Zhang H H, Blanco D R, Exner M M, Shang E S, Champion C I, Phillips M L, Miller J N, Lovett M A
Department of Microbiology, Immunology, and Molecular Genetics, School of Medicine, University of California at Los Angeles, Los Angeles, California 90095, USA.
J Bacteriol. 1999 Dec;181(23):7168-75. doi: 10.1128/JB.181.23.7168-7175.1999.
We have previously observed that while native Treponema pallidum rare outer membrane protein 1 (Tromp1) is hydrophobic and has porin activity, recombinant forms of Tromp1 do not possess these properties. In this study we show that these properties are determined by conformation and can be replicated by proper renaturation of recombinant Tromp1. Native Tromp1, but not the 47-kDa lipoprotein, extracted from whole organisms by using Triton X-114, was found to lose hydrophobicity after treatment in 8 M urea, indicating that Tromp1's hydrophobicity is conformation dependent. Native Tromp1 was purified from 0.1% Triton X-100 extracts of whole organisms by fast-performance liquid chromatography (FPLC) and shown to have porin activity in planar lipid bilayers. Cross-linking studies of purified native Tromp1 with an 11 A cross-linking agent showed oligomeric forms consistent with dimers and trimers. For renaturation studies of recombinant Tromp1 (rTromp1), a 31,109-Da signal-less construct was expressed in Escherichia coli and purified by FPLC. FPLC-purified rTromp1 was denatured in 8 M urea and then renatured in the presence of 0.5% Zwittergent 3,14 during dialysis to remove the urea. Renatured rTromp1 was passed through a Sephacryl S-300 gel exclusion column previously calibrated with known molecular weight standards. While all nonrenatured rTromp1 eluted from the column at approximately the position of the carbonic anhydrase protein standard (29 kDa), all renatured rTromp1 eluted at the position of the phosphorylase b protein standard (97 kDa), suggesting a trimeric conformation. Trimerization was confirmed by using an 11 A cross-linking agent which showed both dimers and trimers similar to that of native Tromp1. Triton X-114 phase separations showed that all of renatured rTromp1, but none of nonrenatured rTromp1, phase separated exclusively into the hydrophobic detergent phase, similar to native Tromp1. Circular dichroism of nonrenatured and renatured rTromp1 showed a marked loss in alpha-helical secondary structure of renatured rTromp1 compared to the nonrenatured form. Finally, renatured rTromp1, but not the nonrenatured form, showed porin activity in planar liquid bilayers. These results demonstrate that proper folding of rTromp1 results in a trimeric, hydrophobic, and porin-active conformation similar to that of the native protein.
我们之前观察到,尽管梅毒螺旋体天然稀有外膜蛋白1(Tromp1)具有疏水性且有孔蛋白活性,但重组形式的Tromp1并不具备这些特性。在本研究中,我们表明这些特性由构象决定,并且可以通过重组Tromp1的适当复性来重现。通过使用Triton X - 114从全菌体中提取的天然Tromp1,而非47 kDa脂蛋白,在8 M尿素中处理后被发现失去疏水性,这表明Tromp1的疏水性依赖于构象。天然Tromp1通过快速液相色谱(FPLC)从全菌体的0.1% Triton X - 100提取物中纯化得到,并在平面脂双层中显示出孔蛋白活性。用一种11 Å交联剂对纯化的天然Tromp1进行交联研究,结果显示其寡聚形式与二聚体和三聚体一致。对于重组Tromp1(rTromp1)的复性研究,一个31,109 Da无信号构建体在大肠杆菌中表达并通过FPLC纯化。FPLC纯化的rTromp1在8 M尿素中变性,然后在透析过程中于0.5%两性离子去污剂3,14存在的情况下复性以去除尿素。复性的rTromp1通过先前用已知分子量标准物校准过的Sephacryl S - 300凝胶排阻柱。虽然所有未复性的rTromp1在柱上的洗脱位置大约在碳酸酐酶蛋白标准物(29 kDa)处,但所有复性的rTromp1在磷酸化酶b蛋白标准物(97 kDa)的位置洗脱,这表明其为三聚体构象。使用一种11 Å交联剂证实了三聚化,该交联剂显示出与天然Tromp1类似的二聚体和三聚体。Triton X - 114相分离表明,所有复性的rTromp1,但没有未复性的rTromp1,专门相分离到疏水去污剂相中,这与天然Tromp1相似。未复性和复性的rTromp1的圆二色性显示,与未复性形式相比,复性的rTromp1的α - 螺旋二级结构明显丧失。最后,复性的rTromp1,但不是未复性形式,在平面液体脂双层中显示出孔蛋白活性。这些结果表明,rTromp1的正确折叠导致了一种与天然蛋白相似的三聚体、疏水且具有孔蛋白活性的构象。