Lactobacillales

Els Lactobacillales són un ordre de bacteris gram positius, dins la divisió de les Firmicutes. Gèneres representatius són Enterococcus, Streptococcus, Lactococcus, Leuconostoc i Lactobacillus.

Produeixen àcid làctic com el major metabòlit de la fermentació del sucre. Són usats en la producció de productes lactis fermentats, com iogurt, formatge, mantega, crema de llet, crema agra, quefir i koumiss.

Els LAB són també responsables de la fermentació malolàctica en la producció de vi, i en la fermentació del cabdell a sauerkraut xucrut.

Bacteris generadors d'àcid làctic

• Carnobacterium
• Enterococcus
• Lactobacillus
• Lactococcus
• Leuconostoc
• Oenococcus
• Pediococcus
• Streptococcus
• Weissella

A Wikimedia Commons hi ha contingut multimèdia relatiu a: Lactobacil·lal

Ang Lactobacillales ay isang uri ng pamilya sa bakterya kahariang Protista. Ito ay Gram-positive bacteria.

Ang lathalaing ito na tungkol sa Bakterya ay isang usbong. Makatutulong ka sa Wikipedia sa nito.

D Milchsüürbakterie (au Lactobacillales oder Suurmilchbakterie) bilde en Ordnig vo grampositive Bakterie, wo fakultativ anaerob si aber mäistens aerotolerant. Si baue Choolegüdrat zu Milchsüüri ab im ene Brozäss, wo mä as Milchsüürigäärig bezäichnet. Milchsüüribakterie chömme bi Mensche und bin e Hufe Dier vor. Si besiidle under anderem dr Verdauigsdrakt. Au in dr Milch cha mä sä finde. Wenn s gnueg von ene het, foot d Milch afo dick wärde wäge dr Milchsüüri, wo si broduziere. Das isch d Grundlag für d Härstellig vo Joghurt, Chääs und andere Milchbrodukt. Näbe de Milchsüüribakterie, wo industriell gnützt wärde, git s in dere Ordnig au Erreger vo Chrankhäite, z. B. e baar Streptococcus-Arte (Streptokokke).

Litratuur

• Martin Dworkin, Stanley Falkow, Eugene Rosenberg, Karl-Heinz Schleifer, Erko Stackebrandt (Hrsg.): The Prokaryotes. A Handbook on the Biology of Bacteria, Volume 4: Bacteria: Firmicutes, Cyanobacteria. 3. Auflage. Springer-Verlag, New York 2006, ISBN 978-0-387-25494-4.
• Paul Vos, George Garrity, Dorothy Jones, Noel R. Krieg, Wolfgang Ludwig, Fred A. Rainey, Karl-Heinz Schleifer, William B. Whitman: Bergey's Manual of Systematic Bacteriology: Volume 3: The Firmicutes. 2. Auflage. Springer-Verlag, New York 2009, ISBN 978-0-387-95041-9.

Sut kislotali bakteriyalar (Lactobacterium) — sut kislotali bijgʻishni paydo qiladigan grammusbat tayoqchalar va kokklar. Sut kislotali bakteriyalarb. uglevodlarni sut kislotagacha bijgitadi; spora hosil qilmaydi, sut, sut mahsulotlari, oʻsimliklar, oʻsimlik qoldiklari, odam va hayvonlar ichagida uchraydi. Gomofermentativ Sut kislotali bakteriyalarb. kandni parchalab, asosan, sut kislotasi hosil qiladi, ulardan qatiq mahsulotlari tayyorlashda, sanoat mikiyosida sut kislotasi olishda foydalaniladi. Geteroferment Sut kislotali bakteriyalarb. uglevodlarni sut kislota, sirka kislota SO2 baʼzan etanolgacha parchalaydi. Yemxashaklarni siloslash, karamni tuzlashda ishtirok etadi. Sut kislotali bakteriyalardan sunʼiy qon plazmasi tayyorlash uchun zarur boʻladigan dekstranlar olishda foydalaniladi.

लैक्टोबैसिलेलीस (Lactobacillales) या लैक्टिक अम्ल बैक्टीरिया (lactic acid bacteria, LAB) ग्राम-धनात्मक बैक्टीरिया का एक जीववैज्ञानिक गण है, जो बैक्टीरिया के फ़र्मीक्यूटीस संघ के बैसिलाए वर्ग का भाग है। य बैक्टीरिया अक्सर अपघटित हो रहे वनस्पतियों और दूध व दूध उत्पादनों में पाए जाते हैं और इन चीज़ों पर अपनी चयापचय प्रक्रिया में कार्बोहाइड्रेट किण्वन (फ़र्मेन्टेशन) कर के लैक्टिक अम्ल बनाते हैं। मानव इतिहास में लैक्टोबैसिलेलीस की कुछ जातियों का प्रयोग इनके इस गुण के लिए करा गया है, क्योंकि खाद्य पदार्थों का अम्लीकरण उनकी ख़राब होने की प्रक्रिया को धीमा कर देता है और उनमें रोगजनक सूक्ष्मजीवियों की बढ़त में भी बाधा डालता है।^[1]

इन्हें भी देखें

• फ़र्मीक्यूटीस
• बैसिलाए
• लैक्टिक अम्ल

सन्दर्भ

Lactobacillales are an order of gram-positive, low-GC, acid-tolerant, generally nonsporulating, nonrespiring, either rod-shaped (bacilli) or spherical (cocci) bacteria that share common metabolic and physiological characteristics. These bacteria, usually found in decomposing plants and milk products, produce lactic acid as the major metabolic end product of carbohydrate fermentation, giving them the common name lactic acid bacteria (LAB).

Production of lactic acid has linked LAB with food fermentations, as acidification inhibits the growth of spoilage agents. Proteinaceous bacteriocins are produced by several LAB strains and provide an additional hurdle for spoilage and pathogenic microorganisms. Furthermore, lactic acid and other metabolic products contribute to the organoleptic and textural profile of a food item. The industrial importance of the LAB is further evidenced by their generally recognized as safe (GRAS) status, due to their ubiquitous appearance in food and their contribution to the healthy microbiota of animal and human mucosal surfaces.

