in Mira leaf were identified by mass spectrometry. The results show:(1)Three from five new specific proteins were found to be cysteine proteinase inhibitors, the rest two proteins were a threonine deaminase and unknown protein. (2)Two down-regulated proteins were found to be ribulose-1,5- bisphosphate carboxylase/oxygenase(Rubisco) large subunit. (3)Three up-regulated proteins were found to be hypothetical proteins, computer match score is respectively 47,70 and 56. One protein has a match score of 72 for calcium binding protein. Another one protein has a match score of 79 for Rubisco small subunit 3 in the photosynthetic and respiration. One protein matches H+-ATPase B with a score of 170, the amino acids sequence homology was 39%. The protein may have function in signal transduction process of Ca+ and play further roles in plant growth and development or the response to stresses. (4) Histidine kinase, alcohol dehydrogenase and thioredoxin peroxidase were found lost after exogenous JA treatment.
In conclusion, exogenous JA has two main effects on potato resistance to P. infestans. Firstly, exogenous JA directly inhibits P.infestans. Secondly, enzyme synthesis of resistance related enzymes is stimulated by exogenous JA as a signal molecule. These enzymes include the cysteine proteinase inhibitor, threonine deaminase and calcium binding proteins for photosynthesis. Therefore, potato shows resistance under exogenous JA by inhibiting P. infestans and improving the life vigor of itself.
1.用液体培养基培养P. infestan 30 d,过滤获得粗毒素,硫酸铵沉淀获得精毒素,进行葡聚糖G-75分离和SDS聚丙烯酰胺凝胶电泳,并进行酶活性和生物活性测定、叶片超微结构分析,试验结果表明：(1)P. infestans及其毒素都可以毒害马铃薯,其毒素是由多种蛋白质组成的群体,分子量在20.709kDa-71.763kDa之间,等电点4.32-9.10之间,对寄主有非专化性毒害作用,稀释200倍也能引起毒害症状发生。毒素的毒力由P. infestans生理小种所决定,即小种不同产生的毒素其毒力也不同。马铃薯对毒素的抵抗能力因品种抗性不同而异,具有高抗晚疫病的地上部分植株或块茎对毒素的抵抗能力高于水平抗性品种和感病品种。(2) P.infestans及其毒素影响马铃薯叶片的光合作用,并破坏马铃薯叶片细胞膜和叶绿体,导致叶片坏死。
2.系统进行了茉莉酸(JA)对P. infestans抑制作用的测定、JA处理马铃薯植株的抗性评价、抗性相关酶(PAL、CAT和POD)活性测定等研究,试验结果表明：(1)JA存在于马铃薯叶片中,含量大小因品种而异,滇薯6号叶片中JA含量高于合作88号和米拉；外源JA引起马铃薯植株内源JA的改变,但变化幅度随马铃薯品种和外源JA的浓度而异。同时,外源茉莉酸对晚疫病菌有直接的抑制作用。(2)外源JA能够诱导马铃薯对晚疫病菌产生抵抗作用,不同品种、同一品种地上部和块茎对外源JA的浓度反应也不同,滇薯6号地上部经100μmol·L-1 JA处理抗性最好,块茎经200μmol·L-1 JA处理抗性最好；米拉地上部经150μmol·L-1 JA处理抗性最好,块茎是200μmol·L-1 JA处理抗性最好；合作88号地上部和块茎都是经150μmol·L-1 JA处理抗性最好。JA浓度的高低与马铃薯植株的抗性不呈相关性,地上部表现出了抗性的提高,而块茎的抗性未必提高。因此,可以利用不同浓度JA诱导不同品种的抗性。在大田将JA做为诱导剂,在马铃薯的不同生长时期喷施,提高马铃薯对晚疫病的抗性,从而达到防治病害的目的。(3)外源JA能够增强马铃薯叶片中抗性相关酶的活性。马铃薯品种不同,酶活性变化不同,酶活性强度与病害发生程度呈负相关关系。外源JA可以使米拉叶片中PAL活性显著增加,使合作88号和滇薯6号叶片中的POD活性显著增强。推测JA作为信号分子启动了植株防御酶活性来抵抗病原菌的入侵。(4) P.infestans在侵染马铃薯时将受到寄主不同程度的抵抗,马铃薯品种抗性不同其抵抗的能力不同。供试的3个品种中,滇薯6号具有较好的抵抗晚疫病菌侵染的能力,其气孔数量少,减少了病原菌入侵的几率,从而抑制了入侵能力和入侵速度。合作88号次之,而米拉气孔数量多,有利于病原菌的入侵。外源喷施JA后在马铃薯的细胞超微结构方面未检测到相应抗性结构的形成,但细胞壁明显加厚,叶绿体中出现空洞状结构。
