浙江衢州地区柑橘绿霉菌对抑霉唑和多菌灵的抗性及其抗性分子机制的研究
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摘要
柑橘绿霉病(Penicillium digitatum)是贮藏柑橘的最主要病害,引起的损失占采后损失的90%以上。化学防治是控制柑橘绿霉病的最有效的方法,但随着药剂的长期使用,抗药性菌株不断产生,从而带来防治效果下降甚至失效的问题。为探讨浙江衢州产区柑橘绿霉菌的对常用药剂抑霉唑和多菌灵的抗性现状和分子机制,本研究评价了来自衢州市,包括柯城区、衢江区和开化县12个贮藏库的70个柑橘绿霉病菌菌株对抑霉唑和多菌灵的抗性水平及其分子机制。结果表明,柯城区和衢江区的抑霉唑抗性菌株(最低抑制浓度(MIC)≥0.5μg/ml)的比例分别为77.1%和62.5%。两地抗性菌株的平均ECso值分别为2.07±1.04Hg/mL和2.35±0.73μg/mL,分别是当地敏感菌株EC50的41.4和47倍。而来自开化县的菌株均对抑霉唑敏感(MIC≤0.1μg/mL),平均EC50值为0.04±0.02μg/mL。柯城区和衢江区的多菌灵抗性菌株(MIC≥10μg/mL)的比例分别为54.3%和54.2%,而开化县的抗性菌株比例仅为9.1%。
本研究克隆测定8个多菌灵敏感菌株和10个多菌灵抗性菌株的β-微管蛋白基因全长序列,比对分析后发现敏感菌株的1255位核苷酸序列为TTC,其编码的氨基酸(200位)为苯丙氨酸,而在抗性菌株中相应位点的核苷酸为序列为TAC,编码的氨基酸为络氨酸。除此外,没有发现其他与抗性相关的突变。因此推断来自浙江衢州的柑橘绿霉病菌对多菌灵的抗性与其β-微管蛋白200位氨基酸由苯丙氨酸突变成络氨酸相关。
已知抑霉唑靶标基因(CYP51)启动区的4个额外的126bp片段简单串联重复插入(IMZ-R1)和其中一个199bp片段的插入(IMZ-R2)与柑橘绿霉菌抑霉唑抗性相关。使用检测区分这两种抗性分子机制的引物对CYP51A1/CYP51R2, PCR扩增发现,来自衢州的所有抗抑霉唑绿霉菌的抗性分子机制既不属于IMZ-R1也不属于IMZ-R2;测序比较敏感菌株和抗性菌株CYP51基因的全长序列,也未发现与抗性相关的点突变。
通过查找柑橘绿霉菌转录组数据库,发现两个CYP51同源基因的EST序列,命名为PdCYP51B和PdCYP51C基因。结合NCBI上产黄青霉(NS_000201)的同源物种信息,设计了CYP51B-R/CYP51B-F和CYP51C-R/CYP51C-F两对引物,成功从抑霉唑敏感菌株PdKH8菌株中克隆到包括启动子和编码区在内PdCYP51B和PdCYP51C两个全长基因。PdCYP51B包含2919bp,包括1751bp完整的编码区,以及它上游742bp的序列和下游425bp的序列,含有3个分别为73bp、51bp、52bp的外显子,分别位于989bp与1061bp之间,1260bp与1310bp之间,和2376bp与2427bp之间,编码485个氨基酸序列。PdCYP51B在GenBank中的登录号为HQ724322.PdCYP51C,包括一个1497bp的开放阅读框,含有分别位于541bp与596bp,682bp与733bp,823bp与885bp,1177bp与1242bp,和1292bp与1348bp之间的5个外显子,编码496个氨基酸序列。PdCYP51C在GenBank中的登录号为HQ724324。序列分析表明PdCYP51B基因与烟曲霉(AfCYP51B, XP_749134.1)和小麦赤霉病菌(FgCYP51B, FGSG_01000)有78%和61%一致性。PdCYP51C与禾谷镰刀菌(FgCYP51C, FGSG_11024)和尖孢镰刀菌(FoCYP51C, FOXG_13138)有19%的一致性。PdCYP51B与PdCYP51A和PdCYP51C有59%和22%的一致性,而PdCYP51C基因与PdCYP51A有23%的一致性。
克隆、测序并比较8个抑霉唑敏感(IMZ-S)和1个IMZ-R1、1个IMZ-R2,以及6个IMZ-R3菌株的CYP51B和CYP51C基因序列,发现这些菌株的CYP51C基本一致,未发现与抗性相关的点突变,但是,在所有IMZ-R3菌株的CYP51B基因启动子区均发现有一个199bp序列的插入,而该片段在所分析的IMZ-S、IMZ-R1和IMZ-R2菌株中均不存在。因此推测,IMZ-R3菌株的抗性机制与该菌株的CYP51基因启动子去的199bp插入相关。基于该分子机制,我们设计了一对检测IMZ-R3的引物,对来自衢州的IMZ-R3菌株进行PCR扩增,结果发现,能从所有这些抗性菌株中扩增到预期的片段。因此,认为衢州柑橘绿霉菌的抑霉唑抗性菌株的抗性分子机制均属于CYP51B基因启动子区有199bp序列的插入。
Penicillium digitatum is the most devastating pathogen of citrus fruit, being responsible for about 90% of production losses during post-harvest handling. Fungieide treatment is the main measure for postharvest disease control of citrus. However. the emergence and prevalence of fungicide-resistant biotypes followed by the successively and intensively using of fungieides often results in the decrease of control effect. In this study,70 isolates of P digitatum from 12 citrus storage houses in Quzhou city (included Kecheng and Qujiang districts, as well as Kaihua county), Zhejiang province, were assayed for their resistance to imazalil and carbendazim, and thier resistance machnisms were studied further. The results demonstrated that the frequency of imazalil-resistant isolates (minimum inhibitory concentration (MIC)≥0.5μg/mL) in Kecheng and Qujiang districts were 77.1% and 62.5%, respectively. The mean EC50 values of imazalil-resistant isolates from Kecheng and Qujiang distracts were 2.07±1.04μg/mLand 2.35±0.73μg/mL, respectively,41.4 and 47 times higher than those of imazalil-sensitive isolates (MIC≤0.1μg/mL). In Kaihua, however, none imazalil-resistant isolates was detected, and the mean EC50 values of P. digitatum was 0.04±0.02μg/mL. The frequency of carbendazim-resistant isolates (MIC>10μg/mL) in Kecheng and Qujiang ware 54.3% and 54.2% whereas 9.1% in Kaihua. These results indicated that the frequencies of imazalil and carbendazim resistance of P. digitatum were much higher in main citrus producing areas (Kecheng and Qujiang districts) than that in non-main citrus producing areas (Kaihua).
