番茄灰霉病菌对丙烷脒的抗药性风险研究
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摘要
近年来,由于杀菌剂抗药性问题日趋严重,许多杀菌剂对植物病害防治效果逐渐下降,某些原已被很好控制的植物病害也有加重发生的趋势,随着传统内吸性杀菌剂的持续应用,抗药性问题势必更加突出。研究新型杀菌剂抗药性风险对延长药剂使用寿命,指导田间科学用药,避免和延缓抗药性的发生具有十分重要的意义。番茄灰霉病菌(Botrytis cinerea)是一种寄主范围广、繁殖速度快、遗传变异大、危害极其严重的病原菌,目前抗药性问题已成为该病菌防治中所面临的突出问题。丙烷脒是西北农林科技大学无公害农药研究服务中心研制的具有我国自主知识产权的新型芳香二脒类内吸性杀菌剂,主要用于防治由番茄灰霉病菌引起的灰霉病。目前其抗药性方面的研究还未见报道。基于此,本研究以番茄灰霉病菌为靶标菌,主要从敏感基线的建立、抗药性风险评价、抗药突变体产生抗药性的机制等方面进行了较为系统的测试和探讨,取得如下结果:
1.采用菌丝生长速率法测定了采自西安、咸阳、杨凌、渭南、宝鸡等5个地区的41株番茄灰霉病菌对丙烷脒的敏感性。结果表明,丙烷脒对所有供试番茄灰霉病菌菌株均具有良好的抑制作用,不同菌株间EC50值差异较小,其中最大为1.460μg/mL,最小为0.182μg/mL,相差8.0倍,平均EC50值为0.791±0.269μg/mL。不同番茄灰霉病菌菌株对丙烷脒的敏感性频率分布呈连续单峰曲线,接近正态分布。因此,可以采用这些菌株的平均EC50值作为田间抗药性检测的敏感基线。
2.通过紫外诱变和药剂选择共获得10株抗药突变体,其中紫外诱导获得1株,突变频率为1.98×10-9,药剂选择获得9株,突变频率为0.025。根据FAO对抗性的划分标准,结合敏感基线及抗药突变体的抗性水平,本研究将抗性表型划分为4类:EC50值小于5.0μg/mL时,为敏感菌株(S);EC50值在5.0~15.0μg/mL时,为低抗菌株(LR);EC50值在15.1~75.0μg/mL时,为中抗菌株(MR);EC50值大于75.0μg/mL时,为高抗菌株(HR)。据此,本试验共获得了8株低抗菌株和2株中抗菌株,未获得高抗菌株。获得抗性菌株的突变频率和抗性菌株的抗性水平均很低,表明番茄灰霉病菌对丙烷脒的抗药性发展非常缓慢。适合度研究结果显示,抗性菌株的生长量、产孢量、孢子萌发率、致病力等适合度指标均明显低于敏感菌株。交互抗药性测定结果表明,丙烷脒与嘧霉胺、多菌灵、速克灵、扑海因等常用杀灰霉剂无交互抗药性。综合分析以上研究结果可见,番茄灰霉病菌对丙烷脒具有低等抗药性风险。
3.线粒体复合酶Ⅲ活性测定结果表明,离体条件下丙烷脒对敏感菌株和抗性菌株线粒体复合酶Ⅲ的活性均没有影响,活体条件下丙烷脒对敏感菌株线粒体复合酶Ⅲ的活性具有明显抑制作用,而抗性菌株的活性基本没有变化。结合前人研究结果可见丙烷脒对病菌线粒体复合酶Ⅲ相关基因表达的抑制是其抑菌的主因;番茄灰霉病菌对丙烷脒的抗药性与线粒体复合酶Ⅲ活性的变化密切相关。
4.渗透压敏感性测定结果表明,敏感菌株对低渗透压和高渗透压均表现敏感,低抗菌株仅对低渗透压敏感,中抗菌株对低渗透压和高渗透压均不敏感,说明抗性菌株的渗透调节能力要大于敏感菌株。细胞膜透性测定结果表明,用各供试浓度丙烷脒处理番茄灰霉病菌敏感菌株和低抗菌株,以及用高浓度丙烷脒(25μg/mL和50μg/mL)处理中抗菌株后,所有菌株细胞膜的透性均有所增大;敏感菌株和低抗菌株最终(240min)的相对渗率虽然比较接近,但丙烷脒对敏感菌株细胞膜的损伤速度明显快于低抗菌株,说明两者的细胞膜透性及对丙烷脒的适应能力存在一定差异;此外,中抗菌株最终(240min)的相对渗率明显低于敏感菌株和低抗菌株,说明中抗菌株适应丙烷脒的能力明显强于敏感菌株和低抗菌株,其细胞膜受损伤的程度相对较小,使其细胞膜的透性相对低于敏感菌株和低抗菌株,且这种适应能力随着丙烷脒处理浓度的增大而降低。对渗透压的研究结果也证实了这一点。可见,渗透压敏感性下降和细胞膜透性的改变可能是番茄灰霉病菌对丙烷脒产生抗药性的生理机制之一。
5.通过聚丙烯酰胺凝胶电泳比较了抗性菌株和敏感菌株酯酶同工酶和可溶性蛋白质的差异,发现抗性菌株和敏感菌株之间无论在谱带数上还是含量上都存在明显的差异,说明番茄灰霉病菌对丙烷脒的抗药性与酯酶同工酶和可溶性蛋白的组成及含量变化有关。
6.利用cDNA-AFLP技术,对敏感菌株(091)和中抗菌株(F091-16)的基因表达进行了分析,共分离得到38个差异表达的TDFs片段,并对其中6个典型的TDFs片段进行克隆、测序和序列分析。结果表明,在6个差异片段中,有4个找到了同源序列,其中TDF2、TDF4、TDF5是同一基因片段,其与细胞色素P450烷烃羟化酶基因序列有很高的同源性,另一片段TDF1与C-1-四氢叶酸复合酶有较高的同源性,其余两个基因片段TDF3、TDF6未找到与之同源的序列。C-1-四氢叶酸复合酶基因的上调表达,必将促进核酸生物合成速率的加快,进而补偿因药物分子作用导致的靶基因表达水平的下降,使线粒体复合酶Ⅲ的活性恢复正常。结合丙烷脒抑菌机理研究结果,推测C-1-四氢叶酸复合酶基因的上调表达可能是番茄灰霉病菌对丙烷脒产生抗药性的主要因素。另外,细胞色素P450烷烃羟化酶基因的上调表达也与抗药性的产生有关。
综上所述,番茄灰霉病菌对丙烷脒具有低等抗药性风险。番茄灰霉病菌对丙烷脒的抗药性是多种因素综合作用的结果,其中C-1-四氢叶酸复合酶基因的上调表达可能是主要因素,具体的抗性机制可能为:渗透压敏感性和细胞膜透性下降,使药物分子难以穿过细胞膜进入菌丝体内,进入菌丝体内的部分药物又在酯酶同工酶、细胞色素P450烷烃羟化酶等解毒代谢酶的作用下被代谢分解;C-1-四氢叶酸复合酶基因的上调表达,使核酸生物合成速率加快,补偿了因药物分子作用导致的靶基因表达水平的下降,使线粒体复合酶Ⅲ的活性恢复正常,从而使抗性菌株表现出一定的抗药性。此外,结合前人研究结果可以进一步确认丙烷脒对病菌线粒体复合酶Ⅲ相关基因表达的抑制是其抑菌的主因。
