Lactobacillus

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Lactobacillus
Lactobacillus sp 01.png
Lactobacillus near a squamous epithelial cell
Scientific classification
Domain: Bacteria
Phylum: Firmicutes
Class: Bacilli
Order: Lactobacillales
Family: Lactobacillaceae
Genus: Lactobacillus
Beijerinck 1901
Species
L. acetotolerans
L. acidifarinae
L. acidipiscis
L. acidophilus
L. agilis
L. algidus
L. alimentarius
L. amylolyticus
L. amylophilus
L. amylotrophicus
L. amylovorus
L. animalis
L. antri
L. apodemi
L. aviarius
L. bifermentans
L. brevis
L. buchneri
L. camelliae
L. casei
L. catenaformis
L. ceti
L. coleohominis
L. collinoides
L. composti
L. concavus
L. coryniformis
L. crispatus
L. crustorum
L. curvatus
L. delbrueckii subsp. bulgaricus
L. delbrueckii subsp. delbrueckii
L. delbrueckii subsp. lactis
L. dextrinicus
L. diolivorans
L. equi
L. equigenerosi
L. farraginis
L. farciminis
L. fermentum
L. fornicalis
L. fructivorans
L. frumenti
L. fuchuensis
L. gallinarum
L. gasseri
L. gastricus
L. ghanensis
L. graminis
L. hammesii
L. hamsteri
L. harbinensis
L. hayakitensis
L. helveticus
L. hilgardii
L. homohiochii
L. iners
L. ingluviei
L. intestinalis
L. jensenii
L. johnsonii
L. kalixensis
L. kefiranofaciens
L. kefiri
L. kimchii
L. kitasatonis
L. kunkeei
L. leichmannii
L. lindneri
L. malefermentans
L. mali
L. manihotivorans
L. mindensis
L. mucosae
L. murinus
L. nagelii
L. namurensis
L. nantensis
L. oligofermentans
L. oris
L. panis
L. pantheris
L. parabrevis
L. parabuchneri
L. paracasei
L. paracollinoides
L. parafarraginis
L. parakefiri
L. paralimentarius
L. paraplantarum
L. pentosus
L. perolens
L. plantarum
L. pontis
L. protectus
L. psittaci
L. rennini
L. reuteri
L. rhamnosus
L. rimae
L. rogosae
L. rossiae
L. ruminis
L. saerimneri
L. sakei
L. salivarius
L. sanfranciscensis
L. satsumensis
L. secaliphilus
L. sharpeae
L. siliginis
L. spicheri
L. suebicus
L. thailandensis
L. ultunensis
L. vaccinostercus
L. vaginalis
L. versmoldensis
L. vini
L. vitulinus
L. zeae
L. zymae
Lactobacillus is a genus of Gram-positive, facultative anaerobic or
microaerophilic, rod-shaped, non-spore-forming bacteria.[1] They are a major part
of the lactic acid bacteria group (i.e. they convert sugars to lactic acid). In
humans, they constitute a significant component of the microbiota at a number of
body sites, such as the digestive system, urinary system, and genital system. In
women of European ancestry, Lactobacillus species are normally a major part of the
vaginal microbiota.[2][3][4] Lactobacillus forms biofilms in the vaginal and gut
microbiota, allowing them to persist during harsh environmental conditions and
maintain ample populations.[5] Lactobacillus exhibits a mutualistic relationship
with the human body as it protects the host against potential invasions by
pathogens, and in turn, the host provides a source of nutrients.[6] Lactobacillus
is the most common probiotic found in food such as yogurt, and it is diverse in its
application to maintain human well-being as it can help treat diarrhea, vaginal
infections and skin disorders such as eczema.[7]

Contents
1 Metabolism
2 Genome
3 Taxonomy
4 Human health
4.1 Vaginal tract
4.2 Interactions with other pathogens
4.3 Probiotics
4.4 Oral health
5 Food production
6 See also
7 References
8 External links
Metabolism

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Many lactobacilli operate using homofermentative metabolism (they produce only
lactic acid from sugars), and some species use heterofermentative metabolism (they
can produce either alcohol or lactic acid from sugars).[8] They are aerotolerant
despite the complete absence of a respiratory chain.[9][10] This aerotolerance is
manganese-dependent and has been explored (and explained) in Lactobacillus
plantarum.[11] Many species of this genus do not require iron for growth and have
an extremely high hydrogen peroxide tolerance.[citation needed]

Tryptophan metabolism by human gastrointestinal microbiota (v � t � e)