To understand the resistance mechanism of P. digitatum to carbendaim, the sequences ofβ-tubulin gene (PdTUB2) of 8 Cab-S isolates and 10 Cab-R isolates were amplified and sequenced. Alignment of these sequences showed that the nucleotide at 1255 of PdTUB2 was thymine (T) in Cab-S isolates, whereas it was adenine (A) in Cab-R isolates, thus resulted in the change of amino acid at 200 from phenylalanine (F) in Cab-S isolates to tyrosine(Y) in Car-R isolates. No any other mutations correlated to Cab-R were found, thus indicated that the point mutatuion at 200aa from phenylalanine (F) to tyrosine (Y) inβ-tubulin gene must be responsive for resistance mechanism to carbendazim of P. digitatum from Quzhou.
Our previous study indicated that the insertion of a four-times of extra tandem repeat of the 126-bp transcriptional enhancer (termed IMZ-R1) and the insertion of a 199bp unique sequence within the 126bp transcriptional enhancer unit (termed IMZ-R2) in the promoter region of imazalil target gene CYP51 were associated with imazalil resistance of P.digitatum. Based on these machinsms, a pair of primers CYP51A1/CYP51A2 were designed and a PCR detection method was developped for distinguishing the IMZ-R1 and IMZ-R2 P. digitatum isolates. However, there was no expected amplicons were amplified from the imazalil-resistant isolates from Quzhou, and any point mutations in CYP51 coding regions or promoter regions correlated to the imazalil resistance were detected, suggesting that an unknown imazalil resistance machnisms (termed as IMZ-R3) must be presented in these imazalil resistant P. digitatum.
Two EST sequences, which were designated as CYP51B and CYP51C, were found in our annotated P. digitatum transcriptome database (unpublished). To obtain the full, as well as the partial sequences upstream and downstream of the PdCYP51B and PdCYP51C genes, primer pairs of CYP51B-R/CYP51B-F and CYP51C-R/CYP51C-F were designed according to the corresponding sequences in P. chrysogenum (NS_000201). Results indicated that the cloned fragment with CYP51B-R/CYP51B-F from IMZ-S isolate PdKH8 contained 2919bp, encompassing a complete coding region (1751bp) and its upstream (742bp) and downstream (425bp) sequences.This gene contained an opening reading frame (ORF) of 1751bp, with 3 introns of 73,51, and 52bp, located between positions 989bp to 1061bp,1260bp to 1310bp, and 2376bp to 2427bp, respectively.
PdCYP51C and partial sequences of its upstream and downstream were obtained using the same method. The cloned fragment with CYP51C-R/CYP51C-F has 2295nt, containing an ORF of 1497bp, with 5 introns located between 541bp to 596bp,682bp to 733bp,823bp to 885bp,1177bp to 1242bp, and 1292bp to 1348bp, respectively (data not shown). The nucleotide sequences of PdCYP51B and PdCYP51C genes were deposited in the GenBank as accession number HQ724322 and HQ724324.
Phylogenetic analysis showed that the deduced amino acid sequence of PdCYP51B was 78% and 61% identical to Aspergillus fumigatus (AfCYP51B, XP_749134.1) and Fusarium graminearum (FgCYP51B, FGSG_01000), respectively. The PdCYP51C showed 19% identity with Fusarium graminearum (FgCYP51C, FGSG_11024) and F. oxysporum (FoCYP51C, FOXG_13138). Moreover, PdCYP51B was 59% and 22% identical to PdCYP51A and PdCYP51C, respectively, while PdCYP51C was 23% identical to PdCYP51A.
