灰葡萄孢菌激活蛋白的纯化、基因克隆与功能研究
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摘要
激活蛋白是一类从真菌中分离的、具有激发子功能的新型蛋白质,可诱导多种植物产生系统抗性,促进植物生长,改善作物品质。为了进一步发掘具有我国自主知识产权的新型激活蛋白,本研究从灰葡萄孢菌(Botrytis cinerea)中分离获得了纯化的激活蛋白,确定了该激活蛋白在诱导植物抗病性和抗旱性方面的生物功能;克隆并获得了灰葡萄孢菌激活蛋白基因,分别实现了在大肠杆菌(Escherichia coli)和毕赤酵母(Pichia pastoris)中的表达,确定了表达蛋白的生物功能。该研究为利用工程菌株获得大量激活蛋白,研究并分析激活蛋白结构与功能的关系,从而为蛋白药物的分子设计奠定了基础。主要结果如下:
通过沸水浴、离心、阴离子交换层析等方法,从灰葡萄孢菌BC-4-2-2-1菌株的菌丝体中分离纯化出一种激活蛋白,命名为PebC1(protein elicitor botrytis cinerea1)。经SDS-PAGE银染显示该蛋白呈单一条带,相对分子量约为36kD,等电点为4.85。
用胶内酶解的方法,采用MALDI-TOF/TOF等质谱技术对纯化的激活蛋白PebC1进行了肽段序列测定,获得3个重复性较好的可靠肽段。在NCBI上进行比对发现,该蛋白与来源于灰葡萄孢菌的蛋白XP_001551609.1匹配率达95%。
功能研究表明,灰葡萄孢菌激活蛋白PebC1能显著促进小麦幼苗生长、提高小麦抗旱性、诱导番茄对灰霉病的系统抗性。不同浓度激活蛋白浸泡小麦种子,11天株高均高于对照,其中以5μg/mL激活蛋白对小麦生长的促进作用最明显;激活蛋白处理的小麦根系活力比对照增加1.29倍,小麦幼苗抗旱综合系数也从36.53提高到57.08,说明激活蛋白增强小麦的抗旱性可能与增加小麦根系活力有一定的关系;用不同浓度的激活蛋白浸泡番茄种子,幼苗生长45天接种灰葡萄孢菌孢子悬浮液(浓度为10~(4)个孢子/mL),接种后14天番茄的病情指数明显低于对照,对灰霉病的诱抗效果达到31.95%～69.19%,其中10μg/mL激活蛋白处理对番茄的诱抗效果最为明显,接种后21天的诱抗效果仍达66.10%,说明灰葡萄孢菌激活蛋白PebC1不仅能显著增强番茄对灰霉病的抗性,而且诱抗效果持续时间较长。
测定了PebC1处理后的番茄体内苯丙氨酸解氨酶(PAL)、过氧化物酶(POD)和多酚氧化酶(PPO)活性的动态变化。经激活蛋白PebC1处理后,番茄幼苗叶片的PAL、POD和PPO活性均有不同程度的提高,PAL活性在诱导后24小时达最高,较对照增加46.84%;POD活性在诱导72小时出现活性高峰,较对照增加109.5%;PPO活性在72小时出现峰值,比对照增加111.0%。PAL、POD和PPO均是与植物抗病相关的防御酶,番茄防御相关酶活性的提高可能是激活蛋白诱导番茄植株抗灰霉病的主要生理机制之一,该研究为揭示激活蛋白诱导植物抗病性的作用机理提供了理论依据。
通过反向遗传学的方法,获得了灰葡萄孢菌激活蛋白PebC1基因的cDNA序列,它与来源于灰葡萄孢菌的基因XM_001551559完全一致。该基因编码的蛋白属于灰葡萄孢菌的alpha-NAC蛋白(nascent polypeptide-associated complex alpha polypeptide),它含有保守的NAC结构域和UBA结构域,与来源于极细链格孢菌(Alternaria tenuissima)的激活蛋白PeaT1基因的结构域相同,另外,PebC1和PeaT1二者在编码的氨基酸组成上也有79.4%的相似性,其中完全一致的氨基酸占71.0%。PebC1基因全长639bp,编码212个氨基酸,预测分子量为23080.2,等电点为4.57。亲水性分析发现该蛋白属于亲水性蛋白。
进行了PebC1基因的原核表达。以原核表达系统中的pET-28a(+)为表达载体,将重组载体转入大肠杆菌Rossetta(DE3)菌株,通过IPTG诱导获得了PebC1基因的表达产物,利用(?)KTA explorer10蛋白纯化仪进行重组蛋白的纯化,选用Hitrap chelating Column柱进行亲和层析,得到了纯化的重组蛋白,并应用质谱分析鉴定了该基因表达的蛋白质为灰葡萄孢菌激活蛋白PebC1。功能测定结果表明,纯化的重组蛋白能促进番茄幼苗的生长,增强小麦的抗旱性。
构建了PebC1毕赤酵母表达载体,获得了具有生物活性的分泌表达蛋白。用PCR法从灰葡萄孢菌cDNA中扩增获得PebC1的编码序列并亚克隆到毕赤酵母表达系统的表达载体pPIC9K上,得到重组质粒pPIC9K/PebC1。重组质粒经BglⅡ线性化后电击导入毕赤酵母GS115中,采用MD、G418-YPD平板和PCR法进行筛选,获得了分泌表达的重组毕赤酵母菌株。用甲醇诱导PebC1表达,经SDS-PAGE检测证明,PebC1基因在毕赤酵母中成功分泌表达。生物活性试验表明,表达的PebC1蛋白能够诱导黄瓜幼苗对灰霉病的抗性,具有激活蛋白的生物功能。