Tryptophan metabolism diagram
TryptophanClostridium
sporogenesLacto-
bacilliTryptophanase-
expressing
bacteriaIPAI3AIndoleLiverBrainIPAI3AIndoleIndoxyl
sulfateAST-120AhRIntestinal
immune
cellsIntestinal
epitheliumPXRMucosal homeostasis:
?TNF-a
?Junction protein-
coding mRNAsL cellGLP-1T JNeuroprotectant:
?Activation of glial cells and astrocytes
?4-Hydroxy-2-nonenal levels
?DNA damage
�Antioxidant
�Inhibits �-amyloid fibril formationMaintains mucosal reactivity:
?IL-22 productionAssociated with vascular disease:
?Oxidative stress
?Smooth muscle cell proliferation
?Aortic wall thickness and calcificationAssociated with chronic kidney disease:
?Renal dysfunction
�Uremic toxinKidneys
The image above contains clickable links This diagram shows the biosynthesis of
bioactive compounds (indole and certain other derivatives) from tryptophan by
bacteria in the gut.[12] Indole is produced from tryptophan by bacteria that
express tryptophanase.[12] Clostridium sporogenes metabolizes tryptophan into
indole and subsequently 3-indolepropionic acid (IPA),[13] a highly potent
neuroprotective antioxidant that scavenges hydroxyl radicals.[12][14][15] IPA binds
to the pregnane X receptor (PXR) in intestinal cells, thereby facilitating mucosal
homeostasis and barrier function.[12] Following absorption from the intestine and
distribution to the brain, IPA confers a neuroprotective effect against cerebral
ischemia and Alzheimer�s disease.[12] Lactobacillus species metabolize tryptophan
into indole-3-aldehyde (I3A) which acts on the aryl hydrocarbon receptor (AhR) in
intestinal immune cells, in turn increasing interleukin-22 (IL-22) production.[12]
Indole itself triggers the secretion of glucagon-like peptide-1 (GLP-1) in
intestinal L cells and acts as a ligand for AhR.[12] Indole can also be metabolized
by the liver into indoxyl sulfate, a compound that is toxic in high concentrations
and associated with vascular disease and renal dysfunction.[12] AST-120 (activated
charcoal), an intestinal sorbent that is taken by mouth, adsorbs indole, in turn
decreasing the concentration of indoxyl sulfate in blood plasma.[12]
Genome
The genomes of Lactobacillus are highly variable, ranging in size from 1.2 to 3.3
Mb (megabases). Accordingly, the number of protein-coding genes ranges from 1,100
to about 3,200 genes.[16]

Lactobacillus contains a wealth of compound microsatellites in the coding region of

the genome, which are imperfect and have variant motifs.[17]

Taxonomy
The genus Lactobacillus currently contains over 180 species and encompasses a wide
variety of organisms.[18] The genus is polyphyletic, with the genus Pediococcus
dividing the L. casei group, and the species L. acidophilus, L. salivarius, and L.
reuteri being representatives of three distinct subclades. The genus
Paralactobacillus falls within the L. salivarius group. In recent years, other
members of the genus Lactobacillus (formerly known as the Leuconostoc branch of
Lactobacillus) have been reclassified into the genera Atopobium, Carnobacterium,
Weissella, Oenococcus, and Leuconostoc. More recently, the Pediococcus species P.
dextrinicus has been reclassified as a Lactobacillus species.[19] According to
metabolism, Lactobacillus species can be divided into three groups:

Obligately homofermentative (group I) including:

L. acidophilus, L. delbrueckii, L. helveticus, L. salivarius
Facultatively heterofermentative (group II) including:
L. casei, L. curvatus, L. plantarum, L. sakei
Obligately heterofermentative (group III) including:
L. brevis, L. buchneri, L. fermentum, L. reuteri
Human health
Vaginal tract
The female genital tract is one of the principal colonisation sites for human
microbiota, and there is interest in the relationship between the composition of
these bacteria and human health, with a domination by a single species being
correlated with general welfare and good outcomes in pregnancy. In around 70% of
women, a Lactobacillus species is dominant, although that has been found to vary
between American women of European origin and those of African origin, the latter
group tending to have more diverse vaginal microbiota. Similar differences have
also been identified in comparisons between Belgian and Tanzanian women.[2][3][4]

Interactions with other pathogens

Lactobacillus species produce hydrogen peroxide which inhibits the growth and
virulence of the fungal pathogen Candida albicans in vitro and in vivo.[20][21] In
vitro studies have also shown that Lactobacillus sp. reduce the pathogenicity of C.
albicans through the production of organic acids and certain metabolites.[22] Both
the presence of metabolites, such as sodium butyrate, and the decrease in
environmental pH caused by the organic acids reduce the growth of hypha in C.
albicans, which reduces its pathogenicity. [22] Lactobacillus sp. also reduce the
pathogenicity of C. albicans by reducing C. albicans biofilm formation.[22] Biofilm
formation is reduced by both the competition from Lactobacillus sp., and the
formation of defective biofilms which is linked to the reduced hypha growth
mentioned earlier. [22] On the other hand, following antibiotic therapy, certain
Candida species can suppress the regrowth of Lactobacillus sp. at body sites where
they cohabitate, such as in the gastrointestinal tract.[20][21]