3.2-D PAGE电泳技术可以用于分离马铃薯叶片的蛋白质,外源JA的使用改变了马铃薯叶片中部分蛋白质的组成,米拉叶片蛋白点遗失3个,新增特异蛋白点5个,上调在2倍以上蛋白点8个,下调在2倍以上蛋白点3个。合作88号叶片出现特异蛋白点1个,上调2倍以上蛋白点13个,下调2倍以下蛋白点8个。滇薯6号未出现特异蛋白点,上调蛋白点10个,下调蛋白点10个,遗失蛋白点10个。进一步对米拉叶片JA诱导蛋白质进行质谱鉴定,结果表明：(1)特异蛋白点5个中有3个与半胱氨酸蛋白酶抑制剂同源；1个是未命名蛋白；另1个与苏氨酸脱氨酶同源。(2)下调点3个中2个与核酮糖-1,5-二磷酸羧化酶/加氧酶(Rubisco)大亚基同源。(3)上调点8个中有3个为假设蛋白,匹配得分分别是47、70、56,均未达到有效得分；1个与钙结合蛋白匹配得分为72；1个与Rubisco小亚基3匹配得分是79；1个与液泡H+-ATPase B亚基匹配得分170,其中有39%氨基酸序列匹配,可能参与Ca2+信号转导过程,进而参与植物生长发育和响应逆境胁迫过程。(4)遗失蛋白3个,分别与组氨酸激酶、乙醇脱氢酶和硫氧还蛋白过氧化物酶同源。
总之,外源茉莉酸对马铃薯抗晚疫病的影响表现在以下两个方面。首先,茉莉酸对晚疫病菌有直接的抑制作用。其次,外源茉莉酸作为信号分子启动了抗性相关酶的合成,如诱导产生半胱氨酸蛋白酶抑制剂和苏氨酸脱氨酶和一些与光合作用相关的钙结合蛋白。因此,外源茉莉酸通过抑制病原菌和提高马铃薯自身的生命活力来表现抗病性。
Potato late blight is one of the most harmful diseases in the world. The P.infestans variation and disease epidemic occur frequently, bringing difficulty to its research and the development of integrated control technique. At present, less attention is focused on pathogenesis mechanism of P.infestans and its toxins, especially the induced resistance of current potato varieties. In this paper, three potato varieties(Mira, S88 and PB06) and three isolates(ZY15,LSX18 and XH05-5-4)of P.infestans from Yunnan province were employed as materials, the pathogenic factor of P.infestans and the effects of exogenous jasmonic acid on potato resistance to late blight have been systematically studied. The results are as follows:
1. The P.infestans was cultured in the liquid rye medium for 30 d, crude toxin was obtained after filtration, refined toxin was obtained by (NH4)2SO4 precipitation. The refined toxin was further purified by glucan G-75 chromatography column and SDS polyacrylamide gel electrophoresis. And then, enzyme activity and bioactivity were tested, and the ultrastructure of potato leaf was observed. The results indicate:(1) Both P.infestans and its toxins can be toxic to damage potato. The toxins are some enzymes, its molecular weights is from 20.709kDa to 71.763kDa. It has non-specific poisoning effect on the host, the symptoms still appeared by toxin even after diluted for 200 times. The toxicity is determined by the physiological race of P.infestans, indicating different physiological race may have different toxicity. Different potato varieties have different resistance to toxins, vertical resistance variety is more resistant than horizontal resistant variety and susceptible variety. (2) The photosynthesis and respiration of potato leaf were affected by P.infestans and its toxins, membrane and chloroplast of potato leaf were destroyed, causing eventual leaf death.