In recent years, the problem of fungicide resistance causes a decline effect in kinds of diseases. Besides, some kinds of diseases, which can be easily managed before, show a tendency of more and more severe. Along with the advance of agricultural modernization and chemical control of plant diseases, the resistant problem becomes more serious than before. B. cinerea, which has the characteristic of wide hosts, fastly genetic mutation and severe damage, is of a pathogen with serious resistance problem. Propamidine, a kind of aromatic diamidine compound with Chinese intellectual property rights, is developed by Research & Development Center of Biorational Pesticide in Northwest A&F University independently and is used for controlling grey mould caused by B. cinerea. Until now, few articles about resistance of propamidine had been reported. So, this research is based on the target pathogen B. cinerea and in terms of baseline-sensitivity assay, assess of resistance risk and the mechanism of development of resistance on resistant mutants. The main results and conclusions were outlined as follows:
1. The sensitivity of 41 strains of B. cinerea which was isolated from 5 districts of Xi’an, Xianyang, Yangling, Weinan and Baoji was tested by the method of mycelial growth rate. According to the result, all the tested strains were inhibited seriously by propamidine. Besides, low differential of EC50 values was tested among the different strains. The EC50 values were 0.182 to 1.460μg/mL. The mean EC50 was 0.791±0.269μg/mL, representing a range-of-variation factor of 8.0. The frequency distribution of sensitivities of different strains to propamidine was representing a curve of continuous single peak which was approaching a unimodal curve. No low sensitivity of resistant population was detected. So the mean EC50 of these strains could be used as a baseline for observing the sensitivity in B. cinerea populations to propamidine.
2. 10 resistance mutants with high genetic stability were obtained by UV irradiation and selection for resistance to fungicide. One resistant mutant was induced by UV irradiation and nine resistant mutants were obtained by selection for resistance to fungicide with the mutation frequency 1.98×10-9 and 0.025 respectively. Based on the standard of resistance created by FAO and the baseline sensitivity and the resistance level of resistant mutants, the resistant mutants were divided into low (L), moderate (M) and high (H) resistant level, determined by the EC50 values of 5.0-15.0μg/mL, 15.1-75.0μg/mL and more than 75.0μg/mL, respectively. So, 8 LR mutants, 2 MR mutants and no HR mutant were obtained. Low mutation frequency and resistant level