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Phylogenetic Foundation of Spirochetes

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Phylogenetic Foundation of Spirochetes J. Mol. Microbiol. Biotechnol. (2000) 2(4): 341-344. JMMBSpirochete Symposium Phylogeny on341 Spirochete Physiology Phylogenetic Foundation of Spirochetes Bruce J. Paster* and Floyd E. Dewhirst Spirochaetales that is divided into three families; namely the Spirochaetaceae, the Brachyspiraceae, and the Department of Molecular Genetics, The Forsyth Institute, Leptospiraceae. The phylogenetic relationships of 140 Fenway, Boston, Massachusetts 02115, USA representatives of each genus are shown in Figure 1. The Spirochaetaceae are separated into 6 genera— Borrelia, Brevinema, Cristispira, Spirochaeta, “Spironema”, Abstract and Treponema. New genera of termite spirochetes, such as Clevelandina, Diplocalyx, and Hollandina, have been The spirochetes are free-living or host-associated, described on the basis of differences in ultrastructural traits helical bacteria, some of which are pathogenic to man (Breznak, 1984). It has been suggested that they also and animal. Comparisons of 16S rRNA sequences belong in the family Spirochaetaceae, but no sequence demonstrate that the spirochetes represent a information is presently available to determine their monophyletic phylum within the bacteria. The phylogenetic position within the spirochetes. spirochetes are presently classified in the Class The Brachyspiraceae contain the genus Brachyspira Spirochaetes in the order Spirochetales and are (Serpulina). Due to the close phylogenetic relationship of divided into three major phylogenetic groupings, or B. aarlborgi to species characterized as Serpulina, it has families. The first family Spirochaetaceae contains been recommended that a single genus be justified. Thus, species of the genera Borrelia, Brevinema, Cristispira, Brachyspira takes precedence over Serpulina since the Spirochaeta, Spironema, and Treponema. The second former genus was listed first as a valid name (Hovind- family Brachyspiraceae contains the genus Hougen et al., 1983). Brachyspira (Serpulina). The third family The Leptospiraceae are separated into 2 known Leptospiraceae contains species of the genera genera, Leptonema and Leptospira. The putative Leptonema and Leptospira. Novel spirochetal species, misclassified “Leptospira” parvula may represent a third or phylotypes, that can not be presently cultivated in genus of the Leptospiraceae. vitro, have been identified from the human oral cavity, Despite the relatively deep branching of the the termite gut, and other host-associated or free-living spirochetes, they are also defined by single-base signature sources. There are now over 200 spirochetal species analysis (Paster et al., 1991). Thus, bases are found in the or phylotypes, of which more than half is presently 16S rRNA sequences of over 200 species or phylotypes not cultivable. It is likely that there is still a significant of spirochetes, but are not found (or rarely found) in unrecognized spirochetal diversity that should be sequences of approximately 16,000 other bacteria in evaluated. GenBank and the Ribosomal Database Project (Maidak et al., 1999). These data support the conclusions that Introduction spirochetes represent a single monophyletic phylum (Paster et al., 1991) and that they have descended from a Based on comparative analysis of 16S rRNA sequences, common spirochetal ancestor. the spirochetes represent one of about 40 major bacterial Sequences of species of the genera Treponema, phyla (Hugenholtz et al., 1998). The spirochetes are Spirochaeta, and Leptospira are unusual in that they unusual in that they are one of the few phyla whose possess a 20 to 30 base 5' extension of the 16S rRNA phenotypic characteristics reflect its natural phylogenetic molecule (Paster et al., 1991). The region is highly variable relationships, e.g., they possess a cellular ultrastructure and likely to form helices from 2 to 12 bp. The function of that is unique among the bacteria (Paster et al., 1991). this 5' extension is unknown. These 5' regions were The biodiversity of spirochetes is marked inasmuch as they discovered using the older technique of direct RNA are often observed in high numbers in the gastrointestinal sequencing with reverse transcriptase (Paster et al., 1991). tract of man, mammals, insects, and bivalves, as well as in aquatic and marine environments. Some species are Unifying Phenotypic Traits pathogenic to man, other mammals and birds. However, the focus of this paper is to review the phylogenetic All spirochetes are helical or spiral shaped microorganisms relationships of the spirochetes, including those that are with internal organelles of motility called periplasmic flagella presently not cultivable. (Paster and Canale-Parola, 1980). The flagella are inserted subterminally at each end of the cell, wrap around the Phylogenetic Foundation of Spirochetes protoplasmic cylinder, and usually overlap in the center region (Figure 2). Periplasmic flagella are located between Based on 16S rRNA or 16S rDNA sequence comparisons, the protoplasmic cylinder and an outer membrane-like spirochetes form a coherent phylogenetic phylum, as structure called the outer sheath. The number of shown in Figure 1 (Paster et al., 1991). The spirochetes periplasmic flagella can range from 2 to 100’s per cell are presently classified in the Class Spirochetes in the order depending upon the species. In some species under certain growth conditions, the periplasmic flagella can protrude *For correspondence. Email [email protected]; Tel. (617) 262-5200 x288; into the external environment (Charon et al., 1992). Fax. (617) 262-4021. Other unifying characteristics of the spirochetes include © 2000 Horizon Scientific Press Further Reading Caister Academic Press is a leading academic publisher of advanced texts in microbiology, molecular biology and medical research. 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The sequences of the species shown may be obtained through GenBank or by request from the authors. Designations for source of termite spirochetal clones and isolates: RFS and Zas, Reticulitermes
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