PdCYP51B and PdCYP51C were amplified from eight IMZ-S isolates, one IMZ-R1, one IMZ-R2 and six IMZ-R3 isolates and sequenced. Alignment of nucleotide sequences of these PdCYP51B genes showed that an extra of 199bp fragment insertion was present of in promoter region in all 6 IMZ-R3 isolates, but was absent in both IMZ-R1 and IMZ-R2, as well as in all IMZ-S isolates. Apart from this difference, no any point mutation that related to imazalil resistance were detected either in the encoding or in the promoter regions of PdCYP51B of P. digitatum We speculate that this 199bp insertion is corelated to the observed resistance of IMZ-R3. So primer pairs of CYP51B-R/CYP51B-F were designed and used for PCR detected of IMZ-R3 P. digitatum. With this primer pair, the diagnostic band for IMZ-R3 were amplified from the imazalil resistant P. digitatum from Quzhou, indicating that the resistance molecular mechanism to imazalil of P. digitatum from Quzhou is the 199bp nucleotide-insertion mutation occurred in the promotor region of PdCYP51B.
本研究克隆测定8个多菌灵敏感菌株和10个多菌灵抗性菌株的β-微管蛋白基因全长序列,比对分析后发现敏感菌株的1255位核苷酸序列为TTC,其编码的氨基酸(200位)为苯丙氨酸,而在抗性菌株中相应位点的核苷酸为序列为TAC,编码的氨基酸为络氨酸。除此外,没有发现其他与抗性相关的突变。因此推断来自浙江衢州的柑橘绿霉病菌对多菌灵的抗性与其β-微管蛋白200位氨基酸由苯丙氨酸突变成络氨酸相关。
已知抑霉唑靶标基因(CYP51)启动区的4个额外的126bp片段简单串联重复插入(IMZ-R1)和其中一个199bp片段的插入(IMZ-R2)与柑橘绿霉菌抑霉唑抗性相关。使用检测区分这两种抗性分子机制的引物对CYP51A1/CYP51R2, PCR扩增发现,来自衢州的所有抗抑霉唑绿霉菌的抗性分子机制既不属于IMZ-R1也不属于IMZ-R2;测序比较敏感菌株和抗性菌株CYP51基因的全长序列,也未发现与抗性相关的点突变。
通过查找柑橘绿霉菌转录组数据库,发现两个CYP51同源基因的EST序列,命名为PdCYP51B和PdCYP51C基因。结合NCBI上产黄青霉(NS_000201)的同源物种信息,设计了CYP51B-R/CYP51B-F和CYP51C-R/CYP51C-F两对引物,成功从抑霉唑敏感菌株PdKH8菌株中克隆到包括启动子和编码区在内PdCYP51B和PdCYP51C两个全长基因。PdCYP51B包含2919bp,包括1751bp完整的编码区,以及它上游742bp的序列和下游425bp的序列,含有3个分别为73bp、51bp、52bp的外显子,分别位于989bp与1061bp之间,1260bp与1310bp之间,和2376bp与2427bp之间,编码485个氨基酸序列。PdCYP51B在GenBank中的登录号为HQ724322.PdCYP51C,包括一个1497bp的开放阅读框,含有分别位于541bp与596bp,682bp与733bp,823bp与885bp,1177bp与1242bp,和1292bp与1348bp之间的5个外显子,编码496个氨基酸序列。PdCYP51C在GenBank中的登录号为HQ724324。序列分析表明PdCYP51B基因与烟曲霉(AfCYP51B, XP_749134.1)和小麦赤霉病菌(FgCYP51B, FGSG_01000)有78%和61%一致性。PdCYP51C与禾谷镰刀菌(FgCYP51C, FGSG_11024)和尖孢镰刀菌(FoCYP51C, FOXG_13138)有19%的一致性。PdCYP51B与PdCYP51A和PdCYP51C有59%和22%的一致性,而PdCYP51C基因与PdCYP51A有23%的一致性。
克隆、测序并比较8个抑霉唑敏感(IMZ-S)和1个IMZ-R1、1个IMZ-R2,以及6个IMZ-R3菌株的CYP51B和CYP51C基因序列,发现这些菌株的CYP51C基本一致,未发现与抗性相关的点突变,但是,在所有IMZ-R3菌株的CYP51B基因启动子区均发现有一个199bp序列的插入,而该片段在所分析的IMZ-S、IMZ-R1和IMZ-R2菌株中均不存在。因此推测,IMZ-R3菌株的抗性机制与该菌株的CYP51基因启动子去的199bp插入相关。基于该分子机制,我们设计了一对检测IMZ-R3的引物,对来自衢州的IMZ-R3菌株进行PCR扩增,结果发现,能从所有这些抗性菌株中扩增到预期的片段。因此,认为衢州柑橘绿霉菌的抑霉唑抗性菌株的抗性分子机制均属于CYP51B基因启动子区有199bp序列的插入。
Penicillium digitatum is the most devastating pathogen of citrus fruit, being responsible for about 90% of production losses during post-harvest handling. Fungieide treatment is the main measure for postharvest disease control of citrus. However. the emergence and prevalence of fungicide-resistant biotypes followed by the successively and intensively using of fungieides often results in the decrease of control effect. In this study,70 isolates of P digitatum from 12 citrus storage houses in Quzhou city (included Kecheng and Qujiang districts, as well as Kaihua county), Zhejiang province, were assayed for their resistance to imazalil and carbendazim, and thier resistance machnisms were studied further. The results demonstrated that the frequency of imazalil-resistant isolates (minimum inhibitory concentration (MIC)≥0.5μg/mL) in Kecheng and Qujiang districts were 77.1% and 62.5%, respectively. The mean EC50 values of imazalil-resistant isolates from Kecheng and Qujiang distracts were 2.07±1.04μg/mLand 2.35±0.73μg/mL, respectively,41.4 and 47 times higher than those of imazalil-sensitive isolates (MIC≤0.1μg/mL). In Kaihua, however, none imazalil-resistant isolates was detected, and the mean EC50 values of P. digitatum was 0.04±0.02μg/mL. The frequency of carbendazim-resistant isolates (MIC>10μg/mL) in Kecheng and Qujiang ware 54.3% and 54.2% whereas 9.1% in Kaihua. These results indicated that the frequencies of imazalil and carbendazim resistance of P. digitatum were much higher in main citrus producing areas (Kecheng and Qujiang districts) than that in non-main citrus producing areas (Kaihua).