Activator protein is a type of protein elicitor which can induce many kinds of plants to gain systemic resistance,promotes plant growing and improve crop quality.To discover the new activator protein which has independent knowledge property right of China,an activator protein from Botrytis cinerea was isolated and purified.The protein elicitor could induce plant resistance to disease and drought.The protein coding gene was coloned and expressed in Escherichia coli and Pichia pastoris respectively.The function of expressed protein was evaluated.This research provided the genetic engineering way for obtaining an amount of activator protein quickly,which set foundation for molucular structure of protein-pesticide by analysis of relationship between protein structure and its function.The result is the following:
An activitor protein named as PebCl was isolated and purified from the hypha of Botrytis cinerea BC-4-2-2-1 with the method of boiling,centrifugation,ultra-filtration and anion exchange chromatography,etc.The activator protein appeared a single band on SDS-PAGE with silver staining.Molecular weight was about 36 kD.The pI of the purified activator protein was about 4.85 based with 2-DE.
Through enzyme digest in gel,the purified protein was analyzed by biological mass spectrometry(MS).Three certain peptide sequences were gotten.By blsat on NCBI,this protein has high homology to protein XP_001551609.1 from Botrytis cinerea and comparability was 95%.
Function study indicated that activator protein PebCl is able to promote wheat growth, enhance drought tolerance and increase tomato disease resisitance to gray mould.Soaking wheat seeds with different concentration of activator protein PebCl,seedling height was higher than that of the untreatment at 11 days after soaking and the best concentration of PebCl was 5μg/mL.Wheat root activity increased by 1.29 times comparing with CK when wheat seed was soaked with activator protein for 8h and coefficience of wheat drought tolerance increased from 36.53 to 57.08,which indicated that activator protein PebC1 improved wheat drought tolerance might correlate with the increasement of root activity.Tomato seeds were soaked with activator protein suspension at different concentration,then,45-day-old seedlings were sprayed with spores suspension of pathogenic fungus,results revealed that disease index of all the plants was lower than that of CK at 14~(th) day post inoculation with the spores of pathogenic fungus of 10~4/mL concentration and the induced efficiency reached 31.95%～69.19%.The best concentration of PebC1 was 10μg/mL Inducing efficiency was still as high as 66.10%at 21~(th) day. Tnis indicated that PebC1 not only increased tomato resistance to gray mould but also kept induced resisitance for long periods.
