Gabaldón Carlos, Gómez-Ros Laura V, Núñez-Flores María J López, Esteban-Carrasco Alberto, Barceló Alfonso Ros
Department of Plant Biology, University of Murcia, 30100, Murcia, Spain.
Plant Mol Biol. 2007 Sep;65(1-2):43-61. doi: 10.1007/s11103-007-9197-0. Epub 2007 Jun 22.
The major basic peroxidase (ZePrx) from Zinnia elegans suspension cell cultures was purified and cloned. The purification resolved ZePrxs in two isoforms (ZePrx33.44 and ZePrx34.70), whose co-translational and post-translational modifications are characterized. Based on the N-terminal sequence obtained by Edman degradation of mature ZePxs, it may be expected that the immature polypeptides of ZePrxs contain a signal peptide (N-terminal pro-peptide) of 30 amino acids, which directs the polypeptide chains to the ER membrane. These immature polypeptides are co-translationally processed by proteolytic cleavage, and modeling studies of digestions suggested that the processing of the N-terminal pro-peptide of ZePrxs is performed by a peptidase from the SB clan (S8 family, subfamily A) of serine-type proteases. When the post-translational modifications of ZePrxs were characterized by trypsin digestion, and tryptic peptides were analyzed by reverse phase nano liquid chromatography (RP-nanoLC) coupled to MALDI-TOF MS, it was seen that, despite the presence in the primary structure of the protein of several (disulphide bridges, N-glycosylation, phosphorylation and N-myristoylation) potential post-translational modification sites, ZePrxs are only post-translationated modified by the formation of N-terminal pyroglutamate residues, disulphide bridges and N-glycosylation. Glycans of ZePrxs belong to three main types and conduce to the existence of at least ten different molecular isoforms. The first glycans belong to both low and high mannose-type glycans, with the growing structure Man(3-9)(GlcNAc)(2). Low mannose-type glycans, Man(3-4)(GlcNAc)(2), coexist with the truncated (paucimannosidic-type) glycan, Man(3)Xyl(1)Fuc(1)(GlcNAc)(2), in the G(3) and G(4 )sub-isoforms of ZePrx33.44. In ZePrx34.70, on the other hand, the complex-type biantennary glycan, Man(3)Xyl(1)Fuc(3)(GlcNAc)(5), and the truncated (paucimannosidic-type) glycan, Man(3)Xyl(1)Fuc(1)(GlcNAc)(2), appear to fill the two putative sites for N-glycosylation. Since the two N-glycosylation sites in ZePrxs are located in an immediately upstream loop region of helix F'' (close to the proximal histidine) and in helix F'' itself, and are flanked by positive-charged amino acids that produce an unusual positive-net surface electrostatic charge pattern, it may be expected that glycans not only affect reaction dynamics but may well participate in protein/cell wall interactions. These results emphasize the complexity of the ZePrx proteome and the difficulties involved in establishing any fine structure-function relationship.
从百日草悬浮细胞培养物中纯化并克隆了主要的碱性过氧化物酶(ZePrx)。纯化过程中分离出两种ZePrx同工型(ZePrx33.44和ZePrx34.70),并对其共翻译和翻译后修饰进行了表征。根据通过对成熟ZePxs进行埃德曼降解获得的N端序列,可以预期ZePrxs的未成熟多肽包含一个30个氨基酸的信号肽(N端前肽),该信号肽将多肽链导向内质网膜。这些未成熟多肽通过蛋白水解切割进行共翻译加工,消化的建模研究表明,ZePrxs的N端前肽的加工是由丝氨酸型蛋白酶SB家族(S8家族,亚家族A)的一种肽酶进行的。当通过胰蛋白酶消化对ZePrxs的翻译后修饰进行表征,并通过与MALDI-TOF MS联用的反相纳米液相色谱(RP-nanoLC)分析胰蛋白酶肽段时,发现尽管该蛋白质的一级结构中存在几个(二硫键、N-糖基化、磷酸化和N-肉豆蔻酰化)潜在的翻译后修饰位点,但ZePrxs仅通过形成N端焦谷氨酸残基、二硫键和N-糖基化进行翻译后修饰。ZePrxs的聚糖属于三种主要类型,导致至少十种不同分子同工型的存在。第一种聚糖属于低甘露糖型和高甘露糖型聚糖,其生长结构为Man(3-9)(GlcNAc)(2)。低甘露糖型聚糖Man(3-4)(GlcNAc)(2)与截短的(寡甘露糖型)聚糖Man(3)Xyl(1)Fuc(1)(GlcNAc)(2)共存于ZePrx33.44的G(3)和G(4)亚同工型中。另一方面,在ZePrx34.70中,复合型双天线聚糖Man(3)Xyl(1)Fuc(3)(GlcNAc)(5)和截短的(寡甘露糖型)聚糖Man(3)Xyl(1)Fuc(1)(GlcNAc)(2)似乎占据了两个假定的N-糖基化位点。由于ZePrxs中的两个N-糖基化位点位于螺旋F''的紧邻上游环区域(靠近近端组氨酸)以及螺旋F''本身,并且两侧是带正电荷的氨基酸,产生了不寻常的正净表面静电电荷模式,因此可以预期聚糖不仅会影响反应动力学,还很可能参与蛋白质/细胞壁相互作用。这些结果强调了ZePrx蛋白质组的复杂性以及建立任何精细的结构-功能关系所涉及的困难。