To understand the resistance mechanism of P. digitatum to carbendaim, the sequences ofβ-tubulin gene (PdTUB2) of 8 Cab-S isolates and 10 Cab-R isolates were amplified and sequenced. Alignment of these sequences showed that the nucleotide at 1255 of PdTUB2 was thymine (T) in Cab-S isolates, whereas it was adenine (A) in Cab-R isolates, thus resulted in the change of amino acid at 200 from phenylalanine (F) in Cab-S isolates to tyrosine(Y) in Car-R isolates. No any other mutations correlated to Cab-R were found, thus indicated that the point mutatuion at 200aa from phenylalanine (F) to tyrosine (Y) inβ-tubulin gene must be responsive for resistance mechanism to carbendazim of P. digitatum from Quzhou.
Our previous study indicated that the insertion of a four-times of extra tandem repeat of the 126-bp transcriptional enhancer (termed IMZ-R1) and the insertion of a 199bp unique sequence within the 126bp transcriptional enhancer unit (termed IMZ-R2) in the promoter region of imazalil target gene CYP51 were associated with imazalil resistance of P.digitatum. Based on these machinsms, a pair of primers CYP51A1/CYP51A2 were designed and a PCR detection method was developped for distinguishing the IMZ-R1 and IMZ-R2 P. digitatum isolates. However, there was no expected amplicons were amplified from the imazalil-resistant isolates from Quzhou, and any point mutations in CYP51 coding regions or promoter regions correlated to the imazalil resistance were detected, suggesting that an unknown imazalil resistance machnisms (termed as IMZ-R3) must be presented in these imazalil resistant P. digitatum.
Two EST sequences, which were designated as CYP51B and CYP51C, were found in our annotated P. digitatum transcriptome database (unpublished). To obtain the full, as well as the partial sequences upstream and downstream of the PdCYP51B and PdCYP51C genes, primer pairs of CYP51B-R/CYP51B-F and CYP51C-R/CYP51C-F were designed according to the corresponding sequences in P. chrysogenum (NS_000201). Results indicated that the cloned fragment with CYP51B-R/CYP51B-F from IMZ-S isolate PdKH8 contained 2919bp, encompassing a complete coding region (1751bp) and its upstream (742bp) and downstream (425bp) sequences.This gene contained an opening reading frame (ORF) of 1751bp, with 3 introns of 73,51, and 52bp, located between positions 989bp to 1061bp,1260bp to 1310bp, and 2376bp to 2427bp, respectively.
PdCYP51C and partial sequences of its upstream and downstream were obtained using the same method. The cloned fragment with CYP51C-R/CYP51C-F has 2295nt, containing an ORF of 1497bp, with 5 introns located between 541bp to 596bp,682bp to 733bp,823bp to 885bp,1177bp to 1242bp, and 1292bp to 1348bp, respectively (data not shown). The nucleotide sequences of PdCYP51B and PdCYP51C genes were deposited in the GenBank as accession number HQ724322 and HQ724324.
Phylogenetic analysis showed that the deduced amino acid sequence of PdCYP51B was 78% and 61% identical to Aspergillus fumigatus (AfCYP51B, XP_749134.1) and Fusarium graminearum (FgCYP51B, FGSG_01000), respectively. The PdCYP51C showed 19% identity with Fusarium graminearum (FgCYP51C, FGSG_11024) and F. oxysporum (FoCYP51C, FOXG_13138). Moreover, PdCYP51B was 59% and 22% identical to PdCYP51A and PdCYP51C, respectively, while PdCYP51C was 23% identical to PdCYP51A.
PdCYP51B and PdCYP51C were amplified from eight IMZ-S isolates, one IMZ-R1, one IMZ-R2 and six IMZ-R3 isolates and sequenced. Alignment of nucleotide sequences of these PdCYP51B genes showed that an extra of 199bp fragment insertion was present of in promoter region in all 6 IMZ-R3 isolates, but was absent in both IMZ-R1 and IMZ-R2, as well as in all IMZ-S isolates. Apart from this difference, no any point mutation that related to imazalil resistance were detected either in the encoding or in the promoter regions of PdCYP51B of P. digitatum We speculate that this 199bp insertion is corelated to the observed resistance of IMZ-R3. So primer pairs of CYP51B-R/CYP51B-F were designed and used for PCR detected of IMZ-R3 P. digitatum. With this primer pair, the diagnostic band for IMZ-R3 were amplified from the imazalil resistant P. digitatum from Quzhou, indicating that the resistance molecular mechanism to imazalil of P. digitatum from Quzhou is the 199bp nucleotide-insertion mutation occurred in the promotor region of PdCYP51B.