The genera that comprise the LAB are at its core Lactobacillus, Leuconostoc, Pediococcus, Lactococcus, and Streptococcus, as well as the more peripheral Aerococcus, Carnobacterium, Enterococcus, Oenococcus, Sporolactobacillus, Tetragenococcus, Vagococcus, and Weissella. All but Sporolactobacillus are members of the Lactobacillales order, and all are members of the Bacillota phylum.

Although lactic acid bacteria are generally associated with the order Lactobacillales, bacteria of the genus Bifidobacterium (phylum Actinomycetota) also produce lactic acid as the major product of carbohydrate metabolism.^[1]

Characteristics

The lactic acid bacteria (LAB) are either rod-shaped (bacilli), or spherical (cocci), and are characterized by an increased tolerance to acidity (low pH range). This aspect helps LAB to outcompete other bacteria in a natural fermentation, as they can withstand the increased acidity from organic acid production (e.g., lactic acid). Laboratory media used for LAB typically include a carbohydrate source, as most species are incapable of respiration. LAB are catalase-negative. LAB are amongst the most important groups of microorganisms used in the food industry.^[2] Their relative simple metabolism has also prompted their use as microbial cell factories for the production of several commodities for the food and non-food sectors ^[3]

Metabolism

LAB genera are classified in terms of two main pathways of hexose fermentation:

1. Under conditions of excess glucose and limited oxygen, homolactic LAB catabolize one mole of glucose in the Embden-Meyerhof-Parnas pathway to yield two moles of pyruvate. Intracellular redox balance is maintained through the oxidation of NADH, concomitant with pyruvate reduction to lactic acid. This process yields two moles of ATP per mole of glucose consumed. Representative homolactic LAB genera include Lactococcus, Enterococcus, Streptococcus, Pediococcus, and group I lactobacilli ^[4]
2. Heterofermentative LAB use the pentose phosphate pathway, alternatively referred to as the pentose phosphoketolase pathway. One mole of glucose-6-phosphate is initially dehydrogenated to 6-phosphogluconate and subsequently decarboxylated to yield one mole of CO₂. The resulting pentose-5-phosphate is cleaved into one mole glyceraldehyde phosphate (GAP) and one mole acetyl phosphate. GAP is further metabolized to lactate as in homofermentation, with the acetyl phosphate reduced to ethanol via acetyl-CoA and acetaldehyde intermediates. In theory, end products (including ATP) are produced in equimolar quantities from the catabolism of one mole of glucose. Obligate heterofermentative LAB include Leuconostoc, Oenococcus, Weissella, and group III lactobacilli ^[4]

Some members of Lactobacillus appear also able to perform aerobic respiration, making them facultative anaerobes, unlike the other members of the order, which are all aerotolerant. Using oxygen helps these bacteria deal with stress.^[5]

Streptococcus reclassification

In 1985, members of the diverse genus Streptococcus were reclassified into Lactococcus, Enterococcus, Vagococcus, and Streptococcus based on biochemical characteristics, as well as molecular features. Formerly, streptococci were segregated primarily based on serology, which has proven to correlate well with the current taxonomic definitions. Lactococci (formerly Lancefield group N streptococci) are used extensively as fermentation starters in dairy production, with humans estimated to consume 10¹⁸ lactococci annually. Partly due to their industrial relevance, both L. lactis subspecies (L. l. lactis and L. l. cremoris) are widely used as generic LAB models for research. L. lactis ssp. cremoris, used in the production of hard cheeses, is represented by the laboratory strains LM0230 and MG1363. In similar manner, L. lactis ssp. lactis is employed in soft cheese fermentations, with the workhorse strain IL1403 ubiquitous in LAB research laboratories. In 2001, Bolotin et al. sequenced the genome of IL1403, which coincided with a significant shift of resources to understanding LAB genomics and related applications.

Phylogeny

The currently accepted taxonomy is based on the List of Prokaryotic names with Standing in Nomenclature^[6][7] and the phylogeny is based on 16S rRNA-based LTP release 106 by 'The All-Species Living Tree' Project.^[8]

Lactobacillales

Aerosphaera taetra ♠ Hutson & Collins 2000

Carnococcus allantoicus ♠ Tanner et al. 1995

Aerococcaceae

Granulicatella Collins and Lawson 2000

Atopobacter phocae Lawson et al. 2000

Bavariicoccus Schmidt et al. 2009

Trichococcus Scheff et al. 1984 emend. Liu et al. 2002

Lactobacillus algidus Kato et al. 2000

Lactobacillus species group 1

Lactobacillus species group 2 Beijerinck 1901 emend. Cai et al. 2012

Leuconostocaceae

Lactobacillus species group 3

Lactobacillus species group 4

Lactobacillus species group 5

Lactobacillus species group 6

Pediococcus Claussen 1903

Lactobacillus species group 7

Carnobacterium Collins et al. 1987

Isobaculum melis Collins et al. 2002

Carnobacteriaceae 2 [incl. various Carnobacterium sp.]

Desemzia (Steinhaus 1941) Stackebrandt et al. 1999

Enterococcaceae & Streptococcaceae

(continued)

Lactobacillales part 2 (continued)

Lactobacillales part 2

Vagococcus fessus Hoyles et al. 2000

Vagococcus Collins et al. 1990

Catellicoccus marimammalium Lawson et al. 2006

Enterococcus species group 1 (ex Thiercelin and Jouhaud 1903) Schleifer and Kilpper-Bälz 1984

Enterococcus phoeniculicola Law-Brown and Meyers 2003

Enterococcus species group 2 [incl. Melissococcus plutonius & Tetragenococcus]

Enterococcus species group 3

Enterococcus species group 4

Enterococcus raffinosus Collins et al. 1989

Enterococcus avium (ex Nowlan and Deibel 1967) Collins et al. 1984

Enterococcus pallens Tyrrell et al. 2002

Enterococcus hermanniensis Koort et al. 2004

Pilibacter Higashiguchi et al. 2006

Streptococcaceae

Notes:
♠ Strains found at the National Center for Biotechnology Information, but not listed in the List of Prokaryotic names with Standing in Nomenclature

