Genus of bacteria
Mycobacterium
is a
genus
of over 190 species in the phylum
Actinomycetota
, assigned its own family,
Mycobacteriaceae
. This genus includes
pathogens
known to cause serious diseases in mammals, including
tuberculosis
(
M. tuberculosis
) and
leprosy
(
M. leprae
) in humans. The
Greek
prefix
myco-
means 'fungus', alluding to this genus'
mold
-like colony surfaces.
[3]
Since this genus has
cell walls
with a waxy lipid-rich outer layer that contains high concentrations of mycolic acid,
[4]
acid-fast staining
is used to emphasize their resistance to acids, compared to other cell types.
[5]
Mycobacterial species are generally aerobic, non-motile, and capable of growing with minimal nutrients. The genus is divided based on each species' pigment production and growth rate.
[6]
While most
Mycobacterium
species are non-pathogenic, the genus' characteristic complex cell wall contributes to evasion from host defenses.
[7]
Microbiology
[
edit
]
Morphology
[
edit
]
Mycobacteria are
aerobic
with 0.2-0.6 μm wide and 1.0-10 μm long
rod shapes
. They are generally
non-motile
, except for the species
Mycobacterium marinum
, which has been shown to be motile within
macrophages
.
[8]
Mycobacteria possess
capsules
and most do not form
endospores
.
M. marinum
and perhaps
M. bovis
have been shown to
sporulate
;
[9]
however, this has been contested by further research.
[10]
The distinguishing characteristic of all
Mycobacterium
species is a thick,
hydrophobic
, and
mycolic acid
-rich cell wall made of
peptidoglycan
and
arabinogalactan
, with these unique components offering targets for new tuberculosis drugs.
[11]
Physiology
[
edit
]
Many
Mycobacterium
species readily grow with minimal nutrients, using
ammonia
and/or
amino acids
as nitrogen sources and
glycerol
as a carbon source in the presence of mineral salts. Temperatures for optimal growth vary between species and media conditions, ranging from 25 to 45 °C.
[6]
Most
Mycobacterium
species, including most clinically relevant species, can be cultured in
blood agar
.
[12]
However, some species grow very slowly due to extremely long reproductive cycles, such as
M. leprae
requiring 12 days per division cycle compared to 20 minutes for some
E. coli
strains.
[13]
Ecology
[
edit
]
Whereas
Mycobacterium tuberculosis
and
M. leprae
are pathogenic, most mycobacteria do not cause disease unless they enter skin lesions of those with pulmonary and/or immune dysfunction, despite being widespread across aquatic and terrestrial environments. Through
biofilm
formation, cell wall resistance to
chlorine
, and association with
amoebas
, mycobacteria can survive a variety of environmental stressors. The agar media used for most
water testing
does not support the growth of mycobacteria, allowing it to go undetected in municipal and hospital systems.
[14]
Genomics
[
edit
]
Hundreds of
Mycobacterium
genomes have been completely sequenced.
[15]
The genome sizes of mycobacteria range from relatively small ones (e.g. in
M. leprae
) to quite large ones, such as that as
M. vulneris
,
encoding 6,653 proteins, larger than the ~6000 proteins of
eukaryotic
yeast
.
[16]
Protein-Coding Genomic Information
Organism
|
Number of Protein Coding Genes
|
M. intracellulare
|
5,289
[17]
|
M. colombiense
|
5,084
[18]
|
M. leprae
|
1,603
[19]
|
M. tuberculosis
|
3,995
[19]
|
M. smegmatis
|
6,602
[20]
|
M. chelonae
|
4,948
[21]
|
Pathogenicity
[
edit
]
Mycobacterium tuberculosis
complex
[
edit
]
Mycobacterium tuberculosis
can remain latent in human hosts for decades after an initial infection, allowing it to continue infecting others. It has been estimated that a third of the world population has latent tuberculosis (TB).
[22]
M. tuberculosis
has many
virulence factors
, which can be divided across lipid and fatty acid metabolism, cell envelope proteins,
macrophage
inhibitors,
kinase
proteins,
proteases
, metal-transporter proteins, and gene expression regulators.
