Mi-ichi Fumika, Miyamoto Tomofumi, Takao Shouko, Jeelani Ghulam, Hashimoto Tetsuo, Hara Hiromitsu, Nozaki Tomoyoshi, Yoshida Hiroki
Division of Molecular and Cellular Immunoscience, Department of Biomolecular Sciences, Faculty of Medicine, Saga University, Saga, Saga 849-8501, Japan;
Department of Natural Products Chemistry, Graduate School of Pharmaceutical Sciences, Kyushu University, Higashi-ku, Fukuoka 812-8582, Japan;
Proc Natl Acad Sci U S A. 2015 Jun 2;112(22):E2884-90. doi: 10.1073/pnas.1423718112. Epub 2015 May 18.
Hydrogenosomes and mitosomes are mitochondrion-related organelles (MROs) that have highly reduced and divergent functions in anaerobic/microaerophilic eukaryotes. Entamoeba histolytica, a microaerophilic, parasitic amoebozoan species, which causes intestinal and extraintestinal amoebiasis in humans, possesses mitosomes, the existence and biological functions of which have been a longstanding enigma in the evolution of mitochondria. We previously demonstrated that sulfate activation, which is not generally compartmentalized to mitochondria, is a major function of E. histolytica mitosomes. However, because the final metabolites of sulfate activation remain unknown, the overall scheme of this metabolism and the role of mitosomes in Entamoeba have not been elucidated. In this study we purified and identified cholesteryl sulfate (CS) as a final metabolite of sulfate activation. We then identified the gene encoding the cholesteryl sulfotransferase responsible for synthesizing CS. Addition of CS to culture media increased the number of cysts, the dormant form that differentiates from proliferative trophozoites. Conversely, chlorate, a selective inhibitor of the first enzyme in the sulfate-activation pathway, inhibited cyst formation in a dose-dependent manner. These results indicate that CS plays an important role in differentiation, an essential process for the transmission of Entamoeba between hosts. Furthermore, we show that Mastigamoeba balamuthi, an anaerobic, free-living amoebozoan species, which is a close relative of E. histolytica, also has the sulfate-activation pathway in MROs but does not possess the capacity for CS production. Hence, we propose that a unique function of MROs in Entamoeba contributes to its adaptation to its parasitic life cycle.
氢化酶体和线粒体相关细胞器(MROs)是在厌氧/微需氧真核生物中功能高度退化且具有差异的细胞器。溶组织内阿米巴是一种微需氧的寄生变形虫物种,可导致人类肠道和肠外阿米巴病,它拥有线粒体相关细胞器(mitosomes),其存在和生物学功能在线粒体进化过程中一直是个长期未解之谜。我们之前证明,通常不局限于线粒体的硫酸盐激活是溶组织内阿米巴线粒体相关细胞器的主要功能。然而,由于硫酸盐激活的最终代谢产物尚不清楚,这种代谢的整体方案以及线粒体相关细胞器在溶组织内阿米巴中所起的作用尚未阐明。在本研究中,我们纯化并鉴定出硫酸胆固醇(CS)是硫酸盐激活的最终代谢产物。然后我们鉴定出了负责合成CS的胆固醇硫酸转移酶的编码基因。向培养基中添加CS会增加囊肿的数量,囊肿是从增殖性滋养体分化而来的休眠形式。相反,氯酸盐作为硫酸盐激活途径中第一种酶的选择性抑制剂,以剂量依赖的方式抑制囊肿形成。这些结果表明,CS在分化过程中发挥重要作用,而分化是溶组织内阿米巴在宿主之间传播的一个关键过程。此外,我们表明,巴氏鞭毛虫是一种厌氧的、自由生活的变形虫物种,是溶组织内阿米巴的近亲,其线粒体相关细胞器中也有硫酸盐激活途径,但不具备产生CS的能力。因此,我们认为线粒体相关细胞器在溶组织内阿米巴中的独特功能有助于其适应寄生生活周期。