Uses

Probiotics

Probiotics are products aimed at delivering living, potentially beneficial, bacterial cells to the gut ecosystem of humans and other animals, whereas prebiotics are indigestible carbohydrates delivered in food to the large bowel to provide fermentable substrates for selected bacteria. Most strains used as probiotics belong to the genus Lactobacillus. (Other probiotic strains used belong to the genus Bifidobacterium).^[2][9]

Probiotics have been evaluated in research studies in animals and humans with respect to antibiotic-associated diarrhea, travellers' diarrhea, pediatric diarrhea, inflammatory bowel disease, irritable bowel syndrome^[10] and Alzheimer's disease.^[11] Future applications of probiotics have been conjectured to include delivery systems for vaccines and immunoglobulins, and the treatment of different gastrointestinal diseases and vaginosis.^[10]

Foods

The quest to find food ingredients with valuable bioactive properties has encouraged interest in exopolysaccharides from LAB. Functional food products that offer health and sensory benefits beyond their nutritional composition are becoming progressively more important to the food industry. The sensory benefits of exopolysaccharides are well established, and there is evidence for the health properties that are attributable to exopolysaccharides from LAB. However, there is a wide variation in molecular structures of exopolysaccharides and the complexity of the mechanisms by which physical changes in foods and bioactive effects are elicited.^[12]

Some LAB produce bacteriocins which limit pathogens by interfering with cell wall synthesis or causing pore formation in the cell membrane.^[13] Nisin, a bacteriocin produced by LAB, was first researched as a food preservative in 1951 and has since been widely commercially used in foods due to its antimicrobial activity against Gram positive bacteria.^[14] Nisin is utilized as a food additive in at least 50 countries.^[14] In addition to having antibacterial activity, LAB can inhibit fungal growth. Various LAB, largely from genus Lactococcus and Lactobacillus, suppress mycotoxigenic mold growth due to the production of anti-fungal metabolites.^[15] Furthermore, LAB have the potential to reduce the abundance of mycotoxins in foods by binding to them.^[15] In a study for postharvest food product safety conducted with 119 LAB isolated from the rhizosphere of olive trees and desert truffles, mostly within the genera of Enterococcus and Weissella, researchers found strong antibacterial activity against Stenotrophomonas maltophilia, Pantoea agglomerans, Pseudomonas savastanoi, Staphylococcus aureus and Listeria monocytogenes, and anti-fungal activity against Botrytis cinerea, Penicillium expansum, Verticillium dahliae and Aspergillus niger.^[16]

Fertilizer

Researchers have studied the impact of lactic acid bacteria on indoleacetic acid production, phosphate solubilization, and nitrogen fixation on citrus. While most of the bacterial isolates, were able to produce IAA, phosphate-solubilization was limited to only one of the eight LAB isolates.^[17]

Fermentation

Lactic acid bacteria are used in the food industry for a variety of reasons such as the production of cheese and yogurt products. Popular drinks such as kombucha are made using lactic acid bacteria, with kombucha having been known to have traces of Lactobacillus and Pediococcus once the drink is made.^[18]

The beer and wine-making process utilizes certain lactic acid bacteria, mostly Lactobacillus. Lactic acid bacteria is used to start the wine-making process by starting the malolactic fermentation. After the malolactic fermentation, yeast cells are used to start the alcoholic fermentation process in grapes. The malolactic fermentation mechanism is mainly transformation of L-malic acid (dicarboxylic acid) to an lactic acid (monocarboxylic acid).^[19] This change occurs due to the presence of malolactic and malic enzymes. All malic acid are degraded and this increase the pH levels which changes the taste of the wine.^[19] Not only do they start the process but they are responsible for the different aromas produced in wine by the nutrients presence and the quality of the grapes. Also, the presence of different strains can change the desirability of aromas' presence. The different availability of enzymes that contribute to the vast spectrum of aromas in wine are associated with glycosidases, β-glucosidases, esterases, phenolic acid decarboxylases and citrate lyases.^[20]

By using molecular biology, researchers can help pick out different desirable strains that help improve the quality of wine and help with the removable of the undesirable strains. The same can be said about brewing beer as well which uses yeast with some breweries using lactic acid bacteria to change the taste of their beer.^[21]

Management of bacteriophages in industry

A broad number of food products, commodity chemicals, and biotechnology products are manufactured industrially by large-scale bacterial fermentation of various organic substrates. Because this involves cultivating enormous quantities of bacteria each day in large fermentation vats, a serious threat in these industries is the risk of contamination by bacteriophages, which can rapidly bring fermentations to a halt and cause economical setbacks. Areas of interest in managing this risk include the sources of phage contamination, measures to control their propagation and dissemination, and biotechnological defense strategies developed to restrain them. In the context of the food fermentation industry, the relationship between bacteriophages and their bacterial hosts is very important. The dairy fermentation industry has openly acknowledged the problem of phage contamination, and has worked for decades with academia and starter-culture manufacturers to develop defence strategies and systems to curtail phages' propagation and evolution.^[22]

Bacteriophage–host interaction

The first contact between an infecting phage and its bacterial host is the phage's attaching to the host cell. This attachment is mediated by the phage's receptor binding protein (RBP), which recognizes and binds to a receptor on the bacterial surface. RBPs are also referred to as host-specificity proteins, host determinants, and antireceptors. A variety of molecules have been suggested to act as host receptors for bacteriophages infecting LAB; among those are polysaccharides and (lipo)teichoic acids, as well as a single-membrane protein. A number of RBPs of LAB phages have been identified by the generation of hybrid phages with altered host ranges. These studies, however, also found additional phage proteins to be important for successful phage infection. Analysis of the crystal structure of several RBPs indicates that these proteins share a common tertiary folding, and support previous indications of the saccharide nature of the host receptor. Gram-positive LAB have a thick peptidoglycan layer, which must be traversed to inject the phage genome into the bacterial cytoplasm. Peptidoglycan-degrading enzymes are expected to facilitate this penetration, and such enzymes have been found as structural elements of a number of LAB phages.^[22]