showed the fact that it is hard for B. cinerea developing resistance to propamidine. According to the result of fitness parameters of the mutants, the hyphal growth, sporulation, spore germination and pathogenicity, showed significantly lower than the sensitive strains. Besides, with the result of cross resistance test, propamidine has no cross resistance with the botryticides in common use like pyrimethanil, carbendazim, iprodione and procymidone. Above all, B. cinerea has low resistance risk to propamidine.
3. According to the result of activity of mitochondrial complexⅢ, propamidine had no effect on both sensitive strain and resistance-strains during the test of post-treatment on mitochondrial. However, during the pre-treatment activity test, propamidine possessed obvious inhibition on sensitive strains and had no significant effect on resistance-strains. The inhibition of propamidine on mitochondrial complexⅢrelative genes expression is a key factor, according to previous research.
4. During the osmolarity sensitivity test, sensitive strain was sensitive against low and high osmolarity. And, LR (low resistance) strain was sensitive against low osmolarity only. Besides, MR (moderate resistance) strain possessed no sensitivity against both low and high osmolarity. Basing on the result of cell membrane permeability, both sensitive strains and LR strains showed high cell membrane permeability after treated by different concentration of propamidine, especially treated by high concentration of propamidine (25μg/mL and 50μg/mL). Although the final relative leakage rate (240min) of sensitive strains and LR strains were close, the membrane injury speed of sensitive strains showed more fast than LR strains during the treatment of propamidine, which indicates that the cell membrane permeability and the adaptation to propamidine between sensitive strains and LR strains is different. Further more, the final relative leakage rate (240min) of MR strains showed more lower than sensitive strains and LR strains, which indicates that MR strains with much higher ability of adapting to propamidine than sensitive strains and LR strains. The lower cell membrane injury degree makes the lower cell membrane permeability of MR strains comparing with sensitive strains and LR strains, and the ability of adapting to propamidine become lower with the concentration increasing, which also can be indicated by the result of osmolarity test. So, the decrease of osmotic sensitivity and the change of cell membrane permeability may be one of the physiological and biochemical mechanisms on resistance of B. cinerea to propamidine.
