Studer Alex, McAnulla Craig, Büchele Rainer, Leisinger Thomas, Vuilleumier Stéphane
Institut für Mikrobiologie, ETH Zürich, CH-8092 Zürich, Switzerland.
J Bacteriol. 2002 Jul;184(13):3476-84. doi: 10.1128/JB.184.13.3476-3484.2002.
Methylobacterium chloromethanicum CM4 is an aerobic alpha-proteobacterium capable of growth with chloromethane as the sole carbon and energy source. Two proteins, CmuA and CmuB, were previously purified and shown to catalyze the dehalogenation of chloromethane and the vitamin B12-mediated transfer of the methyl group of chloromethane to tetrahydrofolate. Three genes located near cmuA and cmuB, designated metF, folD and purU and encoding homologs of methylene tetrahydrofolate (methylene-H4folate) reductase, methylene-H4folate dehydrogenase-methenyl-H4folate cyclohydrolase and formyl-H4folate hydrolase, respectively, suggested the existence of a chloromethane-specific oxidation pathway from methyl-tetrahydrofolate to formate in strain CM4. Hybridization and PCR analysis indicated that these genes were absent in Methylobacterium extorquens AM1, which is unable to grow with chloromethane. Studies with transcriptional xylE fusions demonstrated the chloromethane-dependent expression of these genes. Transcriptional start sites were mapped by primer extension and allowed to define three transcriptional units, each likely comprising several genes, that were specifically expressed during growth of strain CM4 with chloromethane. The DNA sequences of the deduced promoters display a high degree of sequence conservation but differ from the Methylobacterium promoters described thus far. As shown previously for purU, inactivation of the metF gene resulted in a CM4 mutant unable to grow with chloromethane. Methylene-H4folate reductase activity was detected in a cell extract of strain CM4 only in the presence of chloromethane but not in the metF mutant. Taken together, these data provide evidence that M. chloromethanicum CM4 requires a specific set of tetrahydrofolate-dependent enzymes for growth with chloromethane.
氯甲烷甲基杆菌CM4是一种好氧α-变形菌,能够以氯甲烷作为唯一碳源和能源进行生长。之前已纯化出两种蛋白质,即CmuA和CmuB,它们可催化氯甲烷的脱卤反应以及维生素B12介导的氯甲烷甲基向四氢叶酸的转移。位于cmuA和cmuB附近的三个基因,分别命名为metF、folD和purU,它们编码亚甲基四氢叶酸还原酶、亚甲基-四氢叶酸脱氢酶-亚甲基-四氢叶酸环水解酶和甲酰-四氢叶酸水解酶的同源物,这表明在CM4菌株中存在一条从甲基-四氢叶酸到甲酸的氯甲烷特异性氧化途径。杂交和PCR分析表明,在无法利用氯甲烷生长的扭脱甲基杆菌AM1中不存在这些基因。利用转录型xylE融合蛋白进行的研究表明,这些基因的表达依赖于氯甲烷。通过引物延伸确定了转录起始位点,并确定了三个转录单元,每个转录单元可能包含几个基因,它们在CM4菌株利用氯甲烷生长期间特异性表达。推导的启动子的DNA序列显示出高度的序列保守性,但与迄今为止描述的甲基杆菌启动子不同。如之前对purU的研究所示,metF基因失活导致CM4突变体无法利用氯甲烷生长。仅在存在氯甲烷的情况下,在CM4菌株的细胞提取物中检测到亚甲基-四氢叶酸还原酶活性,而在metF突变体中未检测到。综上所述,这些数据提供了证据,表明氯甲烷甲基杆菌CM4利用氯甲烷生长需要一组特定的依赖四氢叶酸的酶。