Lactic acid bacteria and dental plaque

LAB are able to synthesize levans from sucrose, and dextrans from glucose.^[23] Dextrans, like other glucan, enable bacteria to adhere to the surface of teeth, which in turn can cause tooth decay through the formation of dental plaque and production of lactic acid.^[24] While the primary bacteria responsible for tooth decay is Streptococcus mutans, LAB do feature among the other most common oral bacteria that cause decay.^[25]

Lactic acid bacteria genera

• Abiotrophia
• Aerococcus
• Carnobacterium
• Enterococcus
• Lactobacillus
• Lactococcus
• Leuconostoc
• Oenococcus
• Pediococcus
• Streptococcus
• Tetragenococcus
• Vagococcus
• Weissella

See also

• Gaffkaemia
• Malolactic fermentation
• Lactic acid fermentation
• Lacto-2 RNA motif

References

Laktobacilaloj (latine lactobacillales) estas ordo de Gram-pozitivaj bakterioj, kiuj karakteriziĝas per tio ke ili digestas sukerojn kaj aliajn karbonhidratojn, kaj en tiu digesto kreas laktoacidon.

Ecoj

Morfologie la unuopaj genroj kaj specioj de laktobacilaloj apenaŭ similas, sed ili fiziologie unuece priskribeblas, ĉar ĉiuj por atingo de energio bezonas karbonhidratojn. La grupo de homofermentativaj bakterioj en tio kreas nur laktoacidon, la grupo de heterofermentativaj bakterioj krom tio ankaŭ aliajn finproduktojn, plejparte etanolo kaj karbondioksido. Per sia specialiĝo de kreskado en lakto kaj aliaj simile medioj riĉaj je karbonhidratoj, ekzemple en la korpoj de homoj kaj de aliaj mamuloj, la laktobacilaloj perdis la eblecon de sintezo de multaj vivnecesaj substancoj, ekzemple de aminacidoj aŭ la por la spirado necesaj porfirinoj kaj citokromoj. Inter alie ili ne kreas enzimojn nomatajn "katalazoj", tio signifas ke ili ne kapablas disigi la kombinaĵon H₂O₂ (hidrogenperoksido). Sekve la laktobacilaloj troveblas nur en la intesta sistemo respektive en mukozoj de mamuloj, kaj en lakto kaj ĉiuj lokoj kiuj havis kontakton kun lakto.

Signifo por homoj

Laktobacilaloj kutime troveblas en la homa intestaro kaj vagino, kaj helpas tie teni iom acidan medion. Iuj specioj kaŭras malsanojn, ekzemple el la genro de streptokokoj la specio streptococcus pneumoniae estas inter tiuj bakterioj kiuj povas kaŭzi pneŭmonion, kaj la specio streptococcus mutans kontribuas en la ekesto de kario en la homa dentaro.

Multaj specioj kaj subspecioj de laktobacilaloj uzatas en la konservigo de nutraĵoj, ekzemple de jogurto, kefiro, acida lakto kaj fromaĝo, sed ankaŭ en acida brasiko, kimĉio kaj en bakaĵoj.

Klasifikigo

Al la ordo de laktobacilaloj apartenas la biologiaj familioj

• Lactobacillaceae, inter ili la laktobaciloj, ekzemple lactobacillus bulgaricus
• Aerococcaceae
• Carnobacteriaceae
• Enterococcaceae
• Leuconostocaceae
• Streptococcaceae, inter ili la streptokokoj
• kaj pliaj, ne klare kateorigitaj unuopaj specioj (latine nomataj incertae sedis).

Gravaj specioj estas

• lactobacillus acidophilus, ero de la normala vagina bakeriaro,
• lactobacillus bulgaricus, uzata en farado de jogurto kaj kefiro,
• lactobacillus casei, grava en farado de fromaĝo, kazeo kaj aliaj laktaĵoj,
• lactobacillus helveticus, same grava en farado de fromaĝoj,

Lactobacillales es un orden de bacterias Gram-positivas, dentro de la División Firmicutes. Géneros representativos son Enterococcus, Streptococcus, Lactococcus, Leuconostoc y Lactobacillus.

Producen ácido láctico como el mayor metabolito de la fermentación del azúcar. Son usados en la producción de productos lácteos fermentados, como yogur, queso, mantequilla, crema de leche, kéfir y koumiss.

Las LAB son también responsables de la fermentación maloláctica en la producción de vino, y en la fermentación del repollo a sauerkraut chucrut.

Bacterias generadoras de ácido láctico

• Carnobacterium
• Enterococcus
• Lactobacillus
• Lactococcus
• Leuconostoc
• Oenococcus
• Pediococcus
• Streptococcus

Filogenia

El análisis filogenético de ARNr 16S muestra que varias familias son parafiléticas, las cuales están indicadas entre comillas:^[1]​

Lactobacillales  

Aerococcaceae

     

"Carnobacteriaceae"

       

"Enterococcaceae"

   

Streptococcaceae

       

"Lactobacillaceae"

   

Leuconostocaceae

         

Referencias

Les Lactobacillales sont un ordre de bactéries à gram positif, placé parmi les Firmicutes. Elles comprennent les bactéries lactiques.

Dans la classification phylogénétique actuellement acceptée^[2], cet ordre est divisé en six familles : Aerococcaceae, Carnobacteriaceae, Enterococcaceae, Lactobacillaceae, Leuconostocaceae, Streptococcaceae.

Très largement présentes dans la nature, on les trouve aussi bien dans le sol, l'eau, que chez les plantes et les animaux. Elles sont utilisées depuis le néolithique par l'homme, pour la fabrication d'aliments fermentés.

Écologie

Les lactobacillales se rencontrent dans l'eau, les égouts, les plantes et les animaux. Elles sont largement employées en agro-alimentaires.

