小麦赤霉病菌甲硫氨酸代谢途径中关键基因的生物学功能研究
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摘要
小麦赤霉病菌是农业生产上极为重要的植物病原菌之一,由其引起的小麦赤霉病是小麦和大麦上的主要病害之一,不仅对麦类粮食生产造成巨大的损失,而且病麦粒中含有多种毒素直接威胁人畜生命安全。近年来,小麦赤霉病菌对常用杀菌剂的抗性逐年加重,因此,迫切需要开发新药剂用于赤霉病的化学防治。甲硫氨酸的代谢在真菌的生长和致病等方面起重要作用,而且人和动物体内没有甲硫氨酸合成途径,因此,研究小麦赤霉病菌甲硫氨酸代谢途径中的关键元件对开发新型杀菌剂具有重要的理论和现实意义。
本课题利用原生质体转化和农杆菌介导的真菌转化系统,通过基因敲除和互补的方法研究了小麦赤霉病菌甲硫氨酸合成途径上4个关键元件(胱硫醚γ-合成酶FgCGS、丝氨酸乙酰转移酶FgSAT、高丝氨酸乙酰转移酶FgHAT和胱硫醚β-裂解酶FgCBL)和氧化还原过程中3个关键元件(甲硫氨酸亚砜还原酶FgMsrA、 FgMsrB和FgfMsr)的生物学功能,结果发现：
1) FgCGS、FgSAT、FgHAT和FgCBL的缺失突变体在PDA培养基上表现菌落变黄或气生菌丝减少。在FGA培养基上,FgCGS、FgHAT和FgCBL的缺失突变体表现甲硫氨酸营养缺陷型,外源添加1mM甲硫氨酸或0.5mg/ml高半胱氨酸可恢复它们的生长,而FgSAT的缺失不会引起甲硫氨酸营养缺陷,但使菌体的气生菌丝明显减少。产孢量测定发现,FgCGS、FgSAT和FgCBL缺失突变体在绿豆培养液中的产孢量与野生菌株没有显著差异,而FgHAT缺失突变体的产孢与营养有关。孢子萌发分析结果显示,在2%蔗糖水溶液中,FgCGS缺失突变体的孢子萌发率显著低于野生菌株,而添加1mM甲硫氨酸或0.5mg/ml高半胱氨酸后使孢子萌发率恢复正常；FgSAT缺失突变体的孢子萌发率比野生菌株稍低,没有显著变化。致病力测定发现,FgCGS、FgHAT和FgCBL的缺失突变体对小麦穗和番茄的致病力显著下降,而FgSAT缺失突变体的致病力没有明显变化。对FgCGS、FgHAT和FgCBL缺失突变体在麦穗培养基上的生长测定发现,与野生菌株相比,这些突变体的气生菌丝明显减少。Real-time PCR分析表明,编码毒素脱氧雪腐镰刀菌烯醇(DON)合成的基因Tri5表达量显著下降。对FgCGS缺失突变体中毒素DON合成量的进一步测定发现,与野生菌株相比,该突变体中DON的合成量下降了约4倍。对药剂敏感性的测定发现,与野生菌株相比,FgCGS, FgSAT, FgHAT和FgCBL的缺失突变体对嘧霉胺的敏感性都没有显著变化,这表明,FgCGS、FgSAT、FgHAT和FgCBL不可能为嘧霉胺在小麦赤霉病菌中的作用靶标。FgCGS和FgHAT的缺失突变体对DMI类药剂的敏感性显著增加,并通过Real-time PCR进一步验证发现,FgCGS缺失突变体中DMI类药剂靶标基因CYP51A/B/C的表达量下降。胁迫压力敏感性测定发现,FgCGS、FgSAT、FgHAT和FgCBL的缺失突变体对渗透压、氧化压力、细胞壁(膜)压力的敏感性与野生菌株没有显著差异。这些结果表明,FgCGS、FgSAT、 FgHAT和FgCBL在小麦赤霉病菌的生长发育和致病过程中起关键作用。
2)对FgMsrA、FgMsrB和FgfMsr缺失突变体生长表型的测定发现,与野生菌株相比,这些突变体在PDA和FGA培养基上的生长正常,在绿豆培养液中的产孢也正常。致病力测定发现,FgMsrA、FgMsrB和FgfMsr缺失突变体在小麦穗和番茄上的致病力与野生菌株没有明显差异。药剂敏感性测定发现,与野生菌株相比,FgMsrA、FgMsrB和FgfMsr缺失突变体对嘧霉胺、DMI类药剂和多菌灵的敏感性都没有明显变化,这表明,FgMsrA、FgMsrB和FgfMsr不可能为嘧霉胺的作用靶标。氧化压力敏感性测定发现,FgMsrA、FgMsrB和FgfMsr缺失突变体对氧化压力的敏感性也没有明显变化,但在氧化压力下,FgMsrA、FgMsrB和FgfMsr基因的表达量显著上升,这些结果表明,FgMsrA、FgMsrB和FgfMsr在小麦赤霉病菌的生长发育和致病力方面的作用不明显,但可能参与菌体对氧化压力的反应过程。
Fusarium graminearum Schw. is the primary causal agent of Fusarium head blight (FHB) on wheat and barley. Infection of cereal crops with F. graminearum may lead to huge yield loss in severe epidemic years. More importantly, the trichothecene mycotoxins in infected grains pose a serious threat to human and animal health. Currently, emerging fungicide resistance of F. graminearum makes it urgent to explore new compounds for FHB management. Methionine metabolism has important roles in fungal growth, pathogenicity and et al. Moreover, methionine biosynthesis pathway is absent in non-ruminant animals, research on key elements of methionine metabolism in F. graminearum shall contribute to novel fungicide development.
