Lactic Acid Bacteria (LAB)

Taxonomy

The classical approach to bacterial taxonomy was based on morphology and physiology. Currently, molecular characteristics, such as mol% G+C content of the DNA, electrophoretic properties of the gene products, DNA:DNA hybridization studies and sequences of ribosomal RNA (rRNA), have become important taxonomic tools. This has resulted in dramatic changes in taxonomy of the LAB. The classification of LAB remains the focus of intense taxonomic studies.

Phylogenetically the LAB belong to the clostridial branch of the Gram-positive bacteria. They are catalase-negative, non-sporeforming cocci, coccobacilli or rods that have less than 55 mol% G+C content in their DNA.

The bifidobacteria were discovered in 1900 and were included in the genus Lactobacillus as L. bifidus. Only in 1986 they were transfered to the genus Bifidobacterium (Actinomyces subdivision of the Gram-positive bacteria). They are not true lactics because they produce lactic and acetic acids in the ratio of 2:3, they differ from other LAB in the key enzyme of hexose fermentation, and most importantly, they have high (55-57 mol%) G+C content in their DNA.

cellular organisms - Bacteria - Firmicutes - Bacilli - Lactobacillales

• Aerococcaceae
• Carnobacteriaceae
• Enterococcaceae
• Lactobacillaceae
• Leuconostocaceae
• Streptococcaceae

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Brief facts

• Lactic Acid Bacteria (LAB) or Lactics constitute a diverse group of microorganisms associated with plants, meat, and dairy. They are used in the manufacture of dairy products such as acidophilus milk, yogurt, buttermilk, and cheeses. The Lactics are also important commercially in the processing of meats (sausage, cured hams), alcoholic beverages (beer, fortified spirits), and vegetables (pickles, and saukerkraut). Although the LAB have beneficial effects in the food industry, they can be a nuisance as contaminants by producing off-flavors and contributing to spoilage.
• The Lactics have been characterized primarily by their ability to form various isomers of lactic acid from the fermentation of glucose. Lactic acid may be extracted from the fermentation product and a determination made of the ability to optically rotate light. If the rotation is to the right, it is termed Dextrorotary (D-Lactate); if to the left, it is termed Levorotary (L-Lactate), or, if there is a mixture of both D and L, it is termed racemic
• The Lactic Acid bacteria are grouped as either Homofermenters or Heterofermenters based on the end product of their fermentation. The Homofermenters produce lactic acid as the major product of fermentation of glucose. The Heterofermenters produce a number of products besides lactic acid, including carbon dioxide, acetic acid, and ethanol from the fermentation of glucose.
• In general, the LAB may be characterized as Gram-positive, aerobic to facultatively anaerobic, asporogenous rods and cocci which are oxidase, catalase, and benzidine negative, lack cytochromes, do not reduce nitrates to nitrite, are gelatinase negative, and are unable to utilize lactate.
• Orla and Jensen also divided the Lactobacilli into the three groups (the Thermobacteria, Streptobacteria, and the Betabacteria) based on growth temperature and biochemical reactions. Although those three groups have been replaced for the most part, the three names are still in common use and are defined according to growth temperature, ability to ferment pentoses, ability to produce carbon dioxide from glucose or gluconate, requirement for thiamine, formation of lactic acid as a major product of fermentation, Homofermentative or Heterofermentative type of fermentation, reduction of fructose to mannitol, and hydrolysis of arginine.
• The many properties of LAB - ability to provide health benefits, their roles in the food industry (spoilage or starter cultures), potential pathogenicity - are strain-specific. Numerous strains are continuously screened for desirable characteristics.

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Families of order Lactobacillales