Ils comportent la famille type des Lactobacillaceae dont de nombreux éléments font partie du microbiote de la bouche, du tractus gastro-intestinal et du vagin des animaux homéothermes (l'homme y compris). Ces bacilles sont présents dans les aliments comme les produits laitiers, la viande, le poisson, les céréales, les fruits, les jus de fruits, les légumes, les sauces et les plats cuisinés^[3]. Ils servent à la production d'aliments fermentés comme les fromages, les yaourts, le kéfir, le pain au levain, le vin, la bière et le cidre où ils acidifient le substrat et inhibent ainsi la croissance d'autres organismes. Les Aerococcaceae ont été trouvés dans des habitats variés : air, poussière, végétation, sol. Les Carnobacteriaceae se trouvent dans la viande emballée sous vide, le fromage, le poisson, les lacs antarctiques méromictiques^[4]. Les Enterococcaceae font partie de la flore intestinale des mammifères et des oiseaux. Les Leuconostocaceae se trouvent sur les plantes et dans divers aliments comme le lait et les produits laitiers fermentés (fromages, kéfir), les végétaux fermentés (vin, cidre, olives, choucroute) et la viande. Beaucoup de Streptococcaceae sont des commensaux ou des parasites des animaux, certains étant très pathogènes^[4].

Caractères bactériologiques

Les lactobacillales sont des cellules de formes variées : coques, bâtonnets ou de forme intermédiaire entre les coques rondes et les bacilles en bâtonnets (dites coccobacilles)

Ils puisent leur énergie de l’oxydation de composés organiques. Ce sont des chimio-organotrophes^[3].

• à Gram positif
• non sporulé, généralement non mobile
• aéro-anaéorobie, aérobie facultatif ou strict ou micro-aérophile
• Oxydase négatif (absence de l'enzyme oxydase capable d'oxyder)
• Catalase négatif (absence de l'enzyme catalase capable de décomposer l'eau oxygénée)

Phylogénie

Suivant la taxonomie généralement acceptée basée sur la List of Prokaryotic names with Standing in Nomenclature (LSPN) ^[2],[5], l'ordre des Lactobacillales comprend six familles :

Lactobacillales Familles Genres Aerococcaceae Abiotrophia - Aerococcus - Dolosicoccus - Eremococcus - Facklamia - Globicatella - Ignavigranum Carnobacteriaceae Agitococcus - Alkalibacterium - Allofustis - Alloiococcus - Atopobacter - Atopococcus - Atopostipes - Carnobacterium - Desemzia - Dolosigranulum - Granulicatella - Isobaculum - Lacticigenium - Lactosphaera - Marinilactibacillus - Pisciglobus - Trichococcus Enterococcaceae Bavariicoccus - Catellicoccus - Enterococcus - Melissococcus - Pilibacter - Tetragenococcus - Vagococcus Lactobacillaceae Lactobacillus - Paralactobacillus - Pediococcus - Sharpea Leuconostocaceae Fructobacillus - Leuconostoc - Oenococcus - Weissella Streptococcaceae Lactococcus - Lactovum - Streptococcus

L'ordre des Lactobacillales initialement proposé en 2005, a été décrit en 2009 dans le volume 3 de la deuxième édition du Bergey's Manual of Systematic Bacteriology (Ludwig et al.) et validé en mars 2010 par inscription sur la liste de validation 132.

L'ordre des Lactobacillales est défini principalement sur la base des analyses phylogénétiques déduites de l'étude des séquences des ARNr 16S.

Notes

Références

• ITIS
• SNOMED

Lactobacillales é unha orde de bacterias grampositivas que comprende as bacterias do ácido láctico. Encóntranse no solo, auga, plantas e animais. Utilízanse na produción de alimentos fermentados, como iogur, queixo, manteiga, quefir etc., nos que orixinan a formación de ácido láctico a partir de azucres. Tamén son responsables da fermentación maloláctica na produción do viño, e fermentan os repolos para facer sucrut. Hai tamén especies patóxenas como Streptococcus pneumoniae ou Enterococcus faecalis.

Características

• As Lactobacillales teñen células de formas variadas: cocos, bacilos ou cocobacilos.
• Obteñen a enerxía da oxidación de compostos orgánicos.^[1]
• Son grampositivas
• Non esporuladas, xeralmente non móbiles
• Oxidase negativas (ausencia do encima oxidase)
• Catalase negaivas (ausencia do encima catalase).

Ademais, o riboswitch de caixa S_MK encóntrase só na orde Lactobacillales.

Familias

Comprenden as familias Aerococcaceae, Carnobacteriaceae, Enterococcaceae, Lactobacillaceae, Leuconostocaceae, e Streptococcaceae.

A familia tipo é a das Lactobacillaceae que ten numerosas especies que forman parte da microbiota da boca, tracto gastrointestinal e vaxina dos animais homeotermos. Estes bacilos están presentes en alimentos lácteos, carne, peixe, cereais, froitas, zumes de froitas, legumes, prebes e pratos cociñados.^[1] Utilízanse na produción de alimentos fermentados como o queixo ou iogur e outros. As Aerococcaceae encóntranse no aire, po, vexetación e solo. As Carnobacteriaceae encóntranse na carne envasada ao baleiro, o queixo, peixe ou nos lagos antárticos meromícticos^[2]. As Enterococcaceae forman parte da flora intestinal dos mamíferos e aves, e algunhas son patóxenas, como Enterococcus faecalis. As Leuconostocaceae encóntranse nas plantas e en diversos alimentos como o leite e os produtos lácteos fermentados, produtos vexetais fermentados e carne. Moitas Streptococcaceae son comensais ou parasitos de animais, e algunhas moi patóxenas como Streptococcus pneumoniae.^[2].

