Cavalier-Smith T
Department of Botany, University of British Columbia, Vancouver, Canada.
Microbiol Rev. 1993 Dec;57(4):953-94. doi: 10.1128/mr.57.4.953-994.1993.
The demarcation of protist kingdoms is reviewed, a complete revised classification down to the level of subclass is provided for the kingdoms Protozoa, Archezoa, and Chromista, and the phylogenetic basis of the revised classification is outlined. Removal of Archezoa because of their ancestral absence of mitochondria, peroxisomes, and Golgi dictyosomes makes the kingdom Protozoa much more homogeneous: they all either have mitochondria and peroxisomes or have secondarily lost them. Predominantly phagotrophic, Protozoa are distinguished from the mainly photosynthetic kingdom Chromista (Chlorarachniophyta, Cryptista, Heterokonta, and Haptophyta) by the absence of epiciliary retronemes (rigid thrust-reversing tubular ciliary hairs) and by the lack of two additional membranes outside their chloroplast envelopes. The kingdom Protozoa has two subkingdoms: Adictyozoa, without Golgi dictyosomes, containing only the phylum Percolozoa (flagellates and amoeboflagellates); and Dictyozoa, made up of 17 phyla with Golgi dictyosomes. Dictyozoa are divided into two branches: (i) Parabasalia, a single phylum with hydrogenosomes and 70S ribosomes but no mitochondria, Golgi dictyosomes associated with striated roots, and a kinetid of four or five cilia; and (ii) Bikonta (16 unicellular or plasmodial phyla with mitochondria and bikinetids and in which Golgi dictyosomes are not associated with striated ciliary roots), which are divided into two infrakingdoms: Euglenozoa (flagellates with discoid mitochondrial cristae and trans-splicing of miniexons for all nuclear genes) and Neozoa (15 phyla of more advanced protozoa with tubular or flat [usually nondiscoid] mitochondrial cristae and cis-spliced spliceosomal introns). Neozoa are divided into seven parvkingdoms: (i) Ciliomyxa (three predominantly ciliated phyla with tubular mitochondrial cristae but no cortical alveoli, i.e., Opalozoa [flagellates with tubular cristae], Mycetozoa [slime molds], and Choanozoa [choanoflagellates, with flattened cristae]); (ii) Alveolata (three phyla with cortical alveoli and tubular mitochondrial cristae, i.e., Dinozoa [Dinoflagellata and Protalveolata], Apicomplexa, and Ciliophora); (iii) Neosarcodina (phyla Rhizopoda [lobose and filose amoebae] and Reticulosa [foraminifera; reticulopodial amoebae], usually with tubular cristae); (iv) Actinopoda (two phyla with axopodia: Heliozoa and Radiozoa [Radiolaria, Acantharia]); (v) Entamoebia (a single phylum of amoebae with no mitochondria, peroxisomes, hydrogenosomes, or cilia and with transient intranuclear centrosomes); (vi) Myxozoa (three endoparasitic phyla with multicellular spores, mitochondria, and no cilia: Myxosporidia, Haplosporidia, and Paramyxia); and (vii) Mesozoa (multicells with tubular mitochondrial cristae, included in Protozoa because, unlike animals, they lack collagenous connective tissue).
回顾了原生生物界的划分,给出了原生动物界、古原生生物界和色素界完整的修订分类,直至亚纲级别,并概述了修订分类的系统发育基础。由于古原生生物界的祖先缺乏线粒体、过氧化物酶体和高尔基体,将其移除后使原生动物界更加同质化:它们要么都有线粒体和过氧化物酶体,要么后来失去了这些细胞器。原生动物主要是吞噬营养型,与主要进行光合作用的色素界(绿藻虫纲、隐藻纲、不等鞭毛类和定鞭藻纲)不同,原生动物没有纤毛后向丝(刚性的推力反转管状纤毛),并且在叶绿体包膜外没有另外两层膜。原生动物界有两个亚界:无高尔基体亚界,没有高尔基体,仅包含渗养门(鞭毛虫和变形鞭毛虫);有高尔基体亚界,由17个有高尔基体的门组成。有高尔基体亚界分为两个分支:(i)副基体亚门,一个单一的门,有氢化酶体和70S核糖体,但没有线粒体,高尔基体与横纹根相关联,有一个由四或五根纤毛组成的动质体;(ii)双鞭毛亚界(16个单细胞或多核体门,有线粒体和双动质体,其中高尔基体不与横纹纤毛根相关联),分为两个下纲:眼虫纲(鞭毛虫,线粒体嵴呈盘状,所有核基因的微小外显子进行反式剪接)和新原生动物纲(15个更高级的原生动物门,线粒体嵴呈管状或扁平状[通常不是盘状],剪接体内含子进行顺式剪接)。新原生动物纲分为七个小界:(i)纤毛粘菌界(三个主要有纤毛的门,线粒体嵴呈管状,但没有皮层泡,即扁虫藻纲[鞭毛虫,线粒体嵴呈管状]、黏菌门[黏菌]和领鞭毛虫纲[领鞭毛虫,嵴扁平]);(ii)囊泡虫界(三个有皮层泡和管状线粒体嵴的门,即恐龙纲[甲藻纲和原始囊泡虫纲]、顶复门和纤毛虫门);(iii)新肉足虫界(根足纲[叶状和丝状变形虫]和网足虫纲[有孔虫;网状伪足变形虫],通常线粒体嵴呈管状);(iv)辐足虫界(两个有轴伪足的门:太阳虫纲和放射虫纲[放射虫、棘胞亚门]);(v)内阿米巴界(一个变形虫门,没有线粒体、过氧化物酶体、氢化酶体或纤毛,有短暂的核内中心体);(vi)粘孢子虫界(三个内寄生门,有多细胞孢子、线粒体,没有纤毛:粘孢子虫纲、单孢子虫纲和副粘体纲);(vii)中生动物界(多细胞,线粒体嵴呈管状,归入原生动物界,因为与动物不同,它们缺乏胶原结缔组织)。