In this study, target gene disruption and complementary strategy were employed to investigate functions of four key methionine biosynthesis elements (cystathionine y-synthase FgCGS, serine acetyltransferase FgSAT, homoserine O-acetyltransferase FgHAT and cystathionine β-lyase FgCBL) and three key elements (methionine sulfoxide reductases, FgmsrA, FgmsrB and FgfMsr) involved in methionine oxidation and reduction in F. graminearum. Results of our study demonstrated that:
1) Mycelial growth was tested in different conditions. On PDA, compared with wild-type parent, FgCGS, FgSAT, FgHAT and FgCBL deletion mutants showed yellowish or fewer aerial mycelia. On FGA, FgCGS, FgHAT and FgCBL deletion mutants could not grow without supplementation of ether methionine (1mM) or homocysteine (0.5mg/ml). FgSAT deletion mutant showed fewer aerial mycelia, though the radial growth was not distinct from that of wild-type parent. Conidiation assays indicated that FgCGS, FgSAT and FgCBL deletion mutants produced as many conidia as wild-type parent, while conidiation in FgHAT deletion mutant was determined by the nutrient of media. However, compared with wild-type parent, conidial germination rate of FgCGS deletion mutant in2%sucrose solution reduced significantly, but was rescued by supplementation of methionine (1mM) or homocysteine (0.5mg/ml). Conidial germination ability of FgSAT deletion mutant reduced a little. Inoculation tests showed that FgCGS, FgHAT and FgCBL deletion mutants exhibited decreased virulence significantly on wheat heads, while the virulence of FgSAT deletion mutant was comparable to wild-type parent. FgCGS, FgSAT, FgHAT and FgCBL deletion mutants showed fewer aerial mycelia in wheat ear culture, and expression of Tri5gene was decreased significantly, compared with wild-type parent. Deoxynivalenol (DON) production of FgCGS deletion mutant was4-fold less than wild-type parent. Fungicide evaluation assays revealed that sensitivity of FgCGS, FgSAT, FgHAT and FgCBL deletion mutants to anilinopyrimidine fungicide pyrimethanil was not altered, indicating that FgCGS, FgSAT, FgHAT and FgCBL are not likely to be the target of pyrimethanil in F. graminearum. FgCGS and FgHAT deletion mutants showed increased sensitivity to sterol demethylation inhibitors (DMI), which was consistent with a low level of CYP51A/B/C expression in FgCGS deletion mutant. Sensitivity to osmotic, oxidative and cell wall (membrane) stresses revealed that FgCGS, FgSAT, FgHAT and FgCBL deletion mutants did not show observable changes in mycelia growth compared to wild-type parent, which indicate FgCGS, FgSAT, FgHAT and FgCBL are not involved in these stress responses. These results indicate that FgCGS, FgSAT, FgHAT and FgCBL play important roles in the growth, conidiation and pathogenicity of F. graminearum.
2) Mycelial growth tests showed FgMsrA, FgMsrB and FgfMsr deletion mutants had normal growth on PDA and FGA, and produced as many conidia in MBL medium as wild-type parent. Inoculation tests showed that the virulence of FgMsrA、FgMsrB and FgfMsr deletion mutants was comparable to wild-type parent. Fungicide sensitivity assays revealed that FgMsrA、FgMsrB and FgfMsr deletion mutants did not change the sensitivity to pyrimethanil, DMIs and carbendazim, indicating that FgMsrA、FgMsrB and FgfMsr may not be the target of pyrimethanil in F. graminearum. Oxidative stress sensitivity tests revealed that FgMsr A、FgMsr B and FgfMsr deletion mutants did not show observable changes in mycelia growth compared to wild-type parent. However, relative expression of FgMsr A、FgMsrB and FgfMs was increased significantly under oxidative stress. These results suggest that FgMsrA、FgMsrB and FgfMsr may be involed in oxidative stress reaction process of F. graminearum, though their roles in the growth, conidiation and pathogenicity are not obvious.
