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Mycobacteria

 

Mycobacteria

Mycobacterium
TEM micrograph of M. tuberculosis.
Scientific classification
Kingdom: Bacteria
Phylum: Actinobacteria
Order: Actinomycetales
Suborder: Corynebacterineae
Family: Mycobacteriaceae
Genus: Mycobacterium
Lehmann & Neumann 1896
Species

See below.

Mycobacterium is a genus of Actinobacteria, given its own family, the Mycobacteriaceae. The genus includes pathogens known to cause serious diseases in mammals, including tuberculosis (Mycobacterium tuberculosis) and leprosy (Mycobacterium leprae).[1] The Greek prefix myco means fungus, alluding to the way mycobacteria have been observed to grow in a mold-like fashion on the surface of liquids when cultured.[2]

Microbiologic characteristics


Mycobacteria are aerobic and nonmotile bacteria (except for the species Mycobacterium marinum, which has been shown to be motile within macrophages) that are characteristically acid-alcohol-fast.[1] Mycobacteria do not contain endospores or capsules and are usually considered Gram-positive. While mycobacteria do not seem to fit the Gram-positive category from an empirical standpoint (i.e., in general, they do not retain the crystal violet stain well), they are classified as an acid-fast Gram-positive bacterium due to their lack of an outer cell membrane. Mycobacterium marinum and perhaps M. bovis have been shown to sporulate;[3] however, this has been contested by further research.[4] All Mycobacterium species share a characteristic cell wall, thicker than in many other bacteria, which is hydrophobic, waxy, and rich in mycolic acids/mycolates. The cell wall consists of the hydrophobic mycolate layer and a peptidoglycan layer held together by a polysaccharide, arabinogalactan. The cell wall makes a substantial contribution to the hardiness of this genus. The biosynthetic pathways of cell wall components are potential targets for new drugs for tuberculosis.[5]

Many Mycobacterium species adapt readily to growth on very simple substrates, using ammonia or amino acids as nitrogen sources and glycerol as a carbon source in the presence of mineral salts. Optimum growth temperatures vary widely according to the species and range from 25 °C to over 50 °C.

Some species can be very difficult to culture (i.e. they are fastidious), sometimes taking over two years to develop in culture. Further, some species also have extremely long reproductive cycles — M. leprae, may take more than 20 days to proceed through one division cycle (for comparison, some E. coli strains take only 20 minutes), making laboratory culture a slow process.[1] In addition, the availability of genetic manipulation techniques still lags far behind that of other bacterial species.[6]

A natural division occurs between slowly– and rapidly–growing species. Mycobacteria that form colonies clearly visible to the naked eye within seven days on subculture are termed rapid growers, while those requiring longer periods are termed slow growers. Mycobacteria cells are straight or slightly curved rods between 0.2 and 0.6 µm wide by 1.0 and 10 µm long.

Pigmentation

Some mycobacteria produce carotenoid pigments without light. Others require photoactivation for pigment production.

Photochromogens (Group I)
Produce nonpigmented colonies when grown in the dark and pigmented colonies only after exposure to light and reincubation.
  • Ex: M. kansasii, M. marinum, M. simiae.
Scotochromogens (Group II)
Produce deep yellow to orange colonies when grown in the presence of either the light or the dark.
  • Ex: M. scrofulaceum, M. gordonae, M. xenopi, M. szulgai.
Non-chromogens (Groups III & IV)
Nonpigmented in the light and dark or have only a pale yellow, buff or tan pigment that does not intensify after light exposure.
  • Ex: M. tuberculosis, M. avium-intra-cellulare, M. bovis, M. ulcerans
  • Ex: M. fortuitum, M. chelonae

Staining characteristics

Mycobacteria are classical acid-fast organisms.[7] Stains used in evaluation of tissue specimens or microbiological specimens include Fite's stain, Ziehl-Neelsen stain, and Kinyoun stain.

Mycobacteria appear phenotypically most closely related to members of Nocardia, Rhodococcus and Corynebacterium.

