The Thermomicrobia is a group of thermophilicgreen non-sulfur bacteria. Based on species Thermomicrobium roseum and Sphaerobacter thermophilus, this bacteriaclass has the following description: The class Thermomicrobia subdivides into two orders with validly published names: Thermomicrobiales Garrity and Holt 2001 and Sphaerobacterales Stackebrandt, Rainey and Ward-Rainey 1997. Gram negative. Pleomorphic, non-motile, non-spore-forming rods. Non-sporulating. No diamino acid present. No peptidoglycan in significant amount. Atypical proteinaceous cell walls. Hyper-thermophilic, optimum growth temperature at 70-75 °C. Obligatory aerobic and chemoorganotrophic. As thermophilic bacteria, members of this class are usually found in environments which are distant from human activity. However, they have features like improved growth in antibiotics and CO oxidizing activity, making them interesting topics of research.
History
In 1973, a strain of rose-pink thermophilic bacteria was isolated from Toadstool Spring in Yellowstone National Park, which was later named Thermomicrobium roseum and proposed as a novel species of the novel genus Thermomicrobium. At that time the genus was categorized under family Achromobacteraceae, but it became a distinct phylum by 2001. In 2004, it was proposed, on the basis of an analysis of genetic affiliations, that the Thermomicrobia should more properly be reclassified as a class belonging to the phylum Chloroflexi. The bacteria Sphaerobacter thermophilus originally described as an Actinobacteria is now considered a Thermomicrobia. In the same year, another strain of rose-pink thermophilic bacteria was isolated from Yellowstone National Park, which was named Thermobaculum terrenum. Later analysis based on genome put this species under Thermomicrobia class. However, the current standing of Thermobaculum terrenum is disputed. In 2012, a thermo-tolerant nitrite-oxidizing bacterium was isolated from a bioreactor, which was named Nitrolancetus hollandica and proposed as a novel species later in 2014. While it has nitrite-oxidizing activity, which is unique in the Thermomicrobia class, it is placed under the Thermomicrobia class based on 16s rRNA phylogeny. In 2014, two thermophilic, Gram-positive, rod-shaped, non-spore-forming bacteria isolated from geothermally heated biofilms growing on a tumulus in the Kilauea Iki pit crater on the flank of Kilauea Volcano were proposed as representatives of new species based on 16s rRNA phylogeny. The KI3T strain, later named as Thermomicrobium carboxidum, is closely related to Thermomicrobium roseum. The KI4T strain, later named as Thermorudis peleae, was proposed as a type strain of new genus Thermorudis. In 2015, a thermophilic bacteria strain WKT50.2 isolated from geothermal soil in Waitike was proposed to be a novel species, later named Thermorudis pharmacophila. Phylogenic analysis based on 16s rRNA place it within Thermomicrobia class, as close relative to Thermorudis peleae.
Characteristics
Living environment
Members of the class Thermomicrobia are broadly distributed across a wide range of both aquatic and terrestrial habitats. Thermomicrobium roseum was found in geothermally heated hot springs, Thermorudis pharmacophila and Thermobaculum terrenum from heated soils, and Thermomicrobium carboxidum and Thermorudis peleae from heated sediments In addition, Sphaerobacter thermophilus was found in sewage sludge that went through thermophilic treatment. The common features of their habitats include temperature ranging from around 65~75 °C and a pH around 6.0~8.0.
Metabolism
Members of Thermomicrobia class have variation in their basic metabolism. Nitrolancetus hollandica has nitrifying activity that utilize NO2− as energy source, which is unique in the whole Chloroflexi phylum. Thermomicrobium spp. and Sphaerobacter thermophilus have constitutive CO oxidizing not found in other species in this class. However, species of this class do share some features, as listed below:
All members except Thermobaculum terrenum have inability to utilize some common monosaccharides as sole carbon source. The mechanisms behind this inability are currently unknown.
Antibiotic resistance
Members of Thermomicrobia class exhibit certain level of resistance against metronidazole and/or trimethoprim, which are clinically relevant for humans. Thermomicrobium carboxidum and Thermorudis peleae show resistance against both of those antibiotics, while Sphaerobacter thermophilus shows resistance against only metronidazole. Interestingly, Thermomicrobium roseum and Thermorudis pharmacophila have an increased growth in both metronidazole and trimethoprim, a rare trait even within antibiotic resistant bacteria. The mechanisms behind are currently undocumented, and further study is required on this topic.
Members of Thermomicrobia class have various Gram-staining results. Thermomicrobium roseum, Sphaerobacter thermophilus Thermorudis and pharmacophila are reported to be Gram-negative and have a typical layered didermcell envelope structure. However, their cell envelope composition are atypical compared to typical Gram-negative bacteria. Cell envelop of Thermomicrobium roseum lacks significant amount of peptidoglycan, which is fundamental for typical Gram-negative bacteria, while being rich in protein. Membrane lipids of Thermomicrobium roseum are mostly long chain diols instead of glycerol-based lipids commonly found in bacteria. The same feature was found in Sphaerobacter thermophilus and Thermorudis pharmacophila. It was suggested that the high-protein and diol-based lipid composition are responsible for heat resistance of these bacteria. Meanwhile, other members of Thermomicrobia class are reported to be Gram-positive and have typical monodrem cell envelope. There are some possible explanations of the inconsistency of Gram-staining result within the class. For Thermorudis pharmacophila, a possible explanation suggested by Houghton et al. is that it is actually an atypical monoderm bacterium, because its cell envelope contains amino acids usually associated with Gram-positive bacteria, have reaction to KOH, vancomycin and ampicillin, and lacks genes responsible for diderm formation. It is also suggested that further study is required to resolve this problem, since the inconsistent reports of cell envelope structure are found for the whole Chloroflexi phylum.
Taxonomy
Thermomicrobia has the following taxonomy: Class Thermomicrobia