引文
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EL-Gooraili M A and Kasheir G.(1983).Occurance of Thiabendazole-Resistant strains of Penicillium italicum in Egypt.Plant diease.67(6):618-619.
Fujimura M, Kanakura T, Yamaguchi I. (1992).Action mechanism of diethofencarb to a benzimidazole-resistant mutant in Neurospora crassa.Pesti Sci.17:237.
Georgopapadakou N H, Walsh T J.(1994).Human mycoses-drugs and targets for emerging pathogens.Science.264:371-373.
Ghosoph J M,Schmidt L S,Margosan D A,Smilanick J L.(2007).Imazalil resistance linked to a unique insertion sequence in the PdCYP51 promoter region of Penicillium digitatum.Postharvest Biol Tec.44:9-18.
Gisi U, Hermann D, Leonie O and Christoph S. (1997).Sensitivity Profiles of Mycosphaerella graminicola and Phytophthora infestans Populations to Different Classes of Fungicides.Pestic.Sci.51:290-298.
Godwin R,Young J E,Woodward D J,et al.(1997).Azoxystrobin:Effects on the Development of Grapevine Downy Mildew(Plasmopara viticola).Proc-ANNP Cinquieme Conference Internationale sur les Maladies des Plants,Tours,3-5Deeember.871-878.
Gutter A, Shachnai A, Schiffmann-Nadel M and Dinoor A. (1981). Biological aspects of citrus molds tolerant to benzaimidazole fungiacides.Phytopathology.71:482-487.
Gutter Y.(1973).Benzimidazole resistant strains of citrus fruit pathogens. Research Summaries,1971-1973. The Volcani Center,Israel.56-57.
Hamamoto H, Hasegawa K, Nakaune R,Lee Y J, Makizumi Y, Akutsu K, Hibi T.(2000).Tandem repeat of a transcriptional enhancer upstream of the sterol 14 alpha-demethylase gene (CYP51) in Penicillium digitatum.Appl Environ Microb.66:3421-3426.
Hamamoto H, Nawata O, et al. (2001). The role of the ABC transporter gene PMR1 in demethylation inhibitor resistance in Penicillium digitatum. Pesticide Biochemistry and Physiology.70(1):19-26.
Holmes G J, Eckert J W.(1995).Relative fitness of imazalil-resistant and sensitive biotypes of Penicillium digitatum.Plant Dis.79:1068-1073.
Ji H, Zhang W, et al.(2003).Structure-based de novo design, synthesis, and biological evaluation of non-azole inhibitors specific for lanosterol 14alpha-demethylase of fungi. Med Chem.46(4):474-485.
Johnson K B.(1994).Frequency of benzimidazole and dicarboximide-resistant strains of Botrytis cinerea in Western Orgon small fruit Snap Bean plantings.Plant Disease.78:572-577.
Joseph-Horne T and Derek W H.(1997).Molecular mechaniisms of azole resistance in fungi.FEMS Microbiology Letters.149:141-149.
Jung M K, Oakley B R.(1990). Identification of an amino acid substitution in the benA β-tubulin gene of Aspergillus nidulans that confers thiabendazole resistance and benomyl supersensitivity. Cell Motility and the Cytoskeleton.17:87-94.
Kanetis L.(2007).Comparative Efficacy of the New Postharvest Fungicides Azoxystrobin,Fludioxonil,and Pyrimethanil for Managing Citrus Green Mold.Plant Disease.(91)11:1502-1510.
Kanetis L.(2008).Baseline Sensitivities for New Postharvest Fungicides Against Penicillium spp.on Citrus and Multiple Resistance Evaluations in P.digitatum.Plant Disease.92(2):301-309.
Koenraadt H,Somerville S C,Jones A L.(1992).Characterization of mutations in the β-tubulin gene of benomyl-resistant field strains of Venturia inaequalis and other plant pathogenic fungi.Mol Plant Pathol.82:1348-1354.
Koller W.(1987).Isomers of sterol synthesis inhibitors:Funigicidal effects and plant growth regulartor activities.Pestic.Sci.18:129-147.
Kuramoto T.(1976).Resistance to benomyl and thiophanate-mythyl in strains of Penicillium digitatum and P.italicum in Japan.Plant Dis Reptr.60:168-172.
Lepesheva G I, Waterman M R.(2004).CYP51 the omnipotent P450.Mol Cell Endocrinol.215(1-2):165-170.
Lepesheva G I,Waterman M R.(2007).Sterol 14 alpha-demethylase cytochrome P450 (CYP51),a P450 in all biological kingdoms.Biochimica Et Biophysica Acta-General Subjects.1770(3):467-477.
Liu X,Yu F,Schnabel G,Wu J,Wang Z,Ma Z.(2011).Paralogous cyp51 genes in Fusarium graminearum mediate differential sensitivity to sterol demethylation inhibitors.Fungal Genet Biol.48:113-123.
Lupetti A, Danesi R, et al.(2002).Molecular basis of resistance to azole antifungals.Trends Mol Med.8(2):76-81.