To reveal the mechanism of inducing plant disease resistance,the dynamic curve of Phenylalanine ammonia lyase(PAL),Peroxides(POD),polypheol oxidase(PPO) which related with plant resistance metabolism was measured.The activity of PAL,POD and PPO was increased after inducement.The activity of PAL increased by 46.84%at 24h.The activity of POD appeared the peak at 72h after inducement and increased 109.5%.The activity of PPO increased by 111.0%at 72h.The enhancement of the enzyme activity correlated with plant defense is one of the main physiological mechanism of protein activator inducing tomato disease resistance.
With the method of reverse genetics,the PebC1 gene cDNA sequence of Botrytis cinerea was gained and which is identical to gene XM_001551559 from Botrytis cinerea.Protein coding gene belongs to the nascent polypeptide-associated complex alpha polypeptide(alpha-NAC) protein family of Botrytis cinerea,the gene was named as PebC1.It has conserved NAC and UBA construction domains,which is same with activator protein PeaT1 from Alternaria tenuissim in structure domain.These two proteins are highly identical in amino acid composition, the similar amino acids are 79.4%and uniform amino acids were 71.0%.This cDNA is cost of 639bp and codes 212 amino acids.Predicted molecular weight is 23080.2 Dalton and its pI is 4.57.The hydrophilicity analysis revealed that it is a hydrophilic protein.
The prokaryotic expression of PebC1 was accomplished by connecting the gene to vector pET-28a(+),transforming recombined vector to competent cell Rossetta(DE3) and inducing expression by IPTG..The following purification was executed by hitrap chelating column on (?)KTA explorer10.The MS analysis proved that the expressional protein was actually activator protein of Alternaria tenuissima.The biological activity research revealed that this purified recombined protein could promote tomato seedling growth and increase drought resistance of wheat.
We constructed expressed vector of PebC1 Pichia pastoris and got expressed protein that has biological activity.It is the first time that PebC1 gene was amplified from cDNA of Botrytis cinerea and then was connected to the Pichia pastoris expressed vector pPIC9K to get recombination plasmid pPIC9K/PebC1.The plasmid was linearized by BglⅡand then transformed to Pichia pastoris GS115 by electric shock.Through screening by MD、G418-YPD plain and PCR method,we got secreted expressed strain.After methanol inducing and SDS-PAGE determination,PebC1 was successfully secreted expressed in Pichia pastoris.The biological activity experiments revealed that PebC1 protein could induce cucumber to gain resistance against gray mould disease and have the biological function of activator protein.
通过沸水浴、离心、阴离子交换层析等方法,从灰葡萄孢菌BC-4-2-2-1菌株的菌丝体中分离纯化出一种激活蛋白,命名为PebC1(protein elicitor botrytis cinerea1)。经SDS-PAGE银染显示该蛋白呈单一条带,相对分子量约为36kD,等电点为4.85。
