Mathur M, Kolattukudy P E
Ohio State Biotechnology Center, Ohio State University, Columbus 43210.
J Biol Chem. 1992 Sep 25;267(27):19388-95.
Mycocerosyl lipids are found uniquely in the cell walls of pathogenic mycobacteria. Mycocerosic acid synthase (MAS) is a multifunctional protein which catalyzes elongation of n-fatty acyl-CoA with methylamalonyl-CoA as the elongating agent (Rainwater, D. L., and Kolattukudy, P. E. (1985) J. Biol. Chem. 260, 616-623). To understand how the various domains that catalyze the reactions involved in chain elongation are organized, mas gene from Mycobacterium tuberculosis bovis BCG was cloned. A lambda gt11 library of AluI partially digested genomic DNA from the organism was screened with an oligonucleotide probe designed from the N-terminal amino acid sequence of purified MAS. Using terminal segments of inserts from positive clones as the probe, the library was rescreened and the process was repeated. Sequencing of four overlapping clones revealed a contiguous sequence of 9699 base pair(s) (bp) of mycobacterial genome containing a 6330-bp open reading frame that could code for a protein of 2100 amino acids with a molecular mass of 225,437 daltons. The authenticity of the open reading frame as that of MAS was verified by correspondence of the amino acid sequences deduced from the gene with the directly determined amino acid sequences of the N terminus and three different internal peptide fragments. By comparing the MAS amino acid sequence with the sequences in the active site regions of known fatty acid synthases and polyketide synthases the functional domains in MAS were identified. This analysis showed that the domains were organized in the following order: beta-ketoacyl synthase, acyl transferase, dehydratase-enoyl reductase, beta-ketoreductase, acyl carrier protein; no thioesterase-like domain could be found. These results establish MAS as the first case of an elongating multifunctional enzyme composed of two identical subunits that resemble the vertebrate fatty acid synthase in size, subunit structure, and linear organization of functional domains. Southern and Western blot analyses showed absence of mas gene and encoded proteins in Mycobacterium smegmatis and Escherichia coli. This result is consistent with the report that mycocerosic acid is present only in pathogenic mycobacteria.
分枝菌酸脂仅存在于致病性分枝杆菌的细胞壁中。分枝菌酸合酶(MAS)是一种多功能蛋白,它以甲基丙二酰辅酶A作为延伸剂催化n-脂肪酰辅酶A的延伸反应(Rainwater, D. L., and Kolattukudy, P. E. (1985) J. Biol. Chem. 260, 616 - 623)。为了了解催化链延伸反应的各个结构域是如何组织的,克隆了来自牛分枝杆菌卡介苗的mas基因。用根据纯化的MAS的N端氨基酸序列设计的寡核苷酸探针筛选了该生物体经AluI部分消化的基因组DNA的λgt11文库。以阳性克隆插入片段的末端片段为探针,对文库进行再次筛选,并重复该过程。对四个重叠克隆进行测序,揭示了分枝杆菌基因组中一段9699个碱基对(bp)的连续序列,其中包含一个6330 bp的开放阅读框,该开放阅读框可编码一个2100个氨基酸的蛋白质,分子量为225,437道尔顿。通过将从该基因推导的氨基酸序列与直接测定的N端和三个不同内部肽段的氨基酸序列进行比对,验证了该开放阅读框作为MAS的真实性。通过将MAS氨基酸序列与已知脂肪酸合酶和聚酮合酶活性位点区域的序列进行比较,确定了MAS中的功能结构域。该分析表明这些结构域按以下顺序排列:β-酮酰基合酶、酰基转移酶、脱水酶-烯酰还原酶、β-酮还原酶、酰基载体蛋白;未发现类似硫酯酶的结构域。这些结果表明MAS是由两个相同亚基组成的延伸多功能酶的首个实例,其大小、亚基结构和功能结构域的线性组织类似于脊椎动物脂肪酸合酶。Southern和Western印迹分析表明耻垢分枝杆菌和大肠杆菌中不存在mas基因及其编码的蛋白质。这一结果与分枝菌酸仅存在于致病性分枝杆菌中的报道一致。