In addition to its effects on C. albicans, Lactobacillus sp. also interact with

other pathogens. For example, Lactobacillus reuteri can inhibit the growth of many
different bacterial species by using glycerol to produce the antimicrobial
substance called reuterin. [23] Another example is Lactobacillus salivarius, which
interacts with many pathogens through the production of salivaricin B, a
bacteriocin.[24]

Probiotics
Lactobacillus species administered in combination with other probiotics benefits
cases of irritable bowel syndrome (IBS), although the extent of efficacy is still
uncertain.[25] The probiotics help treat IBS by returning homeostasis when the gut
microbiota experiences unusually high levels of opportunistic bacteria.[6] In
addition, Lactobacillus species can be administered as probiotics during cases of
infection by the ulcer-causing bacterium Helicobacter pylori.[26] Helicobacter
pylori is linked to cancer, and antibiotic resistance impedes the success of
current antibiotic-based eradication treatments.[26] When Lactobacillus probiotics
are administered along with the treatment as an adjuvant, its efficacy is
substantially increased and side effects may be lessened.[26] Also, Lactobacillus
is used to help control urogenital and vaginal infections, such as bacterial
vaginosis (BV). Lactobacillus produce bacteriocins to suppress pathogenic growth of
certain bacteria,[27] as well as lactic acid and H2O2 (hydrogen peroxide). Lactic
acid lowers the vaginal pH to around 4.5 or less, hampering the survival of other
bacteria, and H2O2 reestablishes the normal bacterial flora and normal vaginal pH.
[27] In children, Lactobacillus strains such as L. rhamnosus are associated with a
reduction of atopic eczema, also known as dermatitis, due to anti-inflammatory
cytokines secreted by this probiotic bacteria.[6]

Oral health

Dental caries
Some Lactobacillus species have been associated with cases of dental caries
(cavities). Lactic acid can corrode teeth, and the Lactobacillus count in saliva
has been used as a "caries test" for many years. Lactobacilli characteristically
cause existing carious lesions to progress, especially those in coronal caries. The
issue is, however, complex, as recent studies show probiotics can allow beneficial
lactobacilli to populate sites on teeth, preventing streptococcal pathogens from
taking hold and inducing dental decay. The scientific research of lactobacilli in
relation to oral health is a new field and only a few studies and results have been
published.[28][29] Some studies have provided evidence of certain Lactobacilli
which can be a probiotic for oral health.[30] Some species, but not all, show
evidence in defense to dental caries.[30] Due to these studies, there have been
applications of incorporating such probiotics in chewing gum and lozenges.[30]
There is also evidence of certain Lactobacilli that are beneficial in the defense
of periodontal disease such as gingivitis and periodontitis.[30]

Food production
Some Lactobacillus species are used as starter cultures in industry for controlled
fermentation in the production of yogurt, cheese, sauerkraut, pickles, beer, cider,
kimchi, cocoa, kefir, and other fermented foods, as well as animal feeds. The
antibacterial and antifungal activity of Lactobacillus species rely on production
of bacteriocins and low molecular weight compounds that inhibits these
microorganisms.[31][32]

Sourdough bread is made either spontaneously, by taking advantage of the bacteria

naturally present in flour, or by using a "starter culture", which is a symbiotic
culture of yeast and lactic acid bacteria growing in a water and flour medium. The
bacteria metabolize sugars into lactic acid, which lowers the pH of their
environment, creating a signature "sourness" associated with yogurt, sauerkraut,
etc.

In many traditional pickling processes, vegetables are submerged in brine, and

salt-tolerant Lactobacillus species feed on natural sugars found in the vegetables.
The resulting mix of salt and lactic acid is a hostile environment for other
microbes, such as fungi, and the vegetables are thus preserved�remaining edible for
long periods.

Lactobacilli, especially L. casei and L. brevis, are some of the most common beer
spoilage organisms. They are, however, essential to the production of sour beers
such as Belgian lambics and American wild ales, giving the beer a distinct tart
flavor.

See also
Lactobacillus L. anticaries
Lactic acid fermentation
MRS agar
Pediococcus
Probiotics
Proteobiotics
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External links
Data related to Lactobacillus at Wikispecies
List of species of the genus Lactobacillus
Lactobacillus at Milk the Funk Wiki
Lactobacillus at BacDive - the Bacterial Diversity Metadatabase

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