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Applied and Environmental Microbiology, March 2005, p. 1473-1479, Vol. 71, No. 3
0099-2240/05/$08.00+0     doi:10.1128/AEM.71.3.1473-1479.2005
Copyright © 2005, American Society for Microbiology. All Rights Reserved.

"Endomicrobia": Cytoplasmic Symbionts of Termite Gut Protozoa Form a Separate Phylum of Prokaryotes

Ulrich Stingl,1 Renate Radek,2 Hong Yang,1,{dagger} and Andreas Brune1,3*

Fachbereich Biologie, Universität Konstanz, Konstanz,1 Institut für Biologie/Zoologie, Freie Universität Berlin, Berlin,2 Max-Planck-Institut für terrestrische Mikrobiologie, Marburg, Germany3

Received 31 August 2004/ Accepted 8 October 2004

Lignocellulose digestion by wood-feeding termites depends on the mutualistic interaction of unusual, flagellate protists located in their hindgut. Most of the flagellates harbor numerous prokaryotic endosymbionts of so-far-unknown identity and function. Using a full-cycle molecular approach, we show here that the endosymbionts of the larger gut flagellates of Reticulitermes santonensis belong to the so-called termite group 1 (TG-1) bacteria, a group of clones previously obtained exclusively from gut homogenates of Reticulitermes speratus that are only distantly related to other bacteria and are considered a novel bacterial phylum based on their 16S rRNA gene sequences. Fluorescence in situ hybridization with specifically designed oligonucleotide probes confirmed that TG-1 bacteria are indeed located within the flagellate cells and demonstrated that Trichonympha agilis (Hypermastigida) and Pyrsonympha vertens (Oxymonadida) harbor phylogenetically distinct populations of symbionts (<95% sequence similarity). Transmission electron microscopy revealed that the symbionts are small, spindle-shaped cells (0.6 µm in length and 0.3 µm in diameter) surrounded by two membranes and located within the cytoplasm of their hosts. The symbionts of the two flagellates are described as candidate species in the candidate genus "Endomicrobium." Moreover, we provide evidence that the members of the TG-1 phylum, for which we propose the candidate name "Endomicrobia," are phylogenetically extremely diverse and are present in and also restricted to the guts of all lower termites and wood-feeding cockroaches of the genus Cryptocercus, the only insects that are in an exclusive, obligately mutualistic association with such unique cellulose-fermenting protists.

* Corresponding author. Mailing address: Max-Planck-Institut für terrestrische Mikrobiologie, Karl-von-Frisch-Strasse, 35043 Marburg, Germany. Phone: 49-6421-178701. Fax: 49-6421-178709. E-mail: brune{at}staff.uni-marburg.de.

{dagger} Present address: College of Life Science, Central China Normal University, Wuhan 430079, People's Republic of China.

Applied and Environmental Microbiology, March 2005, p. 1473-1479, Vol. 71, No. 3
0099-2240/05/$08.00+0     doi:10.1128/AEM.71.3.1473-1479.2005
Copyright © 2005, American Society for Microbiology. All Rights Reserved.



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