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植物病原卵菌对重要抑制剂的抗性分子机制研究进展

来源：花匠小妙招时间：2024-12-17 18:23

摘要: 植物病原卵菌可导致多种毁灭性的植物病害，由于缺乏有效的抗病品种，目前化学防治仍是防治卵菌病害最有效的方法之一，但随着一些内吸性杀菌剂的不合理及频繁使用，抗性问题已日益突出。文章对目前主要的卵菌抑制剂苯酰胺类 (PAs)、羧酸酰胺类 (CAAs)、苯醌外部抑制剂类 (QoIs)、氧化固醇结合蛋白抑制剂类 (OSBPIs) 以及微管蛋白抑制剂类 (tubulin inhibitors) 的作用机制、抗性遗传机制、抗性分子机制的最新研究进展进行了综述，旨在为当前中国植物病原卵菌的抗性研究和防控提供参考。现有文献报道表明：5类重要卵菌抑制剂的抗性主要是由于病原菌中药剂靶标蛋白的点突变引起，且苯酰胺类及微管蛋白抑制剂的抗性由多个基因控制；不同羧酸酰胺类药剂的抗性遗传机制存在差异；苯醌外部抑制剂的抗性具有母系遗传的特征；推测氧化固醇结合蛋白抑制剂的抗性由单个显性基因控制。

Abstract: Plant pathogenic oomycetes could cause many devastating diseases. Currently, chemical control is still one of the most effective methods for the control of oomycete diseases. However, many fungicides against oomycetes have generated serious resistance problems because of unreasonable and frequent applications in fields. The mode of action, resistant genetic mechanism and molecular resistance mechanism of several most commonly used fungicides were reviewed, such as phenylamides (PAs), carboxylic acid amides (CAAs), quinone outside inhibitors (QoIs), oxysterol binding protein inhibitors (OSBPIs) and tubulin inhibitors. The current knowledge summaried in this review will contribute to the scientific control of plant pathogenic oomycetes and provide a valuable reference for future research. Existing literature reports demosntaretd that point mutation in target protein could cause resistance to PAs, CAAs, QoIs, OSBPIs and tubulin inhibitors. PAs and tubulin inhibitors resistance were controlled by more than one gene, and QoIs resisitance was maternal inheritance. It was speculated that OSBPIs resistance was controlled by one dominant gene. However, there was distinct difference in genetic mechanism for different CAAs fungicides.

图 1 与CAAs药剂抗性相关的不同卵菌纤维素合酶A3亚基突变位点

注：图中Pi、Pc、Pm、Ps、Pvi、Pcu分别为致病疫霉、辣椒疫霉、瓜类疫霉、大豆疫霉、葡萄霜霉及黄瓜霜霉。

Figure 1. Point mutation in CesA3 from different oomycetes that confers CAAs resistance

Note: Pi, Pc, Pm, Ps, Pvi, Pcu are Phytophthora infestans, Ph. capsici, Ph. melonis, Ph. sojae, Plasmopara viticola, Pseudoperonospora cubensis, respectively.

图 2 QoI类药剂抗性相关的Cytb突变位点

注：图中Ztr、Pvi、Bla、Pcu、Pli、Pin、Py分别代表小麦壳针孢叶枯菌、葡萄霜霉、莴苣霜霉、黄瓜霜霉、荔枝霜疫霉、致病疫霉及腐霉。

Figure 2. Point mutation in Cytb from different pathogens that confer QoIs resistance

Note: Ztr, Pvi, Bla, Pcu, Pli, Pin, Py are Zymoseptoria tritici, Plasmopara viticola, Bremia lactucae, Pseudoperonospora cubensis, Peronophythora litchi, Phytophthora infestans, Pythium, respectively.

图 3 与氟噻唑吡乙酮抗性相关的ORPs位点

注：图中Pi、Pc、Ps、Pu、Pa分别代表致病疫霉、辣椒疫霉、大豆疫霉、终极腐霉及瓜果腐霉。

Figure 3. Major point in ORPs from different oomycetes that relates with oxathiapiprolin resistance

Note: Pi, Pc, Ps, Pu, Pa are Phytophthora infestans, Ph. capsici, Ph. sojae, Pythium ultimum and Py. aphanidermatum, respectively.

表 1 卵菌与真菌的生理生化特征比较[3]

Table 1 Physiological and biochemical characteristics of oomycetes and fungi[3]

特征
Characteristic卵菌　　　　
Oomycetes　　　　真菌 (接合菌门、子囊菌门和担子菌门)
Fungi (zygo-, asco-, basidiomycetes) 染色体倍性
Ploidity 二倍体 (2 n，少有多倍体)
Diploid (2 n, rarely also polyploid) 单倍体 (n)
Haploid (n) 细胞壁结晶区组分
Cell wall, fibrillar fraction 纤维素 (10%~30%)
Cellulose (10%~30% of cell wall) 几丁质，壳聚糖
Chitin, chitosan 细胞壁非结晶区组分
Cell wall, amorphous fraction 1-3/1-6-β-葡聚糖 (70%~90%)
1-3/1-6-β-glucans (70%~90 % of cel wall) 1-3/1-6-β-葡聚糖
1-3/1-6-β-glucans 细胞膜功能甾醇
Cell membrane, functional sterol 胆固醇 (甾醇营养缺陷型)
Cholesterol, sterol auxotrophy 麦角甾醇
Ergosterol 赖氨酸生物合成途径
Lysine biosynthesis 二氨基庚二酸途径
DAP-pathway 氨基己二酸途径
AAP-pathway 菌丝隔膜
Hyphae (septa) 没有或仅有很少隔膜
No (few) septa 有隔膜
Regular septation 无性孢子阶段
Asexual spore stages 游动孢子 (孢子囊)；厚垣孢子；分生孢子
Zoospores (and sporangia), chlamydospores, conidia 分生孢子
Conidia 有性孢子阶段
Sexual spore stages 卵孢子
Oospores 接合孢子，子囊孢子，担孢子
Zygo-, asco-, basidiospores