用胶内酶解的方法,采用MALDI-TOF/TOF等质谱技术对纯化的激活蛋白PebC1进行了肽段序列测定,获得3个重复性较好的可靠肽段。在NCBI上进行比对发现,该蛋白与来源于灰葡萄孢菌的蛋白XP_001551609.1匹配率达95%。
功能研究表明,灰葡萄孢菌激活蛋白PebC1能显著促进小麦幼苗生长、提高小麦抗旱性、诱导番茄对灰霉病的系统抗性。不同浓度激活蛋白浸泡小麦种子,11天株高均高于对照,其中以5μg/mL激活蛋白对小麦生长的促进作用最明显;激活蛋白处理的小麦根系活力比对照增加1.29倍,小麦幼苗抗旱综合系数也从36.53提高到57.08,说明激活蛋白增强小麦的抗旱性可能与增加小麦根系活力有一定的关系;用不同浓度的激活蛋白浸泡番茄种子,幼苗生长45天接种灰葡萄孢菌孢子悬浮液(浓度为10~(4)个孢子/mL),接种后14天番茄的病情指数明显低于对照,对灰霉病的诱抗效果达到31.95%～69.19%,其中10μg/mL激活蛋白处理对番茄的诱抗效果最为明显,接种后21天的诱抗效果仍达66.10%,说明灰葡萄孢菌激活蛋白PebC1不仅能显著增强番茄对灰霉病的抗性,而且诱抗效果持续时间较长。
测定了PebC1处理后的番茄体内苯丙氨酸解氨酶(PAL)、过氧化物酶(POD)和多酚氧化酶(PPO)活性的动态变化。经激活蛋白PebC1处理后,番茄幼苗叶片的PAL、POD和PPO活性均有不同程度的提高,PAL活性在诱导后24小时达最高,较对照增加46.84%;POD活性在诱导72小时出现活性高峰,较对照增加109.5%;PPO活性在72小时出现峰值,比对照增加111.0%。PAL、POD和PPO均是与植物抗病相关的防御酶,番茄防御相关酶活性的提高可能是激活蛋白诱导番茄植株抗灰霉病的主要生理机制之一,该研究为揭示激活蛋白诱导植物抗病性的作用机理提供了理论依据。
通过反向遗传学的方法,获得了灰葡萄孢菌激活蛋白PebC1基因的cDNA序列,它与来源于灰葡萄孢菌的基因XM_001551559完全一致。该基因编码的蛋白属于灰葡萄孢菌的alpha-NAC蛋白(nascent polypeptide-associated complex alpha polypeptide),它含有保守的NAC结构域和UBA结构域,与来源于极细链格孢菌(Alternaria tenuissima)的激活蛋白PeaT1基因的结构域相同,另外,PebC1和PeaT1二者在编码的氨基酸组成上也有79.4%的相似性,其中完全一致的氨基酸占71.0%。PebC1基因全长639bp,编码212个氨基酸,预测分子量为23080.2,等电点为4.57。亲水性分析发现该蛋白属于亲水性蛋白。
进行了PebC1基因的原核表达。以原核表达系统中的pET-28a(+)为表达载体,将重组载体转入大肠杆菌Rossetta(DE3)菌株,通过IPTG诱导获得了PebC1基因的表达产物,利用(?)KTA explorer10蛋白纯化仪进行重组蛋白的纯化,选用Hitrap chelating Column柱进行亲和层析,得到了纯化的重组蛋白,并应用质谱分析鉴定了该基因表达的蛋白质为灰葡萄孢菌激活蛋白PebC1。功能测定结果表明,纯化的重组蛋白能促进番茄幼苗的生长,增强小麦的抗旱性。
构建了PebC1毕赤酵母表达载体,获得了具有生物活性的分泌表达蛋白。用PCR法从灰葡萄孢菌cDNA中扩增获得PebC1的编码序列并亚克隆到毕赤酵母表达系统的表达载体pPIC9K上,得到重组质粒pPIC9K/PebC1。重组质粒经BglⅡ线性化后电击导入毕赤酵母GS115中,采用MD、G418-YPD平板和PCR法进行筛选,获得了分泌表达的重组毕赤酵母菌株。用甲醇诱导PebC1表达,经SDS-PAGE检测证明,PebC1基因在毕赤酵母中成功分泌表达。生物活性试验表明,表达的PebC1蛋白能够诱导黄瓜幼苗对灰霉病的抗性,具有激活蛋白的生物功能。
Activator protein is a type of protein elicitor which can induce many kinds of plants to gain systemic resistance,promotes plant growing and improve crop quality.To discover the new activator protein which has independent knowledge property right of China,an activator protein from Botrytis cinerea was isolated and purified.The protein elicitor could induce plant resistance to disease and drought.The protein coding gene was coloned and expressed in Escherichia coli and Pichia pastoris respectively.The function of expressed protein was evaluated.This research provided the genetic engineering way for obtaining an amount of activator protein quickly,which set foundation for molucular structure of protein-pesticide by analysis of relationship between protein structure and its function.The result is the following:
An activitor protein named as PebCl was isolated and purified from the hypha of Botrytis cinerea BC-4-2-2-1 with the method of boiling,centrifugation,ultra-filtration and anion exchange chromatography,etc.The activator protein appeared a single band on SDS-PAGE with silver staining.Molecular weight was about 36 kD.The pI of the purified activator protein was about 4.85 based with 2-DE.