Luo C X and Schnabel G.(2008).The Cytochrome P450 Lanosterol 14-Demethylase Gene Is a Demethylation Inhibitor Fungicide Resistance Determinant in Monilinia fructicola Field Isolates from Georgia. Applied And environmental microbiology.74(2):359-366.
Ma Z, Proffer T J,et al. (2006). Overexpression of the 14alpha-demethylase target gene (CYP51) mediates fungicide resistance in Blumeriella jaapii. Appl Environ Microbiol.72(4):2581-2585.
Maarten A,DeWaard,Alan C A,et al.(2006),Review Impact of fungal drug transporters on fungicide sensitivity, multidrug resistance and virulence.Pest Manag Sci,62:195-207.
Mcdoald R E,Rises L A and Hillebrand B M.(1979).Resistance to thiabendazole citrus fruits from several countries.J.Am.Soc.Hortic.Sci.104:333-335.
Muirhead I F.(1974).Resistance to benzimidazole fungicides in blue mould of citrus in Queensland.Aus.J.Exp.Agric.Anim.Husb.14:698-701.
Nakaune R, Hamamoto H, et al.(2002). A novel ABC transporter gene, PMR5, is involved in multidrug resistance in the phytopathogenic fungus Penicillium digitatum. Molecular.Genetics and Genomics.267(2):179-185.
Nakaune R, Adachi K, Nawata O, et al.(1998). A novel ATP-binding cassette transporter involved in multidrug resistance in the phytopathogenic fungus Penicillium digitatum. Appl Environ Microbiol.64:3983-3988.
Paloma A, Carmen AN, et al.(2011).Granulocytic sarcoma of the small bowel, greater omentum and peritoneum associated with a CBFβ/MYH11 fusion and inv(16) (p13q22):a case report.International Archives of Medicine.4(3):1-5.
Pierre L, Rene F, Daniele D, Catherine A,et al.(2002).Mechanisms of resistance to fungicides in field strains of Botrytis cinerea.Pest Management Science.58:876-888.
Robbertse B,Vander M, et al.(2001).DMI sensitivity and cross-resistance patterns of Rhynchosporium secalis isolates from South Africa.Crop Protection.97-102.
Roohparvar R, Mehrabi R, Johannes G M,Van N,et al.(2008).The drug transporter MgMfsl can modulate sensitivity of field strains of the fungal wheat pathogen Mycosphaerella graminicola to the strobilurin fungicide trifloxystrobin.Pest Management Science.64:685-693.
Rebellato L H, and Monteiro M C.(1984).Report of an imazalil resistant strain of Penicillium digitatum (Sacc) in citrus in Uruguay. Proc. Int.Soc. Citric. 1:588-590.
Russell D J, Graeme W, Huby, Carlile,et al.(1995).Interactions between the Protein-tyrosine Kinase ZAP-70, the Proto-oncoprotein Vav, and Tubulin in Jurkat T Cells. Biological Chemistry.270(51):30241-30244.
Sanchez-Torres P, Tuset J J.(2011).Molecular insights into fungicide resistance in sensitive and resistant Penicillium digitatum strains infecting citrus. Postharvest Biol Tec.59:159-165.
Schirra M D,Aquino S, Palma A, et al.(2005).Residue Level,Persistence, and Storage Performance of Citrus Fruit Treated with Fludioxonil. Agric. Food Chem.53: 6718-6724.
Schepers H T.(1985).Changes during a three-year period in the sensitivity to ergosterol biosynthesis inhibitors of Sphaerotheca fuliginea in the Netherlands.Neth.J.P1.Path.91:105-118.
Schmidt L S,Ghosoph J M,Margosan D A,et al.(2006).Mutation at β-tubulin codon 200 indicated thiabendazole resistance in Penicillium digitatum collected from California citrus packinghouses.Plant Dis.90:765-770.
Schnabel G. and Jones A L.(2001).The 14alpha-Demethylasse(CYP51A1) Gene is Overexpressed in Venturia inaequalis Strains Resistant to Myclobutanil. Phytopathology 91(1):102-110.
Scholberg P L,Harlton C,Haag P,et al.(2005).Benzimidazole and diphenylamine sensitivity and identity of Penicllium spp.that cause postharvest blue mold of apples using β-tubulin gene sequences.Postharv Biol Technol.36:41-49.
Smilanick J L, Mansour M F, et al.(2005).Influence of pH and NaHCO3 on Effectiveness of Imazalil to Inhibit Germination of Penicillium digitatum and to Control Postharvest Green Mold on Citrus Fruit.Plant Disease.(89):640-645.
Smilanick J L, Mansour M F, Gabler F M, Goodwine W R.(2006).The effectiveness of pyrimethanil to inhibit germination of Penicillium digitatum and to control citrus green mold after harvest. Postharvest Biol Tec 42:75-85.
Smoot J J and Brown G E.(1974).Occurance of benzaimidazole-resistant strains of Penicillium digitatum in Florida citrus Packinghouse.Plant diease.58:933-934.
Spalding D H.(1982).Resistance of mango pathogensto fungicides used to control postharvest diseases.PlantDiseases.66:1185-1186.
Stanis V F and Jones A L.(1985).Reduced Sensitivity to Sterol-Inhibiting Fungicides in Field Isolates of Venturia inaequalis.Diease Control and Pest Management.75(10)1098-1102.