[23]
Several lineages such as
M. t.
var.
bovis
(bovine TB) were considered separate species in the
M, tuberculosis
complex
until they were finally merged into the main species in 2018.
[24]
Leprosy
[
edit
]
The development of
Leprosy
is caused by infection with either
Mycobacterium leprae
or
Mycobacterium lepromatosis
, two closely related bacteria. Roughly 200,000 new cases of infection are reported each year, and 80% of new cases are reported in Brazil, India, and Indonesia.
[25]
M. leprae
infection localizes within the skin macrophages and Schwann cells found in peripheral nerve tissue.
Nontuberculosis
Mycobacteria
[
edit
]
Nontuberculosis Mycobacteria (NTM), which exclude
M. tuberculosis
,
M. leprae
,
and
M. lepromatosis,
can infect mammalian hosts. These bacteria are referred to as "atypical mycobacteria." Although person-to-person transmission is rare, transmission of
M. abscessus
has been observed between patients with
cystic fibrosis
.
[26]
The four primary diseases observed in humans are chronic pulmonary disease, disseminated disease in immunocompromised patients, skin and soft tissue infections, and superficial lymphadenitis. 80-90% of recorded NTM infections manifest as pulmonary diseases.
[27]
M. abscessus
is the most virulent rapidly-growing mycobacterium (RGM), as well as the leading cause of RGM based pulmonary infections. Although it has been traditionally viewed as an opportunistic pathogen like other NTMs, analysis of various virulence factors (VFs) have shifted this view to that of a true pathogen. This is due to the presence of known mycobacterial VFs and other non-mycobacterial VFs found in other prokaryotic pathogens.
[27]
Virulence factors
[
edit
]
Mycobacteria have cell walls with
peptidoglycan
,
arabinogalactan
, and
mycolic acid
; a waxy outer mycomembrane of mycolic acid; and an outermost
capsule
of
glucans
and secreted proteins for virulence. It constantly remodels these layers to survive in stressful environments and avoid host immune defenses. This cell wall structure results in colony surfaces resembling fungi, leading to the genus' use of the Greek prefix
myco-
.
[28]
This unique structure makes
penicillins
ineffective, instead requiring a multi-drug antibiotic treatment of
isoniazid
to inhibit mycolic acid synthesis,
rifampicin
to interfere with transcription,
ethambutol
to hinder arabinogalactan synthesis, and
pyrazinamide
to impede Coenzyme A synthesis.
[7]
Mycobacterial
Infection Information
Organism
|
Common Symptoms of Infection
|
Known Treatments
|
Reported Cases (Region, Year)
|
M. tuberculosis
|
Fatigue, weight loss, fever, hemoptysis, chest pain.
[29]
|
isoniazid INH, rifampin, pyrazinamide, ethambutol.
[30]
|
1.6 Million (Global, 2021)
[31]
|
M. leprae
M. lepromatosis
|
Skin discoloration, nodule development, dry skin, loss of eyebrows and/or eyelashes, numbness, nosebleeds, paralysis, blindness, nerve pain.
[32]
|
dapson, rifampicin, clofazimine.
[32]
|
133,802 (Global, 2021)
[33]
|
M. avium
complex
|
Tender skin, development of boils or pus-filled vesicles, fevers, chills, muscle aches.
[34]
|
clarithromycin, azithromycin, amikacin, cefoxitin, imipenem.
[35]
|
3000 (US, Annual
estimated
)
[36]
|
M. abscessus
complex
|
Coughing, hemoptysis, fever, cavitary lesions.
[37]
|
clarithromycin, amikacin, cefoxitin, imipenem.
[37]
|
Unknown
|
History
[
edit
]
Mycobacteria have historically been categorized through
phenotypic testing
, such as the
Runyon classification
of analyzing growth rate and production of yellow/orange
carotenoid
pigments. Group I contains
photochromogens
(pigment production induced by light), Group II comprises
scotochromogens
(constitutive pigment production), and the
non-chromogens
of Groups III and IV have a pale yellow/tan pigment, regardless of light exposure. Group IV species are "rapidly-growing" mycobacteria compared to the "slowly-growing" Group III species because samples grow into visible colonies in less than seven days.