Ecological characteristics

Mycobacteria are widespread organisms, typically living in water (including tap water treated with chlorine) and food sources. Some, however, including the tuberculosis and the leprosy organisms, appear to be obligate parasites and are not found as free-living members of the genus.

Pathogenicity

Mycobacteria can colonize their hosts without the hosts showing any adverse signs. For example, billions of people around the world have asymptomatic infections of M. tuberculosis.

Mycobacterial infections are notoriously difficult to treat. The organisms are hardy due to their cell wall, which is neither truly Gram negative nor positive. In addition, they are naturally resistant to a number of antibiotics that disrupt cell-wall biosynthesis, such as penicillin. Due to their unique cell wall, they can survive long exposure to acids, alkalis, detergents, oxidative bursts, lysis by complement, and many antibiotics. Most mycobacteria are susceptible to the antibiotics clarithromycin and rifamycin, but antibiotic-resistant strains have emerged.

As with other bacterial pathogens, surface and secreted proteins of M. tuberculosis contribute significantly to the virulence of this organism. There is an increasing list of extracytoplasmic proteins proven to have a function in the virulence of M. tuberculosis.[8]

Medical classification

Mycobacteria can be classified into several major groups for purpose of diagnosis and treatment: M. tuberculosis complex, which can cause tuberculosis: M. tuberculosis, M. bovis, M. africanum, and M. microti; M. leprae, which causes Hansen's disease or leprosy; Nontuberculous mycobacteria (NTM) are all the other mycobacteria, which can cause pulmonary disease resembling tuberculosis, lymphadenitis, skin disease, or disseminated disease.

Mycosides

Mycosides are phenolic alcohols (such as phenolphthiocerol) that were shown to be components of Mycobacterium glycolipids that are termed glycosides of phenolphthiocerol dimycocerosate (Smith DW et al., Nature 1960, 186, 887) There are 18 and 20 carbon atoms in mycosides A, and B, respectively.[9]

Genomics

Comparative analyses of mycobacterial genomes have identified several conserved indels and signature proteins that are uniquely found in all sequenced species from the genus Mycobacterium.[10][11] Additionally, 14 proteins are found only in the species from the genera Mycobacterium and Nocardia, suggesting that these two genera are closely related.[11]

Species

Phenotypic tests can be used to identify and distinguish different Mycobacteria species and strains. In older systems, mycobacteria are grouped based upon their appearance and rate of growth. However, these are symplesiomorphies, and more recent classification is based upon cladistics.

Slowly growing

Mycobacterium tuberculosis complex

Mycobacterium avium complex

Mycobacterium gordonae clade

Mycobacterium kansasii clade

Mycobacterium nonchromogenicum/terrae clade

Mycolactone-producing mycobacteria

Mycobacterium simiae clade

Ungrouped

Intermediate growth rate

Rapidly growing

Mycobacterium chelonae clade

Mycobacterium fortuitum clade

Mycobacterium parafortuitum clade

Mycobacterium vaccae clade

CF

Ungrouped

Ungrouped

Mycobacteriophage

Mycobacteria can be infected by Mycobacteriophage, bacterial viruses that may be used in the future to treat tuberculosis and related diseases by phage therapy.

References

Further reading

  • Diagnosis and Treatment of Disease Caused by Nontuberculous Mycobacteria. American Thoracic Society. Am J Respiratory and Critical Care Medicine. Aug 1997 156(2) Part 2 Supplement
  • RIDOM: Ribosomal Differentiation of Medical Microorganisms
  • J.P. Euzéby: List of Prokaryotic Names with Standing in Nomenclature -

External links

  • Tuberculist: Genome annotation database
  • MTB Sysborg: Genome annotation database from the Institute of Genomics and Integrative Biology
  • TB Structural Genomics Consortium: Structures of Mycobacterium tuberculosis proteins
  • MycDB: Mycobacterium database
  • TBDB: Tuberculosis database
  • NIAID
  • Frequently Asked Questions about NTM Lung Disease
  • PRASITE: Identification of mycobacteria
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