Filoxenia

A orde Lactobacillales foi proposta inicialmente en 2005, e foi descrita en 2009 no volume 3 da segunda edición do Manual de bacterioloxía sistemática de Bergey (Ludwig et al.) e validado en marzo de 2010 e inscrito na lista de validación 132. A taxonomía aceptada actualmente desta orde está baseada na LPSN ^{[3] [4]} e a filoxenia foi elaborada segundo as secuencias do ARNr 16S da LTP 106 do 'The All-Species Living Tree' Project ^[5]

Lactobacillales  

Aerosphaera taetra ♠ Hutson & Collins 2000

   

Carnococcus allantoicus ♠ Tanner et al. 1995

   

Aerococcaceae

     

Granulicatella Collins e Lawson 2000

       

Atopobacter phocae Lawson et al. 2000

   

Bavariicoccus seileri Schmidt et al. 2009

       

Trichococcus Scheff et al. 1984 emend. Liu et al. 2002

       

Lactobacillus algidus Kato et al. 2000

     

Lactobacillus grupo de especies 2

     

Lactobacillus Beijerinck 1901 emend. Cai et al. 2012

     

Leuconostocaceae

     

Lactobacillus grupo de especies 3

     

Lactobacillus grupo de especies 4

     

Lactobacillus grupo de especies 5

     

Lactobacillus grupo de especies 6

     

Pediococcus Claussen 1903

   

Lactobacillus grupo de especies 7

                       

Carnobacterium Collins et al. 1987

     

Isobaculum melis Collins et al. 2002

     

Carnobacteriaceae 2 [incl. varios Carnobacterium sp.]

     

Desemzia incerta (Steinhaus 1941) Stackebrandt et al. 1999

  Enterococcaceae & Streptococcaceae

(continúa)

                   

Lactobacillales parte 2 (continuación)

Lactobacillales parte 2  

Vagococcus fessus Hoyles et al. 2000

     

Vagococcus Collins et al. 1990

     

Catellicoccus marimammalium Lawson et al. 2006

     

Enterococcus (ex Thiercelin e Jouhaud 1903) Schleifer e Kilpper-Bälz 1984

     

Enterococcus phoeniculicola Law-Brown e Meyers 2003

     

Enterococcus grupo de especies 2 [incl. Melissococcus plutonius & Tetragenococcus]

     

Enterococcus grupo de especies 3

     

Enterococcus grupo de especies 4

     

Enterococcus raffinosus Collins et al. 1989

   

Enterococcus avium (ex Nowlan e Deibel 1967) Collins et al. 1984

     

Enterococcus pallens Tyrrell et al. 2002

     

Enterococcus hermanniensis Koort et al. 2004

     

Pilibacter termitis Higashiguchi et al. 2006

   

Streptococcaceae

                         

Notas:
♠ Cepas que se encontran no NCBI pero non están na LPSN.

Xéneros de bacterias do ácido láctico

• Abiotrophia
• Aerococcus
• Carnobacterium
• Enterococcus
• Lactobacillus
• Lactococcus
• Leuconostoc
• Oenococcus
• Pediococcus
• Streptococcus
• Tetragenococcus
• Vagococcus
• Weissella

Notas

Lactobacillales je red Gram-pozitivnih bakterija, široko rasprostranjenih u prirodi.

Nedovršeni članak Lactobacillales koji govori o biologiji treba dopuniti. Dopunite ga prema pravilima Wikipedije.

Lactobacillales er ættbálkur Gram-jákvæðra gerla sem einkennast meðal annars af því að stunda mjólkursýrumyndandi gerjun sem meginleið til orkuefnaskipta. Þeir teljast því til loftfirrtra gerla, en þola þó flestir að súrefni sé til staðar (eru loftþolnir loftfirringar). Þeir eru algengir í vistkerfum þar sem lífræn efni eru auðfengin, svo sem í jarðvegi og rotnandi dýra- og plöntuleifum. Mjólkursýrugerlarnir tilheyra þessum ættbálki og er það hugtak stundum notað sem safnheiti yfir allan ættbálkinn.

I Lactobacillales sono un ordine di batteri Gram-positivi appartenenti alla classe dei Bacilli. Ampiamente diffusi in natura, si trovano infatti nel suolo, nell'acqua, nelle piante e negli animali, sono largamente utilizzati nei processi fermentativi per la produzione di derivati del latte quali yogurt, formaggio, burro, latticello, Kéfir e Kumis.

Intervengono anche nella fermentazione malolattica nella produzione del vino.

Alcuni Lactobacillales

• Abiotrophia
• Aerococcus
• Carnobacterium
• Lactobacillus
• Lactococcus
• Leuconostoc
• Oenococcus
• Pediococcus
• Tetragenococcus
• Vagococcus
• Weissella

Šeimos

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

Lactobacillales – bacilų (Bacilli) klasės bakterijų eilė.

Lactobacillales - gramteigiamos bakterijos.