• Aerococcaceae
  • Abiotrophia A heterogeneous group of fastidious cocci that show a dependence on pyridoxal hydrochloride analogs for growth (Kanamoto T et al. J Clin Microbiol. 2000).
    • A. defectiva Synonym: Streptococcus defectivus. Associated with pathologies, most commonly infective endocarditis. Referred as nutritionally variant streptococci (NVS).
      Relevant literature
  • Aerococcus Microaerophilic cocci, usually commensal, implicated in various inflammatory pathologies. Most infections are mild, but serious infections, such as such as endocarditis and septicemia/urosepsis, have been described.
    Relevant literature
  • Eremococcus Single species E. coleocola was isolated rom the reproductive ract of horses (Collins MD et al. Int J Syst Bacteriol. 1999).
  • Facklamia Gram-positive cocci, arranged in short chains or diplos, and resembling viridans streptococci on 5% sheep blood agar. Eighteen strains representing four species of Facklamia were isolated from blood cultures, an abscess, bone, cerebrospinal fluid, gall bladder, vaginal swab, and one unknown source (LaClaire L, Facklam R. Antimicrob Agents Chemother. 2000).
  • Globicatella Globicatella spp. isolated from animal sources and human clinical samples. Implicated in purulent infections of domestic animals.
    Relevant literature
  • Ignavigranum Single species I. ruoffiae, facultatively anaerobic coccus, was isolated from human sources (Collins MD et al. Int J Syst Bacteriol. 1999).

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• Carnobacteriaceae
  • Alkalibacterium Haplophilic and alkaliphilic lactic bacteria of marine origin (Ishikawa M et al. Int J Syst Evol Microbiol. 2009). Found in association with some cheeses (Ishikawa M et al. Lett Appl Microbiol. 2007).
  • Alloiococcus Single species A. otitis found in the normal bacterial flora of the outer ear canal. Suspected in potential pathogenicity (Tano K et al. APMIS. 2008).
  • Atopobacter Single species A. phocae, Gram-positive, catalase-negative, facultatively anaerobic, rod-shaped bacterium, was isolated from common seals (Lawson PA et al. Int J Syst Evol Microbiol. 2000).
  • Atopostipes Single species A. suicloacale solated from an underground swine manure storage pit (Cotta MA et al. Anaerobe. 2004).
  • Carnobacterium The genus was proposed by Collins et al. (1987). Commonly associated with meat, particularly poultry; also found in refrigerated, vacuum-packaged, unprocessed beef and lamb; sometimes referred to as "atypical meat lactics"; psychrophilic (grow at refrigeration temp.), non-aciduric, grow at pH 9.5 (Leisner JJ et al. FEMS Microbiol Rev. 2007, Laursen BG et al. Syst Appl Microbiol. 2005 ).
    • C. divergens Synonym: Lactobacillus divergens.
    • C. maltaromaticum Synonyms: Lactobacillus maltaromicus, Lactobacillus piscicola, Lactobacillus carnis, Carnobacterium piscicola.
  • Desemzia Species D. incerta (formerly Brevibacterium incertum, Bacterium incertum) was re-classified from genus Brevibacterium (Stackebrandt E et al. Int J Syst Bacteriol. 1999).
  • Dolosigranulum Single species D. pigrum is a rare gram-positive opportunistic pathogen capable of causing pneumonia and septicemia (Lécuyer H et al. J Clin Microbiol. 2007).
  • Granulicatella Representative species (G. adiacens and G. elegans) were re-classified from genus Abiotrophia, associated with various inflammatory pathologies.
    Relevant literature.
  • Isobaculum Single species I. melis was isolated from the intestine of a badger (Collins MD et al. Int J Syst Evol Microbiol. 2002).
  • Lacticigenium Single species L. naphtae, a halotolerant and motile lactic acid bacterium, was isolated from crude oil (Iino T et al. Int J Syst Evol Microbiol. 2009).
  • Marinilactibacillus M. piezotolerans (Toffin L et al. Int J Syst Evol Microbiol. 2005 ) and M. psychrotolerans (Ishikawa M et al. Int J Syst Evol Microbiol. 2003) are marine lactic acid bacteria isolated from sub-seafloor sediments and marine organisms in Japan.
  • Trichococcus A few species of related lactic bacteria isolated from various media, some of them were re-classified from other genera (Liu JR et al. Int J Syst Evol Microbiol. 2002 ). T. patagoniensis was isolated from penguin guano (Pikuta EV et al. Int J Syst Evol Microbiol. 2006).