5. The difference of esterase isoenzyme and soluble proteins were compared by the PAGE between resistance-strains and sensitive strains . The result showed obvious difference of the band amount and content between resistance-strains and sensitive strains, which indicates that the resistance of B. cinerea to propamidine is relative to the formation or content change of esterase isoenzyme and soluble proteins.
6. Gene expression of resistance strain (091) and MR strain (F091-16) was analyzed by the cDNA-AFLP. About 38 differentially expressed TDFs were isolated and 6 of them were cloned and sequenced. The result showed that 4 of 6 differentially expressed TDFs were found with homological sequences. Besides, TDF2, TDF4 and TDF4, which belongs to the same gene fragment, possess high homology with the gene sequence of cytochrome P450 alkane hydroxylase. And, another differential TDF named TDF1 possesses high homology with gene sequence of C-1-tetrahydrofolate synthase and the others two TDFs, TDF3 and TDF6, possesses no homology with existing sequences. Up-regulated expression of C-1-tetrahydrofolate synthase gene can promote the synthesis speed of nucleic acid which lead to the decrease of target gene expression which caused by retrieving of medical molecular effect, and the activity of mitochondrial complexⅢcan be recovered. Along with the result of mechanism on agriculture fungal inhibition of propamidine, the conclusion is speculated that the up-regulated expression of C-1-tetrahydrofolate synthase gene may be the main fact of resistance development on B. cinerea to propamidine. Further more, the up-regulated expression of cytochrome P450 alkane hydroxylase gene is also relation to the resistance development.
Above all, the resistance risk is low on B. cinerea to propamidine. The resistance of B. cinerea to propamidine is leaded by lots of factors. Among the factors, the up-regulated expression of C-1-tetrahydrofolate synthase gene may be the main factor: The decrease of osmolarity and cell membrane permeability makes propamidine hard to pass through the cell membrane into plasma. Part of some medical molecular in plasma can be degraded by esterase isoenzyme or cytochrome P450 alkane hydroxylase. C-1-tetrahydrofolate synthase gene can promote the synthesis speed of nucleic acid which leads to the decrease of target gene expression caused by retrieving of medical molecular effect and resistance is possessed finally on resistance-strains. Besides, the gene expression which inhibition of propamidine on mitochondrial complexⅢis a key factor, according to previous research.