Wild B L and Rippon L E.(1975).Repponse of Penicillium digitatum strains to benomyl thiabendazole and sodium O-phenylphenate. Phytopathology.69:1176-1177.
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Eckert J W,Sievert J R,Ratnayake M.(1994).Reduction of imazalil effectiveness against citrus green mold in california packinghouses by resistant biotypes of Penicillium digitatum.Plant Dis.78:971-974.
Eckert J W.(1987).Penicillium digitatum biotypes with reduced sensitivity to imazalil.Phytopathology.77:1728-1728.
EL-Gooraili M A and Kasheir G.(1983).Occurance of Thiabendazole-Resistant strains of Penicillium italicum in Egypt.Plant diease.67(6):618-619.
Fujimura M, Kanakura T, Yamaguchi I. (1992).Action mechanism of diethofencarb to a benzimidazole-resistant mutant in Neurospora crassa.Pesti Sci.17:237.
Georgopapadakou N H, Walsh T J.(1994).Human mycoses-drugs and targets for emerging pathogens.Science.264:371-373.
Ghosoph J M,Schmidt L S,Margosan D A,Smilanick J L.(2007).Imazalil resistance linked to a unique insertion sequence in the PdCYP51 promoter region of Penicillium digitatum.Postharvest Biol Tec.44:9-18.
Gisi U, Hermann D, Leonie O and Christoph S. (1997).Sensitivity Profiles of Mycosphaerella graminicola and Phytophthora infestans Populations to Different Classes of Fungicides.Pestic.Sci.51:290-298.
Godwin R,Young J E,Woodward D J,et al.(1997).Azoxystrobin:Effects on the Development of Grapevine Downy Mildew(Plasmopara viticola).Proc-ANNP Cinquieme Conference Internationale sur les Maladies des Plants,Tours,3-5Deeember.871-878.
Gutter A, Shachnai A, Schiffmann-Nadel M and Dinoor A. (1981). Biological aspects of citrus molds tolerant to benzaimidazole fungiacides.Phytopathology.71:482-487.
Gutter Y.(1973).Benzimidazole resistant strains of citrus fruit pathogens. Research Summaries,1971-1973. The Volcani Center,Israel.56-57.
Hamamoto H, Hasegawa K, Nakaune R,Lee Y J, Makizumi Y, Akutsu K, Hibi T.(2000).Tandem repeat of a transcriptional enhancer upstream of the sterol 14 alpha-demethylase gene (CYP51) in Penicillium digitatum.Appl Environ Microb.66:3421-3426.
Hamamoto H, Nawata O, et al. (2001). The role of the ABC transporter gene PMR1 in demethylation inhibitor resistance in Penicillium digitatum. Pesticide Biochemistry and Physiology.70(1):19-26.
Holmes G J, Eckert J W.(1995).Relative fitness of imazalil-resistant and sensitive biotypes of Penicillium digitatum.Plant Dis.79:1068-1073.
Ji H, Zhang W, et al.(2003).Structure-based de novo design, synthesis, and biological evaluation of non-azole inhibitors specific for lanosterol 14alpha-demethylase of fungi. Med Chem.46(4):474-485.
Johnson K B.(1994).Frequency of benzimidazole and dicarboximide-resistant strains of Botrytis cinerea in Western Orgon small fruit Snap Bean plantings.Plant Disease.78:572-577.
Joseph-Horne T and Derek W H.(1997).Molecular mechaniisms of azole resistance in fungi.FEMS Microbiology Letters.149:141-149.
Jung M K, Oakley B R.(1990). Identification of an amino acid substitution in the benA β-tubulin gene of Aspergillus nidulans that confers thiabendazole resistance and benomyl supersensitivity. Cell Motility and the Cytoskeleton.17:87-94.
Kanetis L.(2007).Comparative Efficacy of the New Postharvest Fungicides Azoxystrobin,Fludioxonil,and Pyrimethanil for Managing Citrus Green Mold.Plant Disease.(91)11:1502-1510.
Kanetis L.(2008).Baseline Sensitivities for New Postharvest Fungicides Against Penicillium spp.on Citrus and Multiple Resistance Evaluations in P.digitatum.Plant Disease.92(2):301-309.
Koenraadt H,Somerville S C,Jones A L.(1992).Characterization of mutations in the β-tubulin gene of benomyl-resistant field strains of Venturia inaequalis and other plant pathogenic fungi.Mol Plant Pathol.82:1348-1354.
Koller W.(1987).Isomers of sterol synthesis inhibitors:Funigicidal effects and plant growth regulartor activities.Pestic.Sci.18:129-147.
Kuramoto T.(1976).Resistance to benomyl and thiophanate-mythyl in strains of Penicillium digitatum and P.italicum in Japan.Plant Dis Reptr.60:168-172.
Lepesheva G I, Waterman M R.(2004).CYP51 the omnipotent P450.Mol Cell Endocrinol.215(1-2):165-170.
Lepesheva G I,Waterman M R.(2007).Sterol 14 alpha-demethylase cytochrome P450 (CYP51),a P450 in all biological kingdoms.Biochimica Et Biophysica Acta-General Subjects.1770(3):467-477.
Liu X,Yu F,Schnabel G,Wu J,Wang Z,Ma Z.(2011).Paralogous cyp51 genes in Fusarium graminearum mediate differential sensitivity to sterol demethylation inhibitors.Fungal Genet Biol.48:113-123.