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• Enterococcaceae
  • Bavariicoccus Single species B. seileri isolated from the surface and smear water of German red smear soft cheese (Schmidt VS et al. Int J Syst Evol Microbiol. 2009).
  • Catellicoccus Single species C. marimammalium was isolated from porpoise and grey seal (Lawson PA et al. Int J Syst Evol Microbiol. 2006).
  • Enterococcus Homofermentative Gram-positive, facultatively anaerobic cocci. Can grow at 10°C and at 45°C in broth with 6.5% NaCl, at pH 9.6, and survive heating at 60°C for 30 minutes. The genus' name emphasizes intestinal origin of these bacteria. The nosocomial pathogenicity of enterococci has emerged in recent years, as well as increasing resistance to glycopeptide antibiotics (Fisher K, Phillips C. Microbiology. 2009, Giridhara Upadhyaya PM et al. Indian J Med Microbiol. 2009 ).
  • Melissococcus Single species M. plutonius is a causative agent of European foulbrood in honeybee colonies (Waite R et al. Lett Appl Microbiol. 2003).
  • Pilibacter Single species P. termitis was isolated from from the hindgut of the Formosan subterranean termite (Higashiguchi DT et al. Int J Syst Evol Microbiol. 2006).
  • Tetragenococcus Several species of halophilic lactic acid bacteria. T. halophilus can cause sugar thick juice degradation (Justé A et al. Food Microbiol. 2008). T. muriaticus and T. halophilus are important in biopreservation during the manufacture of fermented food products (Kobayashi T et al. J Appl Microbiol. 2004).
  • Vagococcus Motile cocci (Collins MD et al. J Appl Bacteriol. 1989). Found in feces, river water, diseased fish, animal as well as human sources, ground beef and spoilage of cooked schrimp.
    Relevant literature

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• Lactobacillaceae
  • Lactobacillus Strictly fermenative with complex nutritional requirements; grow in and are associated with many different habitats; aciduric or acidophilic, produce pH 4.0 in foods containing a fermentable carbohydrate; often suppress growth or kill other bacteria; important in fermented food manufacture (dairy, meats, sourdough, beer and wine) as well as spoilage.
    • L. plantarum Synonyms: L. arizonensis, Lactobacterium plantarum, Streptobacterium plantarum, L. arabinosus, L. plantari, L. arizonae. Often found in association with cabbage products and with Cheddar cheeses; can cause spoilage of pickles, acidity and off-flavor in wine. Ferments lactose, maltose and mannitol ("typical Streptobacteria"); grows predominantly at 15 °C; forms small chains with short round ends.
      Relevant literature
    • L. delbrueckii Forms an important complex of homofermentative lactics that previously had species status: L. delbrueckii subsp. lactis (synonyms: L. lactis, L. lactis-acidi, Lactobacterium caucasicum var. lactis, Thermobacterium lactis, Bacterium lactis acidi, Bacillus lactis acidi), L. delbrueckii subsp. bulgaricus (synomyms: L. bulgaricus, Thermobacterium bulgaricum), L. delbrueckii subsp. delbrueckii, and L. delbrueckii subsp. indicus. L. delbrueckii subsp. lactis is used as starter culture in the manufacture of Swiss-type cheeses (because of its heat tolerance). L. delbrueckii subsp. bulgaricus is used as starter culture for yoghurt manufacture.
      Relevant literature
    • L. casei Synonyms: L. casei casei, Streptobacterium casei, Bacillus casei a, Bacillus a. Found in association with milk products. Ferments lactose, maltose and mannitol ("typical Streptobacteria"). Forms short to long rods with square ends. Probiotic, used in yoghurts.
      Relevant literature
    • L. helveticus Synonyms: L. suntoryeus, L. helveticum, Plocamobacterium helveticum, Thermobacterium helveticum, Caseobacterium e, Bacillus casei e, Bacillus e. Belongs to Thermobacteria; important probiotic, used in production fermented dairy products, can cause spoilage of beer
      Relevant literature
    • L. curvatus Synonym: Bacterium curvatum. Found in association with red meat and summer sausage. Psychrophilic (grows at temp. 2-8 °C). Occurs in pairs, has a curved shape, and may form unusual closed rings consisting of four cells making up a horse shoe form.
      Relevant literature
    • L. sakei Synonyms: L. sake, L. bavaricus. Found in association with red meat and summer sausage. Psychrophilic (grows at temp. 2-8 °C).
      Relevant literature
    • L. fermentum Synonyms: Lactobacterium fermentum, Bacillus casei d, Bacillus d. Can grow at temp. 45 °C; belongs to arginine and ribose fermenting Betabacteria; important spoilage microorganism of beer.
      Relevant literature
    • L. reuteri Synonyms: Lactobacterium fermentum biotype II, Lactobacterium fermentum subsp. reuteri. Belongs to Betabacteria that can grow at temp. 45 °C.
      Relevant literature
    • L. vaccinostercus Synonyms: Lactobacillus durianis. Belongs to Betabacteria that can grow at 45 °C. Was originally isolated from cow dung. Relevant literature
    • L. sanfranciscensis Synonym: L. sanfrancisco. Important in sour dough. Belongs to group of Betabacteria that does not ferment ribose and hydrolize arginine.
      Relevant literature
    • L. mali Synonym: Lactobacillus yamanashiensis. Alcohol-tolerant (10 to 20%).
      Relevant literature
    • L. alimentarius A meat species that produces acetoin, forms internal granule, non-motile, microaerophilic, homofermentative, produces predominantly L(+) lactic acid; grows at 15 °C but not at 45 °C, and grows in 10% NaCl.
      Relevant literature
    • L. paracasei Found in dairy products and forms cells singly or in chains; non-motile, grows at 10 °C and at 37 °C but not at 40 °C.
      Relevant literature
    • L. farciminis May serve as a sausage starter culture; microaerophilic; grows at presence of 10% NaCl.
      Relevant literature
    • L. brevis Synonyms: Lactobacterium breve, Betabacterium breve, Bacillus casei g, Bacillus g. Belongs to Betabacteria that ferment ribose and hydrolize arginine, important in production of fermented meat and dairy products.
      Relevant literature
    • L. buchneri Synonyms: Lactobacterium buchneri, Ulvina buchneri, Bacterium buchneri, Bacillus buchneri. Belongs to Betabacteria (ferments ribose and hydrolizes arginine).
      Relevant literature
    • L. hilgardii Synonyms: Lactobacillus Type II. Belong to Betabacteria that ferment ribose and hydrolize arginine. Associated with grapes and preserves. Capable of tolerating up to 20% of alcohol.
      Relevant literature
    • L. rhamnosus Synonyms: L. casei subsp. rhamnosus, Lactobacillus casei rhamnosus. Heterofermenter, rod-shaped nonmotile cells.
      Relevant literature
    • L. acidophilus Synonyms: Thermobacterium intestinale, Bacillus acidophilus. Belongs to Thermobacteria. Produces factors inhibiting tumor proliferation; restores intestinal balance after desease or antibiotic treatment; shown to inhibit pathogen such as S. aureus, S. typhimutium, and E.coli. believed to increase lactose tolerance in humans.
      Relevant literature
  • Paralactobacillus Single species P. selangorensis was isolated from chili bo, a Malaysian food ingredient (Leisner JJ et al. Int J Syst Evol Microbiol. 2000 ).
  • Pediococcus Homofermentative, readily distinguished from the other homofermentative LAB by tetrad formation, non-motile, commonly associated with plants and their products such as cabbage and sauerkraut, cucumbers and pickles, grapes and wine.
    Relevant literature
  • Sharpea Single species S. azabuensis a Gram-positive, strictly anaerobic bacterium isolated from the faeces of thoroughbred horses (Morita H et al. Int J Syst Evol Microbiol. 2008).