[6]
Because the
International Code of Nomenclature of Prokaryotes (ICNP)
currently recognizes 195
Mycobacterium
species, classification and identification systems now rely on
DNA sequencing
and
computational phylogenetics
. The major disease-causing groups are the
M. tuberculosis
complex
(
tuberculosis
),
M. avium
complex
(
mycobacterium avium-intracellulare infection
),
M. leprae
and
M. lepromatosis
(
leprosy
), and
M. abscessus
(
chronic lung infection
).
[3]
Microbiologist Enrico Tortoli has constructed a phylogenetic tree of the genus' key species based on the earlier genetic sequencing of Rogall, et al. (1990), alongside new phylogentic trees based on Tortoli's 2017 sequencing of 148
Mycobacterium
species:
[38]
Proposed division of the genus
[
edit
]
Gupta
et al.
have proposed dividing
Mycobacterium
into five genera, based on an analysis of 150 species in this genus. Due to controversy over complicating clinical diagnoses and treatment, all of the renamed species have retained their original identity in the
Mycobacterium
genus as a valid taxonomic synonym:
[40]
[41]
- Mycobacterium
based on the Slowly-Growing Tuberculosis-Simiae clade
- Mycobacteroides
based on the Rapidly-Growing Abscessus-Chelonae clade
- Mycolicibacillus
based on the Slowly-Growing Triviale clade
- Mycolicibacter
based on the Slowly-Growing Terrae clade
- Mycolicibacterium
based on the Rapidly-Growing Fortuitum-Vaccae clade
Diagnosis
[
edit
]
The two most common methods for visualizing these acid-fast bacilli as bright red against a blue background are the
Ziehl-Neelsen stain
and
modified Kinyoun stain
. Fite's stain is used to color
M. leprae
cells as pink against a blue background. Rapid Modified Auramine O Fluorescent staining has specific binding to slowly-growing mycobacteria for yellow staining against a dark background. Newer methods include Gomori-Methenamine Silver staining and
Perioidic Acid Schiff staining
to color
Mycobacterium avium complex
(MAC) cells black and pink, respectively.
[5]
While some mycobacteria can take up to eight weeks to grow visible colonies from a cultured sample, most clinically relevant species will grow within the first four weeks, allowing physicians to consider alternative causes if negative readings continue past the first month.
[42]
Growth media include
Lowenstein?Jensen medium
and
mycobacteria growth indicator tube
(MGIT).
-
Mycobacterium tuberculosis
on
Ziehl-Neelsen stain
-
Slant tubes of Lowenstein-Jensen medium.
[note 1]
-
MGIT
samples emitting fluorescence in ultraviolet light
Mycobacteriophages
[
edit
]
Mycobacteria can be infected by
mycobacteriophages
, a class of viruses with high specificity for their targets. By hijacking the cellular machinery of mycobacteria to produce additional phages, such viruses can be used in
phage therapy
for eukaryotic hosts, as they would die alongside the mycobacteria. Since only some mycobacteriophages are capable of penetrating the
M. tuberculosis
membrane, the viral DNA may be delivered through artificial
liposomes
because bacteria uptake, transcribe, and translate foreign DNA into proteins.
[43]
Mycosides
[
edit
]
Mycosides are
glycolipids
isolated from
Mycobacterium
species with Mycoside A found in photochromogenic strains, Mycoside B in bovine strains, and Mycoside C in avian strains.
[44]
Different forms of Mycoside C have varying success as a receptor to inactivate
mycobacteriophages
.
[45]
Replacement of the gene encoding mycocerosic acid synthase in
M. bovis
prevents formation of mycosides.
[46]
Notes
[
edit
]
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[
edit
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.
PMC
39357
.
PMID
8643481
.
External links
[
edit
]