2. The inhibitory effect of JA to P.infestans was tested, including the resistance evaluation of potato under JA treatment, the activity measurement of PAL, CAT and POD. The results show:(1) Naturally occurring JA was found in potato leaf, the JA content in leaf were different in different potato varieties. The JA content of PB06 was higher than S88 or Mira. While the exogenous JA changed the JA content in potato leaf, but the change range varied with potato varieties and JA treatment concentrations. Meanwhile, JA was found to have immediate inhibitory effect on P.infestans. (2) Resistance of potato to late blight was induced by exogenous JA, while the resistant ability was affected by JA concentrations, potato variety and the organ of the plants. In PB06, JA treatment at 100μmol·L-1 and 200μmol·L-1 were found to be the most effective concentrations for upper part and the tuber respectively. In Mira, the most effective concentrations for upper part and the tuber were 150μmol·L-1 and 200μmol·L-1 respectively. The optimal JA treatment concentration for both the upper part and the tuber of S88 is 150μmol·L-1. The JA content was not correlated to resistance of potato, only the upper part of plant showed increased resistance while the tuber did not show any resistance improvement. Therefore, different JA content will be used to induce resistance for different potato varieties. JA can be used as an inducer when sprayed JA on potato plants in field, inducing resistance of potato to late blight. (3) Activities of resistance related enzymes in potato leaf were enhanced by exogenous JA. The change in enzyme activity varied with potato varieties. There was a negative correlation between enzyme activity and degree of the late blight occurrence. The PAL activity of Mira leaf was enhanced by exogenous JA, both the POD activities of PB06 and S88 were also increased by JA. It was speculated that exogenous JA served as a signal molecule activating the defense enzymes against P.infestans invasion. (4) P.infestansw faced resistance somehow when it invaded the host, depending on potato resistance ability. The resistance ability of PB06 was better than S88 and Mira, because it has less stomatic numbers. No special resistance structure in potato leaf was observed after exogenous JA treatment, however the cell wall was found to be thickened and some small cavities in the chloroplast were discovered under electronic microscope.
3. The proteins in potato leaf were separated by 2-D PAGE, protein composition of potato leaf were partly changed by exogenous JA. Three proteins of Mira were lost, while five new specific proteins,8 up-regulated proteins and 3 down-regulated proteins were found in Mira leaf. One new specific protein,13 up-regulated proteins and 8 down-regulated proteins were found in S88 leaf. No new specific protein was found in PB06 leaf, but 10 up-regulated proteins and 10 down-regulated proteins were found, and ten proteins were lost. The proteins induced by exogenous JA in Mira leaf were identified by mass spectrometry. The results show:(1)Three from five new specific proteins were found to be cysteine proteinase inhibitors, the rest two proteins were a threonine deaminase and unknown protein. (2)Two down-regulated proteins were found to be ribulose-1,5- bisphosphate carboxylase/oxygenase(Rubisco) large subunit. (3)Three up-regulated proteins were found to be hypothetical proteins, computer match score is respectively 47,70 and 56. One protein has a match score of 72 for calcium binding protein. Another one protein has a match score of 79 for Rubisco small subunit 3 in the photosynthetic and respiration. One protein matches H+-ATPase B with a score of 170, the amino acids sequence homology was 39%. The protein may have function in signal transduction process of Ca+ and play further roles in plant growth and development or the response to stresses. (4) Histidine kinase, alcohol dehydrogenase and thioredoxin peroxidase were found lost after exogenous JA treatment.
In conclusion, exogenous JA has two main effects on potato resistance to P. infestans. Firstly, exogenous JA directly inhibits P.infestans. Secondly, enzyme synthesis of resistance related enzymes is stimulated by exogenous JA as a signal molecule. These enzymes include the cysteine proteinase inhibitor, threonine deaminase and calcium binding proteins for photosynthesis. Therefore, potato shows resistance under exogenous JA by inhibiting P. infestans and improving the life vigor of itself.
引文
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