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• Leuconostocaceae Relevant literature
  • Fructobacillus Several species were re-classified from genus Leuconostoc on a basis of analysis of 16S rRNA sequences and physiology (Endo A, Okada S. Int J Syst Evol Microbiol. 2008).
  • Leuconostoc Predominant genus among the LAB on plants, with L. mesenteroides subsp. mesenteroides as the principal isolate. In fermented foods of plant origin, L. mesenteroides is generally the first organism to grow and to be succeeded by more acid-tolerant lactobacilli. The Leuconostoc has considerable species-specific commercial importance.
    Relevant literature
    • L. mesenteroides In the dairy industry, L. mesenteroides subsp. dextranicum and L. mesenteroides sbsp. cremoris (formerly, L. dextranicum and L. cremoris) are of vital importance as starter cultures. They produce diacetyl, a flavor compound, in the manufacture of such dairy products as butter, buttermilk, and cheeses. L. mesenteroides subsp. mesenteroides has been used in the manufacture of dextrans, which have been used as blood plasma extenders. It is also a dominant species causing contamination of sugar and meat products with the dextran slime.
      Relevant literature
  • Oenococcus
    • O. oeni Synonyms: L. blayaisense, L. oenos. "Wine Leuconostoc", the most important malolactic bacterium, are used to induce malolactic fermentation in wine (de-acidification of wines), grows at pH 4.2 and 4.7, tolerates 10% ethanol.
      Relevant literature
    • O. kitaharae Non-malolactic-fermenting oenococcus isolated from a composting distilled shochu residue (Endo A, Okada S. Int J Syst Evol Microbiol. 2006).
  • Weissella The genus contains several species that were re-classified from Lactobacillus.
    Relevant literature