本课题利用原生质体转化和农杆菌介导的真菌转化系统,通过基因敲除和互补的方法研究了小麦赤霉病菌甲硫氨酸合成途径上4个关键元件(胱硫醚γ-合成酶FgCGS、丝氨酸乙酰转移酶FgSAT、高丝氨酸乙酰转移酶FgHAT和胱硫醚β-裂解酶FgCBL)和氧化还原过程中3个关键元件(甲硫氨酸亚砜还原酶FgMsrA、 FgMsrB和FgfMsr)的生物学功能,结果发现：
1) FgCGS、FgSAT、FgHAT和FgCBL的缺失突变体在PDA培养基上表现菌落变黄或气生菌丝减少。在FGA培养基上,FgCGS、FgHAT和FgCBL的缺失突变体表现甲硫氨酸营养缺陷型,外源添加1mM甲硫氨酸或0.5mg/ml高半胱氨酸可恢复它们的生长,而FgSAT的缺失不会引起甲硫氨酸营养缺陷,但使菌体的气生菌丝明显减少。产孢量测定发现,FgCGS、FgSAT和FgCBL缺失突变体在绿豆培养液中的产孢量与野生菌株没有显著差异,而FgHAT缺失突变体的产孢与营养有关。孢子萌发分析结果显示,在2%蔗糖水溶液中,FgCGS缺失突变体的孢子萌发率显著低于野生菌株,而添加1mM甲硫氨酸或0.5mg/ml高半胱氨酸后使孢子萌发率恢复正常；FgSAT缺失突变体的孢子萌发率比野生菌株稍低,没有显著变化。致病力测定发现,FgCGS、FgHAT和FgCBL的缺失突变体对小麦穗和番茄的致病力显著下降,而FgSAT缺失突变体的致病力没有明显变化。对FgCGS、FgHAT和FgCBL缺失突变体在麦穗培养基上的生长测定发现,与野生菌株相比,这些突变体的气生菌丝明显减少。Real-time PCR分析表明,编码毒素脱氧雪腐镰刀菌烯醇(DON)合成的基因Tri5表达量显著下降。对FgCGS缺失突变体中毒素DON合成量的进一步测定发现,与野生菌株相比,该突变体中DON的合成量下降了约4倍。对药剂敏感性的测定发现,与野生菌株相比,FgCGS, FgSAT, FgHAT和FgCBL的缺失突变体对嘧霉胺的敏感性都没有显著变化,这表明,FgCGS、FgSAT、FgHAT和FgCBL不可能为嘧霉胺在小麦赤霉病菌中的作用靶标。FgCGS和FgHAT的缺失突变体对DMI类药剂的敏感性显著增加,并通过Real-time PCR进一步验证发现,FgCGS缺失突变体中DMI类药剂靶标基因CYP51A/B/C的表达量下降。胁迫压力敏感性测定发现,FgCGS、FgSAT、FgHAT和FgCBL的缺失突变体对渗透压、氧化压力、细胞壁(膜)压力的敏感性与野生菌株没有显著差异。这些结果表明,FgCGS、FgSAT、 FgHAT和FgCBL在小麦赤霉病菌的生长发育和致病过程中起关键作用。
2)对FgMsrA、FgMsrB和FgfMsr缺失突变体生长表型的测定发现,与野生菌株相比,这些突变体在PDA和FGA培养基上的生长正常,在绿豆培养液中的产孢也正常。致病力测定发现,FgMsrA、FgMsrB和FgfMsr缺失突变体在小麦穗和番茄上的致病力与野生菌株没有明显差异。药剂敏感性测定发现,与野生菌株相比,FgMsrA、FgMsrB和FgfMsr缺失突变体对嘧霉胺、DMI类药剂和多菌灵的敏感性都没有明显变化,这表明,FgMsrA、FgMsrB和FgfMsr不可能为嘧霉胺的作用靶标。氧化压力敏感性测定发现,FgMsrA、FgMsrB和FgfMsr缺失突变体对氧化压力的敏感性也没有明显变化,但在氧化压力下,FgMsrA、FgMsrB和FgfMsr基因的表达量显著上升,这些结果表明,FgMsrA、FgMsrB和FgfMsr在小麦赤霉病菌的生长发育和致病力方面的作用不明显,但可能参与菌体对氧化压力的反应过程。
Fusarium graminearum Schw. is the primary causal agent of Fusarium head blight (FHB) on wheat and barley. Infection of cereal crops with F. graminearum may lead to huge yield loss in severe epidemic years. More importantly, the trichothecene mycotoxins in infected grains pose a serious threat to human and animal health. Currently, emerging fungicide resistance of F. graminearum makes it urgent to explore new compounds for FHB management. Methionine metabolism has important roles in fungal growth, pathogenicity and et al. Moreover, methionine biosynthesis pathway is absent in non-ruminant animals, research on key elements of methionine metabolism in F. graminearum shall contribute to novel fungicide development.