Lupetti A, Danesi R, et al.(2002).Molecular basis of resistance to azole antifungals.Trends Mol Med.8(2):76-81.
Luo C X and Schnabel G.(2008).The Cytochrome P450 Lanosterol 14-Demethylase Gene Is a Demethylation Inhibitor Fungicide Resistance Determinant in Monilinia fructicola Field Isolates from Georgia. Applied And environmental microbiology.74(2):359-366.
Ma Z, Proffer T J,et al. (2006). Overexpression of the 14alpha-demethylase target gene (CYP51) mediates fungicide resistance in Blumeriella jaapii. Appl Environ Microbiol.72(4):2581-2585.
Maarten A,DeWaard,Alan C A,et al.(2006),Review Impact of fungal drug transporters on fungicide sensitivity, multidrug resistance and virulence.Pest Manag Sci,62:195-207.
Mcdoald R E,Rises L A and Hillebrand B M.(1979).Resistance to thiabendazole citrus fruits from several countries.J.Am.Soc.Hortic.Sci.104:333-335.
Muirhead I F.(1974).Resistance to benzimidazole fungicides in blue mould of citrus in Queensland.Aus.J.Exp.Agric.Anim.Husb.14:698-701.
Nakaune R, Hamamoto H, et al.(2002). A novel ABC transporter gene, PMR5, is involved in multidrug resistance in the phytopathogenic fungus Penicillium digitatum. Molecular.Genetics and Genomics.267(2):179-185.
Nakaune R, Adachi K, Nawata O, et al.(1998). A novel ATP-binding cassette transporter involved in multidrug resistance in the phytopathogenic fungus Penicillium digitatum. Appl Environ Microbiol.64:3983-3988.
Paloma A, Carmen AN, et al.(2011).Granulocytic sarcoma of the small bowel, greater omentum and peritoneum associated with a CBFβ/MYH11 fusion and inv(16) (p13q22):a case report.International Archives of Medicine.4(3):1-5.
Pierre L, Rene F, Daniele D, Catherine A,et al.(2002).Mechanisms of resistance to fungicides in field strains of Botrytis cinerea.Pest Management Science.58:876-888.
Robbertse B,Vander M, et al.(2001).DMI sensitivity and cross-resistance patterns of Rhynchosporium secalis isolates from South Africa.Crop Protection.97-102.
Roohparvar R, Mehrabi R, Johannes G M,Van N,et al.(2008).The drug transporter MgMfsl can modulate sensitivity of field strains of the fungal wheat pathogen Mycosphaerella graminicola to the strobilurin fungicide trifloxystrobin.Pest Management Science.64:685-693.
Rebellato L H, and Monteiro M C.(1984).Report of an imazalil resistant strain of Penicillium digitatum (Sacc) in citrus in Uruguay. Proc. Int.Soc. Citric. 1:588-590.
Russell D J, Graeme W, Huby, Carlile,et al.(1995).Interactions between the Protein-tyrosine Kinase ZAP-70, the Proto-oncoprotein Vav, and Tubulin in Jurkat T Cells. Biological Chemistry.270(51):30241-30244.
Sanchez-Torres P, Tuset J J.(2011).Molecular insights into fungicide resistance in sensitive and resistant Penicillium digitatum strains infecting citrus. Postharvest Biol Tec.59:159-165.
Schirra M D,Aquino S, Palma A, et al.(2005).Residue Level,Persistence, and Storage Performance of Citrus Fruit Treated with Fludioxonil. Agric. Food Chem.53: 6718-6724.
Schepers H T.(1985).Changes during a three-year period in the sensitivity to ergosterol biosynthesis inhibitors of Sphaerotheca fuliginea in the Netherlands.Neth.J.P1.Path.91:105-118.
Schmidt L S,Ghosoph J M,Margosan D A,et al.(2006).Mutation at β-tubulin codon 200 indicated thiabendazole resistance in Penicillium digitatum collected from California citrus packinghouses.Plant Dis.90:765-770.
Schnabel G. and Jones A L.(2001).The 14alpha-Demethylasse(CYP51A1) Gene is Overexpressed in Venturia inaequalis Strains Resistant to Myclobutanil. Phytopathology 91(1):102-110.
Scholberg P L,Harlton C,Haag P,et al.(2005).Benzimidazole and diphenylamine sensitivity and identity of Penicllium spp.that cause postharvest blue mold of apples using β-tubulin gene sequences.Postharv Biol Technol.36:41-49.
Smilanick J L, Mansour M F, et al.(2005).Influence of pH and NaHCO3 on Effectiveness of Imazalil to Inhibit Germination of Penicillium digitatum and to Control Postharvest Green Mold on Citrus Fruit.Plant Disease.(89):640-645.
Smilanick J L, Mansour M F, Gabler F M, Goodwine W R.(2006).The effectiveness of pyrimethanil to inhibit germination of Penicillium digitatum and to control citrus green mold after harvest. Postharvest Biol Tec 42:75-85.
Smoot J J and Brown G E.(1974).Occurance of benzaimidazole-resistant strains of Penicillium digitatum in Florida citrus Packinghouse.Plant diease.58:933-934.
Spalding D H.(1982).Resistance of mango pathogensto fungicides used to control postharvest diseases.PlantDiseases.66:1185-1186.
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