1.采用菌丝生长速率法测定了采自西安、咸阳、杨凌、渭南、宝鸡等5个地区的41株番茄灰霉病菌对丙烷脒的敏感性。结果表明,丙烷脒对所有供试番茄灰霉病菌菌株均具有良好的抑制作用,不同菌株间EC50值差异较小,其中最大为1.460μg/mL,最小为0.182μg/mL,相差8.0倍,平均EC50值为0.791±0.269μg/mL。不同番茄灰霉病菌菌株对丙烷脒的敏感性频率分布呈连续单峰曲线,接近正态分布。因此,可以采用这些菌株的平均EC50值作为田间抗药性检测的敏感基线。
2.通过紫外诱变和药剂选择共获得10株抗药突变体,其中紫外诱导获得1株,突变频率为1.98×10-9,药剂选择获得9株,突变频率为0.025。根据FAO对抗性的划分标准,结合敏感基线及抗药突变体的抗性水平,本研究将抗性表型划分为4类:EC50值小于5.0μg/mL时,为敏感菌株(S);EC50值在5.0~15.0μg/mL时,为低抗菌株(LR);EC50值在15.1~75.0μg/mL时,为中抗菌株(MR);EC50值大于75.0μg/mL时,为高抗菌株(HR)。据此,本试验共获得了8株低抗菌株和2株中抗菌株,未获得高抗菌株。获得抗性菌株的突变频率和抗性菌株的抗性水平均很低,表明番茄灰霉病菌对丙烷脒的抗药性发展非常缓慢。适合度研究结果显示,抗性菌株的生长量、产孢量、孢子萌发率、致病力等适合度指标均明显低于敏感菌株。交互抗药性测定结果表明,丙烷脒与嘧霉胺、多菌灵、速克灵、扑海因等常用杀灰霉剂无交互抗药性。综合分析以上研究结果可见,番茄灰霉病菌对丙烷脒具有低等抗药性风险。
3.线粒体复合酶Ⅲ活性测定结果表明,离体条件下丙烷脒对敏感菌株和抗性菌株线粒体复合酶Ⅲ的活性均没有影响,活体条件下丙烷脒对敏感菌株线粒体复合酶Ⅲ的活性具有明显抑制作用,而抗性菌株的活性基本没有变化。结合前人研究结果可见丙烷脒对病菌线粒体复合酶Ⅲ相关基因表达的抑制是其抑菌的主因;番茄灰霉病菌对丙烷脒的抗药性与线粒体复合酶Ⅲ活性的变化密切相关。
4.渗透压敏感性测定结果表明,敏感菌株对低渗透压和高渗透压均表现敏感,低抗菌株仅对低渗透压敏感,中抗菌株对低渗透压和高渗透压均不敏感,说明抗性菌株的渗透调节能力要大于敏感菌株。细胞膜透性测定结果表明,用各供试浓度丙烷脒处理番茄灰霉病菌敏感菌株和低抗菌株,以及用高浓度丙烷脒(25μg/mL和50μg/mL)处理中抗菌株后,所有菌株细胞膜的透性均有所增大;敏感菌株和低抗菌株最终(240min)的相对渗率虽然比较接近,但丙烷脒对敏感菌株细胞膜的损伤速度明显快于低抗菌株,说明两者的细胞膜透性及对丙烷脒的适应能力存在一定差异;此外,中抗菌株最终(240min)的相对渗率明显低于敏感菌株和低抗菌株,说明中抗菌株适应丙烷脒的能力明显强于敏感菌株和低抗菌株,其细胞膜受损伤的程度相对较小,使其细胞膜的透性相对低于敏感菌株和低抗菌株,且这种适应能力随着丙烷脒处理浓度的增大而降低。对渗透压的研究结果也证实了这一点。可见,渗透压敏感性下降和细胞膜透性的改变可能是番茄灰霉病菌对丙烷脒产生抗药性的生理机制之一。
5.通过聚丙烯酰胺凝胶电泳比较了抗性菌株和敏感菌株酯酶同工酶和可溶性蛋白质的差异,发现抗性菌株和敏感菌株之间无论在谱带数上还是含量上都存在明显的差异,说明番茄灰霉病菌对丙烷脒的抗药性与酯酶同工酶和可溶性蛋白的组成及含量变化有关。
6.利用cDNA-AFLP技术,对敏感菌株(091)和中抗菌株(F091-16)的基因表达进行了分析,共分离得到38个差异表达的TDFs片段,并对其中6个典型的TDFs片段进行克隆、测序和序列分析。结果表明,在6个差异片段中,有4个找到了同源序列,其中TDF2、TDF4、TDF5是同一基因片段,其与细胞色素P450烷烃羟化酶基因序列有很高的同源性,另一片段TDF1与C-1-四氢叶酸复合酶有较高的同源性,其余两个基因片段TDF3、TDF6未找到与之同源的序列。C-1-四氢叶酸复合酶基因的上调表达,必将促进核酸生物合成速率的加快,进而补偿因药物分子作用导致的靶基因表达水平的下降,使线粒体复合酶Ⅲ的活性恢复正常。结合丙烷脒抑菌机理研究结果,推测C-1-四氢叶酸复合酶基因的上调表达可能是番茄灰霉病菌对丙烷脒产生抗药性的主要因素。另外,细胞色素P450烷烃羟化酶基因的上调表达也与抗药性的产生有关。
综上所述,番茄灰霉病菌对丙烷脒具有低等抗药性风险。番茄灰霉病菌对丙烷脒的抗药性是多种因素综合作用的结果,其中C-1-四氢叶酸复合酶基因的上调表达可能是主要因素,具体的抗性机制可能为:渗透压敏感性和细胞膜透性下降,使药物分子难以穿过细胞膜进入菌丝体内,进入菌丝体内的部分药物又在酯酶同工酶、细胞色素P450烷烃羟化酶等解毒代谢酶的作用下被代谢分解;C-1-四氢叶酸复合酶基因的上调表达,使核酸生物合成速率加快,补偿了因药物分子作用导致的靶基因表达水平的下降,使线粒体复合酶Ⅲ的活性恢复正常,从而使抗性菌株表现出一定的抗药性。此外,结合前人研究结果可以进一步确认丙烷脒对病菌线粒体复合酶Ⅲ相关基因表达的抑制是其抑菌的主因。
In recent years, the problem of fungicide resistance causes a decline effect in kinds of diseases. Besides, some kinds of diseases, which can be easily managed before, show a tendency of more and more severe. Along with the advance of agricultural modernization and chemical control of plant diseases, the resistant problem becomes more serious than before. B. cinerea, which has the characteristic of wide hosts, fastly genetic mutation and severe damage, is of a pathogen with serious resistance problem. Propamidine, a kind of aromatic diamidine compound with