表 2 卵菌与真菌对药剂的敏感性比较[3]

Table 2 Sensitivity of oomycetes and fungi to different fungicides[3]

杀菌剂
Fungicide卵菌
Oomycetes真菌 (接合菌门、子囊菌门和担子菌门)
Fungi (zygo-, asco-, basidiomycetes) 匹马菌素
pimaricin 不敏感
Tolerant 多数敏感
Mostly sensitive 苯并咪唑类
MBCs 不敏感
Tolerant 多数敏感 (抗性)
Mostly sensitive (resistant) 甾醇脱甲基抑制剂
DMIs 不敏感
Tolerant 多数敏感 (抗性)
Mostly sensitive (resistant) 苯酰胺类
PAs 敏感 (抗药性)
Sensitive (resistant) 不敏感
Tolerant 羧酸酰胺类
CAAs 霜霉目敏感 (抗药性)；其他卵菌不敏感
Sensitive (resistant) for Peronosporales, tolerant for other oomycetes 不敏感
Tolerant 氧化固醇结合蛋白抑制剂类
OSBPIs 敏感 (抗药性) (大部分 Pythium spp.
不敏感)
Sensitive (resistant) (most Pythium spp., tolerant) 不敏感
Tolerant 苯醌外抑制剂类
QoIs 敏感 (抗药性)
Sensitive (resistant) 多数敏感 (抗性)
Mostly sensitive (resistant) 苯醌内抑制剂类
QiIs 敏感 (抗药性)
Sensitive (resistant) 多数不敏感
Mostly tolerant 苯醌内外抑制剂类
QioIs 部分敏感 (抗药性)
Some sensitive (resistant) 不敏感
Tolerant

表 3 主要卵菌抑制剂种类[3-4]

Table 3 Major oomycetes inhibitors[3-4]

杀菌剂类别
Fungicide group作用机制　　　　　
Mode of action　　　　　主要有效成分　　　　　
Major active ingredients　　　　杀菌剂抗性委员会代码
FRAC code抗性风险
Resistant risk 苯酰胺类
PAs RNA聚合酶 I
RNA polymerase I 甲霜灵 (精甲霜灵)、苯霜灵 (精苯霜灵)、呋霜灵等
metalaxyl (mefenoxam), benalaxyl (kiralaxyl), furalaxyl etc. 4 (A1) 高等
High 苯醌外抑制剂类
QoIs 细胞色素 bc1 复合物
Qo 位点
Cytochrome bc1 Qo site 嘧菌酯、吡唑醚菌酯、唑菌酮等
azoxystrobin, pyraclostrobin; famoxadone etc. 11 (C3) 高等
High 羧酸酰胺类
CAAs 纤维素合酶
Cellulose synthase 烯酰吗啉、异丙菌胺、双炔酰菌胺等
dimethomorph, iprovalicarb, mandipropamid etc. 40 (H5) 低到中等
Low to medium 氧化固醇结合蛋白抑制剂类
OSBPIs 氧化固醇结合蛋白同源蛋白
OSBP homologue 氟噻唑吡乙酮
oxathiapiprolin 49 (F9) 中到高等
Medium to high 苯甲酰胺类、噻唑甲酰胺类
Benzamides, thiazole-carboxamide β-微管蛋白
β-tubulin 苯酰菌胺、噻唑菌胺
zoxamide, ethaboxam 22 (B3) 低到中等
Low to medium 苯甲酰胺类
Benzamides 血影蛋白类似蛋白
Spectrin-like protein 氟吡菌胺
fluopicolide 43 (B5) 中等
Medium 硝基苯胺类
Dinitroanilines 氧化磷酸化解偶联剂
Uncouplers of oxidative phosphorylation 氟啶胺
fluazinam 29 (C5) 低等
Low 苯醌内抑制剂类
QiIs 细胞色素 bc1 复合物 Qi 位点
Cytochrome bc1 Qi site 氰霜唑、吲唑磺菌胺
cyazofamid, amisulbrom 21 (C4) 中到高等
Medium to high 苯醌内外抑制剂类
QioIs 细胞色素 bc1 复合物 Qo 和
Qi 位点
Cytochrome bc1 Qi and Qo site 唑嘧菌胺
ametoctradin 45 (C8) 中到高等
Medium to high 磷酸盐类
Phosphonates 植物诱抗剂
Host plant defence induction 乙磷铝
fosetyl-al P 07 (P) 低等
Low 氰基乙酰胺肟类
Cyanoacetamide-oxime 未知
Unknown 霜脲氰
cymoxanil 27 (U) 低到中等
Low to medium 其他
Others 多作用位点
Multisites 代森锰锌、百菌清、铜制剂等
mancozeb, chlorothalonil, copper compounds etc. M1-M5 低等
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