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• Streptococcaceae
  • Lactococcus Lactic streptococci. The genus was proposed by Schleifer et al. in 1985 to re-classify some species of the genera Streptococcus and Lactobacillus. Lactococci are are homofermentative and exclusively produce L-lactate; generally found on plants and the skins of animals (Casalta E, Montel MC. Int J Food Microbiol. 2008). Lactococci are used widely in dairy industry. The principal concern is reliability and stability of their starter cultures because many of the desirable traits of the lactococci are plasmid-dependent and can be unstable.
    • L. lactis Synonyms: Streptococcus lactis, Bacterium lacti. The sunspecies of L. lactis are the most important of the commercially used LAB; commonly isolated from plant material, but the most recognised habitat is dairy products.
      Relevant literature
    • L. garvieae Synonyms: Streptococcus garvieae, Enterococcus seriolicida. Not used in dairy products, fish pathogen (Vendrell D et al. Comp Immunol Microbiol Infect Dis. 2006).
  • Lactovum Single species L. miscens an aerotolerant, psychrotolerant, mixed-fermentative anaerobe was isolated from acidic forest soil (Matthies C et al. Res Microbiol. 2004).
  • Streptococcus Among the earliest bacteria to be recognized because of their involvement in a large number of human and animal diseases. The generic name Streptococcus was first used by Rosenbach (1884) to describe the chain-forming, coccus-shaped bacteria associated with wound infections. The genus was originally described based on morphological, serological, physiological and biochemical characteristics and contains the highly pathogenic S. pneumoniae, S. pyogenes and S. agalactiae; the intestinal group D streptococci S. faecalis and S. faecium; and economically important starter species S. thermophilus.
    Relevant literature

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About probiotics

• The Food and Agriculture Organization of the United Nations and the World Health Organization have defined probiotics as "live microorganisms which when administered in adequate amounts confer a health benefit on the host" (FAO/WHO 2002). For adequate amount of health benefits, a dose of five billion colony forming units a day (5x10⁹ CFU/day) has been recommended for at least five days.
• Dead bacteria, bacteria-derived products or products of bacterial growth may also have beneficial effect, but they are not considered probiotics because they are not alive when administered.
• The most common probiotics are the Lactic Acid Bacteria, but probiotics also include other bacteria (for example, Bifidobacterium, Bacillus, and Escherichia coli) as well as yeasts such as Saccharomyces boulardii. Probiotics are available in capsules, powder, fermented milks and other fermented foods and drinks.
• An organism used as a probiotic agent may not necessarily be part of what is concidered the "normal microbiota". Some researchers distinguish between autochthonous and allochthonous species. The former microorganisms both present and replicating in the gastrointestinal tract (between 800 and 1000 bacterial phylotypes are believed to be present in the healthy human GI tract) in contrast to the latter, which are usually passing through.
• Prebiotics are non-digestible food ingredients that confer benefits on the host by stimulating probiotic microorganisms in the gastrointestinal tract. Prebiotics are commonly found in, or extracted from, plant material including fruits cereal, and vegetables, but are also present in human milk and colostrum. The best characterized prebiotics include inulin, fructooligosaccharadie, galactooligosaccharide, xylooligosaccharide, isomaltooligosaccharide, beta-glucans, and lactulose. Both probiotics and prebiotics combined together are called Synbiotics.
• To be considered as potential probiotics, the bacteria must be able to tolerate extreme gastrointestinal conditions (acid, bile, enzymes, low levels of oxygen), ability to adhere to gastrointestinal mucosa and competitive exclusion of pathogens.

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Probiotics and health

Based on Lomax AR, Calder PC. 2009.