Chinese intellectual property rights, is developed by Research & Development Center of Biorational Pesticide in Northwest A&F University independently and is used for controlling grey mould caused by B. cinerea. Until now, few articles about resistance of propamidine had been reported. So, this research is based on the target pathogen B. cinerea and in terms of baseline-sensitivity assay, assess of resistance risk and the mechanism of development of resistance on resistant mutants. The main results and conclusions were outlined as follows:
1. The sensitivity of 41 strains of B. cinerea which was isolated from 5 districts of Xi’an, Xianyang, Yangling, Weinan and Baoji was tested by the method of mycelial growth rate. According to the result, all the tested strains were inhibited seriously by propamidine. Besides, low differential of EC50 values was tested among the different strains. The EC50 values were 0.182 to 1.460μg/mL. The mean EC50 was 0.791±0.269μg/mL, representing a range-of-variation factor of 8.0. The frequency distribution of sensitivities of different strains to propamidine was representing a curve of continuous single peak which was approaching a unimodal curve. No low sensitivity of resistant population was detected. So the mean EC50 of these strains could be used as a baseline for observing the sensitivity in B. cinerea populations to propamidine.
2. 10 resistance mutants with high genetic stability were obtained by UV irradiation and selection for resistance to fungicide. One resistant mutant was induced by UV irradiation and nine resistant mutants were obtained by selection for resistance to fungicide with the mutation frequency 1.98×10-9 and 0.025 respectively. Based on the standard of resistance created by FAO and the baseline sensitivity and the resistance level of resistant mutants, the resistant mutants were divided into low (L), moderate (M) and high (H) resistant level, determined by the EC50 values of 5.0-15.0μg/mL, 15.1-75.0μg/mL and more than 75.0μg/mL, respectively. So, 8 LR mutants, 2 MR mutants and no HR mutant were obtained. Low mutation frequency and resistant level showed the fact that it is hard for B. cinerea developing resistance to propamidine. According to the result of fitness parameters of the mutants, the hyphal growth, sporulation, spore germination and pathogenicity, showed significantly lower than the sensitive strains. Besides, with the result of cross resistance test, propamidine has no cross resistance with the botryticides in common use like pyrimethanil, carbendazim, iprodione and procymidone. Above all, B. cinerea has low resistance risk to propamidine.