In this study, target gene disruption and complementary strategy were employed to investigate functions of four key methionine biosynthesis elements (cystathionine y-synthase FgCGS, serine acetyltransferase FgSAT, homoserine O-acetyltransferase FgHAT and cystathionine β-lyase FgCBL) and three key elements (methionine sulfoxide reductases, FgmsrA, FgmsrB and FgfMsr) involved in methionine oxidation and reduction in F. graminearum. Results of our study demonstrated that:
1) Mycelial growth was tested in different conditions. On PDA, compared with wild-type parent, FgCGS, FgSAT, FgHAT and FgCBL deletion mutants showed yellowish or fewer aerial mycelia. On FGA, FgCGS, FgHAT and FgCBL deletion mutants could not grow without supplementation of ether methionine (1mM) or homocysteine (0.5mg/ml). FgSAT deletion mutant showed fewer aerial mycelia, though the radial growth was not distinct from that of wild-type parent. Conidiation assays indicated that FgCGS, FgSAT and FgCBL deletion mutants produced as many conidia as wild-type parent, while conidiation in FgHAT deletion mutant was determined by the nutrient of media. However, compared with wild-type parent, conidial germination rate of FgCGS deletion mutant in2%sucrose solution reduced significantly, but was rescued by supplementation of methionine (1mM) or homocysteine (0.5mg/ml). Conidial germination ability of FgSAT deletion mutant reduced a little. Inoculation tests showed that FgCGS, FgHAT and FgCBL deletion mutants exhibited decreased virulence significantly on wheat heads, while the virulence of FgSAT deletion mutant was comparable to wild-type parent. FgCGS, FgSAT, FgHAT and FgCBL deletion mutants showed fewer aerial mycelia in wheat ear culture, and expression of Tri5gene was decreased significantly, compared with wild-type parent. Deoxynivalenol (DON) production of FgCGS deletion mutant was4-fold less than wild-type parent. Fungicide evaluation assays revealed that sensitivity of FgCGS, FgSAT, FgHAT and FgCBL deletion mutants to anilinopyrimidine fungicide pyrimethanil was not altered, indicating that FgCGS, FgSAT, FgHAT and FgCBL are not likely to be the target of pyrimethanil in F. graminearum. FgCGS and FgHAT deletion mutants showed increased sensitivity to sterol demethylation inhibitors (DMI), which was consistent with a low level of CYP51A/B/C expression in FgCGS deletion mutant. Sensitivity to osmotic, oxidative and cell wall (membrane) stresses revealed that FgCGS, FgSAT, FgHAT and FgCBL deletion mutants did not show observable changes in mycelia growth compared to wild-type parent, which indicate FgCGS, FgSAT, FgHAT and FgCBL are not involved in these stress responses. These results indicate that FgCGS, FgSAT, FgHAT and FgCBL play important roles in the growth, conidiation and pathogenicity of F. graminearum.
2) Mycelial growth tests showed FgMsrA, FgMsrB and FgfMsr deletion mutants had normal growth on PDA and FGA, and produced as many conidia in MBL medium as wild-type parent. Inoculation tests showed that the virulence of FgMsrA、FgMsrB and FgfMsr deletion mutants was comparable to wild-type parent. Fungicide sensitivity assays revealed that FgMsrA、FgMsrB and FgfMsr deletion mutants did not change the sensitivity to pyrimethanil, DMIs and carbendazim, indicating that FgMsrA、FgMsrB and FgfMsr may not be the target of pyrimethanil in F. graminearum. Oxidative stress sensitivity tests revealed that FgMsr A、FgMsr B and FgfMsr deletion mutants did not show observable changes in mycelia growth compared to wild-type parent. However, relative expression of FgMsr A、FgMsrB and FgfMs was increased significantly under oxidative stress. These results suggest that FgMsrA、FgMsrB and FgfMsr may be involed in oxidative stress reaction process of F. graminearum, though their roles in the growth, conidiation and pathogenicity are not obvious.
引文
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