Beresford P J, Kam C M, Powers J C, Lieberman J
Center for Blood Research, Harvard Medical School, Boston, MA 02115, USA.
Proc Natl Acad Sci U S A. 1997 Aug 19;94(17):9285-90. doi: 10.1073/pnas.94.17.9285.
The release of cytotoxic granule contents by cytotoxic T lymphocytes triggers apoptotic target cell death. Cytotoxic granules contain a pore-forming protein, perforin, and a group of serine proteases called granzymes. We expressed human granzyme A in bacteria as a proenzyme capable of in vitro activation by enterokinase. The recombinant activated enzyme has catalytic activity against substrates with Arg, preferably, or Lys at the P1 position, comparable to trypsin. An enzymatically inactive recombinant granzyme A, with the active site Ser mutated to Ala, was produced and used with affinity chromatography to identify potential substrates. Two granzyme A-binding cytoplasmic proteins of molecular mass 33 and 44 kDa were isolated and identified by tryptic fragment sequencing as PHAP I and II, ubiquitous putative HLA-associated proteins, previously coisolated by binding to an HLA class II peptide. PHAP II forms an SDS-stable complex with recombinant mutant granzyme A and coprecipitates with it from cytoplasmic extracts. PHAP II, either purified or in cell lysates, is cleaved by the recombinant enzyme at nanomolar concentrations to a 25-kDa fragment. PHAP II begins to be degraded within minutes of initiation of cytotoxic T lymphocyte attack. PHAP I and II are candidate participants in the granzyme A pathway of cell-mediated cytotoxicity.
细胞毒性T淋巴细胞释放细胞毒性颗粒成分会引发靶细胞凋亡死亡。细胞毒性颗粒含有一种成孔蛋白穿孔素和一组称为颗粒酶的丝氨酸蛋白酶。我们在细菌中表达人颗粒酶A,使其成为一种可被肠激酶体外激活的酶原。重组激活酶对P1位置含有精氨酸(最好是)或赖氨酸的底物具有催化活性,与胰蛋白酶相当。我们制备了一种酶活性无活性的重组颗粒酶A,其活性位点丝氨酸突变为丙氨酸,并用于亲和色谱法以鉴定潜在底物。通过胰蛋白酶片段测序分离并鉴定了两种分子量分别为33 kDa和44 kDa的与颗粒酶A结合的细胞质蛋白,分别为PHAP I和II,它们是普遍存在的假定HLA相关蛋白,之前通过与HLA II类肽结合而共同分离出来。PHAP II与重组突变颗粒酶A形成SDS稳定复合物,并从细胞质提取物中与其共沉淀。纯化的或细胞裂解物中的PHAP II在纳摩尔浓度下被重组酶切割成25 kDa片段。在细胞毒性T淋巴细胞攻击开始后的几分钟内,PHAP II开始降解。PHAP I和II是细胞介导的细胞毒性颗粒酶A途径中的候选参与者。