Through enzyme digest in gel,the purified protein was analyzed by biological mass spectrometry(MS).Three certain peptide sequences were gotten.By blsat on NCBI,this protein has high homology to protein XP_001551609.1 from Botrytis cinerea and comparability was 95%.
Function study indicated that activator protein PebCl is able to promote wheat growth, enhance drought tolerance and increase tomato disease resisitance to gray mould.Soaking wheat seeds with different concentration of activator protein PebCl,seedling height was higher than that of the untreatment at 11 days after soaking and the best concentration of PebCl was 5μg/mL.Wheat root activity increased by 1.29 times comparing with CK when wheat seed was soaked with activator protein for 8h and coefficience of wheat drought tolerance increased from 36.53 to 57.08,which indicated that activator protein PebC1 improved wheat drought tolerance might correlate with the increasement of root activity.Tomato seeds were soaked with activator protein suspension at different concentration,then,45-day-old seedlings were sprayed with spores suspension of pathogenic fungus,results revealed that disease index of all the plants was lower than that of CK at 14~(th) day post inoculation with the spores of pathogenic fungus of 10~4/mL concentration and the induced efficiency reached 31.95%～69.19%.The best concentration of PebC1 was 10μg/mL Inducing efficiency was still as high as 66.10%at 21~(th) day. Tnis indicated that PebC1 not only increased tomato resistance to gray mould but also kept induced resisitance for long periods.
To reveal the mechanism of inducing plant disease resistance,the dynamic curve of Phenylalanine ammonia lyase(PAL),Peroxides(POD),polypheol oxidase(PPO) which related with plant resistance metabolism was measured.The activity of PAL,POD and PPO was increased after inducement.The activity of PAL increased by 46.84%at 24h.The activity of POD appeared the peak at 72h after inducement and increased 109.5%.The activity of PPO increased by 111.0%at 72h.The enhancement of the enzyme activity correlated with plant defense is one of the main physiological mechanism of protein activator inducing tomato disease resistance.
With the method of reverse genetics,the PebC1 gene cDNA sequence of Botrytis cinerea was gained and which is identical to gene XM_001551559 from Botrytis cinerea.Protein coding gene belongs to the nascent polypeptide-associated complex alpha polypeptide(alpha-NAC) protein family of Botrytis cinerea,the gene was named as PebC1.It has conserved NAC and UBA construction domains,which is same with activator protein PeaT1 from Alternaria tenuissim in structure domain.These two proteins are highly identical in amino acid composition, the similar amino acids are 79.4%and uniform amino acids were 71.0%.This cDNA is cost of 639bp and codes 212 amino acids.Predicted molecular weight is 23080.2 Dalton and its pI is 4.57.The hydrophilicity analysis revealed that it is a hydrophilic protein.
The prokaryotic expression of PebC1 was accomplished by connecting the gene to vector pET-28a(+),transforming recombined vector to competent cell Rossetta(DE3) and inducing expression by IPTG..The following purification was executed by hitrap chelating column on (?)KTA explorer10.The MS analysis proved that the expressional protein was actually activator protein of Alternaria tenuissima.The biological activity research revealed that this purified recombined protein could promote tomato seedling growth and increase drought resistance of wheat.
We constructed expressed vector of PebC1 Pichia pastoris and got expressed protein that has biological activity.It is the first time that PebC1 gene was amplified from cDNA of Botrytis cinerea and then was connected to the Pichia pastoris expressed vector pPIC9K to get recombination plasmid pPIC9K/PebC1.The plasmid was linearized by BglⅡand then transformed to Pichia pastoris GS115 by electric shock.Through screening by MD、G418-YPD plain and PCR method,we got secreted expressed strain.After methanol inducing and SDS-PAGE determination,PebC1 was successfully secreted expressed in Pichia pastoris.The biological activity experiments revealed that PebC1 protein could induce cucumber to gain resistance against gray mould disease and have the biological function of activator protein.
引文
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