3. According to the result of activity of mitochondrial complexⅢ, propamidine had no effect on both sensitive strain and resistance-strains during the test of post-treatment on mitochondrial. However, during the pre-treatment activity test, propamidine possessed obvious inhibition on sensitive strains and had no significant effect on resistance-strains. The inhibition of propamidine on mitochondrial complexⅢrelative genes expression is a key factor, according to previous research.
4. During the osmolarity sensitivity test, sensitive strain was sensitive against low and high osmolarity. And, LR (low resistance) strain was sensitive against low osmolarity only. Besides, MR (moderate resistance) strain possessed no sensitivity against both low and high osmolarity. Basing on the result of cell membrane permeability, both sensitive strains and LR strains showed high cell membrane permeability after treated by different concentration of propamidine, especially treated by high concentration of propamidine (25μg/mL and 50μg/mL). Although the final relative leakage rate (240min) of sensitive strains and LR strains were close, the membrane injury speed of sensitive strains showed more fast than LR strains during the treatment of propamidine, which indicates that the cell membrane permeability and the adaptation to propamidine between sensitive strains and LR strains is different. Further more, the final relative leakage rate (240min) of MR strains showed more lower than sensitive strains and LR strains, which indicates that MR strains with much higher ability of adapting to propamidine than sensitive strains and LR strains. The lower cell membrane injury degree makes the lower cell membrane permeability of MR strains comparing with sensitive strains and LR strains, and the ability of adapting to propamidine become lower with the concentration increasing, which also can be indicated by the result of osmolarity test. So, the decrease of osmotic sensitivity and the change of cell membrane permeability may be one of the physiological and biochemical mechanisms on resistance of B. cinerea to propamidine.
5. The difference of esterase isoenzyme and soluble proteins were compared by the PAGE between resistance-strains and sensitive strains . The result showed obvious difference of the band amount and content between resistance-strains and sensitive strains, which indicates that the resistance of B. cinerea to propamidine is relative to the formation or content change of esterase isoenzyme and soluble proteins.
6. Gene expression of resistance strain (091) and MR strain (F091-16) was analyzed by the cDNA-AFLP. About 38 differentially expressed TDFs were isolated and 6 of them were cloned and sequenced. The result showed that 4 of 6 differentially expressed TDFs were found with homological sequences. Besides, TDF2, TDF4 and TDF4, which belongs to the same gene fragment, possess high homology with the gene sequence of cytochrome P450 alkane hydroxylase. And, another differential TDF named TDF1 possesses high homology with gene sequence of C-1-tetrahydrofolate synthase and the others two TDFs, TDF3 and TDF6, possesses no homology with existing sequences. Up-regulated expression of C-1-tetrahydrofolate synthase gene can promote the synthesis speed of nucleic acid which lead to the decrease of target gene expression which caused by retrieving of medical molecular effect, and the activity of mitochondrial complexⅢcan be recovered. Along with the result of mechanism on agriculture fungal inhibition of propamidine, the conclusion is speculated that the up-regulated expression of C-1-tetrahydrofolate synthase gene may be the main fact of resistance development on B. cinerea to propamidine. Further more, the up-regulated expression of cytochrome P450 alkane hydroxylase gene is also relation to the resistance development.
Above all, the resistance risk is low on B. cinerea to propamidine. The resistance of B. cinerea to propamidine is leaded by lots of factors. Among the factors, the up-regulated expression of C-1-tetrahydrofolate synthase gene may be the main factor: The decrease of osmolarity and cell membrane permeability makes propamidine hard to pass through the cell membrane into plasma. Part of some medical molecular in plasma can be degraded by esterase isoenzyme or cytochrome P450 alkane hydroxylase. C-1-tetrahydrofolate synthase gene can promote the synthesis speed of nucleic acid which leads to the decrease of target gene expression caused by retrieving of medical molecular effect and resistance is possessed finally on resistance-strains. Besides, the gene expression which inhibition of propamidine on mitochondrial complexⅢis a key factor, according to previous research.
引文
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