Immune function

Function	Description	Findings
Phagocytosis	The process by which phagocytic cells (for example, neutrophils and monocytes) engulf foreign cells and debris; this can lead to presentation of antigens on the phagocyte surface and facilitation a cell-mediated immune response.	L. rhamnosus and L. coryniformis enhanced, other probiotics shown mixed results.
Natural Killer (NK) Cell Activity	NK cells are involved in killing infected or tumor cells by causing apoptosis or necrosis.	L. rhamnosus and L. lactis enhanced NK activity.
Cytokine Production by T Lymphocytes	Cytokines, such as Inteferon, Tumor Necrosis Factor alpha, Interleukins, etc. modulate the function of various cell types.	Probiotics appear to have very little effect on the majority of cytokines measured.
Antibody production in response to vaccination	Antibodies are used by the immune system to identify and neutralize foreign objects, such as bacteria and viruses.	Antibody responses have been increased by probiotics in some, but not all, studies.

Probiotics and infection

Condition	Findings
Enterocolitis and sepsis in very low birth weight infants	L. acidophilis reduced the incidence of necrotizing enterocolitis, sepsis and death.
Antibiotic-Associated Diarrhoea	Some Lactobacilli have been shown to reduce risk of developing the diarrhoea in infants and children.
Helicobacter pylori infection	Although there was no significant effect on H. pylori eradication, severity and frequency of overall symptoms were reduced.
Acute Common Childhood Diarrhoea	Probiotic supplementation was reported to reduce the duration of non-rotavirus diarrhoea. L. casei, L. acidophilus, L. helveticus, and S. thermophilus redused incidence of diarrhoea in healthy children.
Travellers' Diarrhoea	A mostly beneficial effect has been observed regarding incidence, risk of frequency of the diarrhoea.

Probiotics and inflammatory condition

Condition	Findings
Irritable Bowel Syndrome (IBS)	Supplementation with Lactobacillus GG decreased abdominal distention, but other gastrointestinal symptoms and abdominal pain were not altered.
Ulcerative Colitis in Adults	L. acidophilus and L. reuteri alleviated symptoms and decreased exacebrations. Other non-LAB probiotics also had some beneficial effects.
Rheumatoid Arthritis	No significan effects were shown.
Allergies (nasal and food allergies, atopic dermatitis)	Neutral and some beneficial results were achieved in various clinical studies in adults and children.

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Possible mechanisms of action of probiotics

• Increased competition with pathogenic bacteria for nutrients and adhesion sites.
• Production of antimicrobial substances such as bacteriocins.
• Production of short-chain fatty acids by fermentation of carbohydrates, which may:
  • reduce pH of the colon which is detrimental to pathogenic bacteria;
  • provide nutrition to colonocytes.
• Reduction of intestinal permeability.
• Modulation of immune function of gastointestinal mucosa.

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Potential probiotic traditional fermented foods

Modified from Rivera-Espinoza Y, Gallardo-Navarro Y. 2010.

Product	Microorganisms	Description
Ben-saalga	LAB	A traditional gruel from Burkina Faso (E.H. Tou et al. 2007).
Boza	Lactobacillus plantarum, L. brevis, L. rhamnosus, L. fermentum, Leuconostoc mesenteroides subsp. dextranium	A popular fermented beverage in Turkey, Albania, Bulgaria, Kyrgyzstan, Macedonia, Montenegro, Bosnia and Herzegovina (Todorov SD et al. 2007).
Ilambazi lokubilisa	LAB	Fermented maize porridge, traditional in Zimbabwe (Gadaga TH et al. 1999).
Kenkey	Lactobacillus plantarum, Pediococcus pentosaceus, Lactobacillus fermentum/reuteri and Lactobacillus brevis	Fermented maize (Olsen A et al. 1995).
Kimmchi	LAB	Korean traditional Baechu (Chinese cabbage) kimchi (health benefits).
Kishk	LAB	Egyptian traditional porrige prepared from a powdery cereal of burghul (cracked wheat) fermented with milk and laban (yoghurt) (Morcos SR et al. 1973).
Koko	W. confusa and Lact. fermentum	African spontaneously fermented millet porridge and drink (Lei V, Jakobsen M. 2004).
Sauerkraut	LAB	Fermented cabbage products in several European countries (health benefits).
Tarhana	Streptococcus thermophilus, Lactobacillus fermentum, Enterococcus faecium, Pediococcus pentosaceus, Leuconostoc pseudomesenteroides, Weissella cibaria, Lactobacillus plantarum, Lactobacillus delbrueckii spp. bulgaricus, Leuconostoc citreum, Lactobacillus paraplantarum and Lactobacillus casei	A traditional Turkish fermented cereal food (Sengun IY 2009).
Tempeh	LAB	A traditional fermented Malay soy product (health benefits).

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References

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