Klasifikacija

• Šeima. Aerococcaceae
• Šeima. Carnobacteriaceae
• Šeima. Enterococcaceae
  • Gentis. Atopobacter
  • Gentis. Enterococcus
    • Rūšis. Enterococcus avium
    • Rūšis. Enterococcus durans
    • Rūšis. Enterococcus faecalis
    • Rūšis. Enterococcus faecium
    • Rūšis. Enterococcus solitarius
  • Gentis. Melissococcus
  • Gentis. Tetragenococcus
  • Gentis. Vagococcus
• Šeima. Lactobacillaceae
  • Gentis. Lactobacillus
    • Rūšis. Lactobacillus acidophilus
    • Rūšis. Lactobacillus brevis
    • Rūšis. Lactobacillus delbrueckii subsp. bulgaricus
    • Rūšis. Lactobacillus casei
    • Rūšis. Lactobacillus delbrueckii
    • Rūšis. Lactobacillus fermentum
    • Rūšis. Lactobacillus helveticus
    • Rūšis. Lactobacillus plantarum
    • Rūšis. Lactobacillus rhamnosus
    • Rūšis. Lactobacillus reuteri
    • Rūšis. Lactobacillus sanfranciscensis
  • Gentis. Paralactobacillus
  • Gentis. Pediococcus
    • Rūšis. Pediococcus acidilactici
    • Rūšis. Pediococcus cellicola
    • Rūšis. Pediococcus claussenii
    • Rūšis. Pediococcus damnosus
    • Rūšis. Pediococcus dextrinicus
    • Rūšis. Pediococcus ethanolidurans
    • Rūšis. Pediococcus inopinatus
    • Rūšis. Pediococcus parvulus
    • Rūšis. Pediococcus pentosaceus
    • Rūšis. Pediococcus stilesii
• Šeima. Leuconostocaceae
  • Gentis. Leuconostoc
    • Rūšis. Leuconostoc carnosum
    • Rūšis. Leuconostoc citreum
    • Rūšis. Leuconostoc durionis
    • Rūšis. Leuconostoc fallax
    • Rūšis. Leuconostoc ficulneum
    • Rūšis. Leuconostoc fructosum
    • Rūšis. Leuconostoc garlicum
    • Rūšis. Leuconostoc gasicomitatum
    • Rūšis. Leuconostoc gelidum
    • Rūšis. Leuconostoc inhae
    • Rūšis. Leuconostoc kimchii
    • Rūšis. Leuconostoc lactis
    • Rūšis. Leuconostoc mesenteroides
    • Rūšis. Leuconostoc pseudoficulneum
    • Rūšis. Leuconostoc pseudomesenteroides
  • Gentis. Oenococcus
    • Rūšis. Oenococcus oeni
    • Rūšis. Oenococcus kitaharae
  • Gentis. Weissella
• Šeima. Streptococcaceae
  • Gentis. Laktokokai Lactococcus
    • Rūšis. Lactococcus garvieae
    • Rūšis. Lactococcus lactis
    • Rūšis. Lactococcus piscium
    • Rūšis. Lactococcus plantarum
    • Rūšis. Lactococcus raffinolactis
  • Gentis. Lactovum
  • Gentis. Pilibacter
  • Gentis. Streptokokai Streptococcus
    • Rūšis. Streptococcus agalactiae
    • Rūšis. Streptococcus bovis
    • Rūšis. Streptococcus mutans
    • Rūšis. Streptococcus pneumoniae
    • Rūšis. Streptococcus pyogenes
    • Rūšis. Streptococcus salivarius
    • Rūšis. Streptococcus sanguinis
    • Rūšis. Streptococcus suis
    • Rūšis. Streptococcus viridans
    • Rūšis. Streptococcus uberis

Pienskābes baktērijas (Lactobacillales) ir nekustīgu, grampozitīvu, fakultatīvi anaerobu un sporu neveidojošu baktēriju kārta, kas galvenokārt barojas no monosaharīdiem un disaharīdiem (dažas pienskābes baktēriju dzimtas kā barības vielas spēj izmantot arī pentozes vai organiskās skābes, piemēram, citronskābi), kā galveno vielmaiņas galaproduktu izdalot pienskābi, bet mazākos daudzumos arī bioloģiski aktīvas vielas (B vitamīnus) un patogēnu attīstību kavējošas antibiotikas.^[1]

Iedalījums

Pienskābes baktērijas var iedalīt gan pēc to optimālās darbības temperatūras, šūnu formas, gan arī pienskābajā rūgšanā izdalītajiem galaproduktiem. Pēc optimālās darbības temperatūras pienskābes baktērijas iedala:

• mezofilajās pienskābes baktērijās, kuru optimālā darbības temperatūra ir robežās no 20 līdz 30 °C;
• termofilajās pienskābes baktērijās, kuru optimālā darbības temperatūra ir robežās no 40 līdz 45 °C.^[1]

Pēc pienskābās rūgšanas galaproduktiem pienskābes baktērijas iedala:

• homofermentatīvajās pienskābes baktērijās, kas cukurus pārraudzējot veido līdz 90% pienskābes, bet niecīgos daudzumos izdala arī gaistošas skābes un etilspirtu. Atkarībā no to šūnu formas homofermentatīvās pienskābes baktērijas sīkāk iedala:^[1]

• homofermentatīvās lodveida formās, pie kā pieder daļa Lactococcus, Streptococcoccus un Pediococcus ģints baktērijas. Lactococcus sugas baktēriju šūnas ir apaļas vai nedaudz ieapaļas formas 0,5—1 µm diametrā, kas savstarpēji izvietojušās izklaidus, pa pāriem vai ķēdītēs, bet Pediococcus sugas baktēriju šūnas ir izkārtojušās viena no otras izklaidus, pa pāriem vai veido pakešu vai tetraedra veida sakopojumus. Svarīgākie sugu pārstāvji — Lactococcus lactis, Lactococcus cremoris un Streptococcus thermophilus, kas ir nozīmīgi daudzu skābpienu dzērienu ieraugu sastāvdaļa, savukārt Str. thermophilus, kas papildus spēj pārraudzēt arī saharozi, izmanto atsevišķu jogurtu un siera veida ieraugos.^[1]
• homofermentatīvās nūjiņveida formās, pie kā pieder Lactobacillus ģints baktērijas. Baktēriju šūnas var būt dažāda izmēra un garuma nūjiņas, kas izkārtojušās vai nu ķēdītēs vai izklaidus. Svarīgākie sugu pārstāvji — Lactobacillus lactis, Lactobacillus acidophilus, Lactobacillus bulgaricus, Lactobacillus casei un Lactobacillus plantarum, ko izmanto skābpiena dzērienu, skābpiena produktu un siera ieraugos. Lielākā daļa homofermentatīvās nūjiņveida pienskābes baktērijas izdala aptuveni 2—3,5% pienskābes, salīdzinājumā ar homofermentatīvajām lodveida formām, kas izdala vien ap 1% pienskābes un pie pH 3,8—4,0 vairs neattīstās.^[1]

• heterofermentatīvajās pienskābes baktērijās, kas līdztekus pienskābei veido arī aptuveni 50% gaistošo aromātvielu, tostarp, diacetilu, acetoīnu, skudrskābi, dzintarskābi. Arī tās sīkāk iedala pēc baktēriju šūnu formas:

• heterofermentatīvās lodveida formās, pie kā pieskaitāma Leuconostoc ģints un Lactococcus lactis ssp. lactis biovar. diacetylactis baktērijas. Leuconostoc ģints baktērijas spēj pārstrādāt monosaharīdus un disaharīdus. Baktēriju šūnām ir olveida forma un tās vai nu veido blīvus sakopojumus, īsas ķēdītēs vai ir izkārtojušās pa pāriem. Leuconostoc ģints baktērijas lielākoties izmanto skābpiena dzērienu un skābpiena produktu ieraugos..^[1]
• heterofermentīvajās nūjiņveida formās, pie ka pieder bifidobaktērijas un daļa Lactobacillus ģints baktēriju. Bifidobaktērijas ir izliektas formas nūjiņas ar sašķeltiem galiem, kas veido līdz pat aptuveni 95% no jaundzimušo zarnu kopējās mikrofloras. Cilvēkam pieaugot, to skaits pakāpeniski samazinās un pieaugušajiem bifidobaktērijas veido vien ap 25% kopējās zarnu mikrofloras, bet pieaug nevēlamo mikroorganismu skaits, kas var izraisīt dažādas zarnu trakta saslimšanas, tādēļ svarīgi bifidobaktērijas uzņemt ar uzturu.^[1]

Atsauces

Bakterie Lactobacillus na komórkach nabłonka pochwy. Systematyka Królestwo bakterie Typ Firmicutes Klasa Bacilli Rząd Lactobacillales Nazwa systematyczna Lactobacillales

Lactobacillales (bakterie mlekowe) to rząd bakterii z klasy Bacilli.

Charakterystyka

Ich ciała są pałeczkowate lub niemal ziarenkowate. W przypadku form pałeczkowatych widoczny jest podział na człony. W wielu przypadkach z wiekiem rozpadają się one tworząc formy ziarenkowate. Wyniki barwienia Gramma w przypadku tego rzędu są zawsze gramdodatnie bądź gramchwiejne, jednakże pod względem fizjologicznym Lactobacillales zawiera wyłącznie mikroby gramdodatnie. Zaliczają się do niego wyłącznie heterotroficzne bakterie, które nie tworzą endospor. Zwykle występują w glebach, w resztkach roślin bądź jako pasożyty.

Systematyka

Wyróżniamy w nim sześć rodzin: Corynebacteriaceae, Arthobacteraceae, Propionibacteriaceae, Streptococcaceae, Lactobacillaceae oraz Brevibacteriaceae

Rząd pod wieloma względami przypomina Actinomycetales.

Bibliografia

• Systematyka bakterii. W: Władysław J. H. Kunicki-Goldfinger: Życie bakterii. Warszawa: Wydawnictwo Naukowe PWN, 1998, s. 505–6. ISBN 978-83-01-14378-7.

Lactobacillales é uma ordem de bactérias que compõem as bactérias do ácido láctico. Eles são amplamente distribuídos na natureza, e são encontrados no solo, água, plantas e animais. Eles são amplamente utilizados na produção de alimentos fermentados, incluindo os produtos lácteos, como iogurte, queijo, manteiga, manteiga de kefir, e koumiss. LAB também são responsáveis ​​pelo processo de fermentação maloláctica na produção de vinho, e a fermentação do repolho salgado para o chucrute.

Laktobaciller (Lactobacillales) är en ordning av grampositiva bakterier. Familjer som ingår är Lactobacillaceae (mjölksyrabakterier), Enterococcaceae och Streptococcaceae (som omfattar arter som är sjukdomsframkallande).

De finns på många ställen i naturen och återfinns i vatten, jord, växter och djur.

De används vid produktion av mjölkprodukter som yoghurt, ost, smör och fil. Mjölksyrabakterier är också delaktiga i den malolaktiska jäsprocessen vid vinproduktion och vid syrning av surkål.

Lactobacillales, laktik asit bakterilerini kapsayan gram-pozitif bakterilerin bir takımıdır.

Doğada yaygın bir şekilde bulunmaktadır; toprakta, suda, bitkilerde ve hayvanlarda. Yoğurt, peynir, tereyağı, ayran, kefir ve kımız da dahil olmak üzere günlük tüketilen fermente gıdaların üretiminde geniş bir şekilde kullanılmaktadır. Bunlar aynı zamanda şarap üretiminde de malolaktik fermentasyonda ve tuzlanmış lahananın turşuya dönüşmesinde de görev almaktadır.

Lactobacillales — порядок грам-позитивних бактерій типу Firmicutes. Містить більшість молочнокислих бактерій, що розподілені за різними родинами ряду, та велике число бактерій інших родів. Представники ряду дуже поширені в природі, вони мешкають у ґрунті, водоймах, в асоціації з тваринами і рослинами.

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ラクトバシラス目 分類 ドメ
イン : 真正細菌 Bacteria 門 : フィルミクテス門 Firmicutes 綱 : バシラス綱 Bacilli 目 : ラクトバシラス目 Lactobacillales 科

• アエロコッカス科
  Aerococcaceae
• カルノバクテリウム科
  Carnobacteriaceae
• エンテロコッカス科
  Enterococcaceae
• ラクトバシラス科
  Lactobacillaceae
• ロイコノストック科
  Leucinostocaceae
• レンサ球菌科
  Streptococcaceae

ラクトバシラス目(Lactobacillales)とは、乳酸菌を主とした菌を含むフィルミクテス門の細菌の目の一つである。自然界で生成され、土や水、動植物にも存在する。ヨーグルトやチーズ、バター、バターミルクのような乳酸菌を含む醗酵食品にも広く使用されている。

系統樹[編集]

現在受け入れられている分類法は、List of Prokaryotic names with Standing in Nomenclatureの原核生物名リストに基づいており^{[1] [2]}、系統分類は、'The All-Species Living Tree' Projectの16S rRNA系統解析のLTPリリース106に基づいている^[3]。

ラクトバシラス目  

Aerosphaera taetra ♠ Hutson & Collins 2000

 

Carnococcus allantoicus ♠ Tanner et al. 1995

 

Aerococcaceae

 
 

Granulicatella Collins and Lawson 2000

 
 
 

Atopobacter phocae Lawson et al. 2000

 

Bavariicoccus seileri Schmidt et al. 2009

 
 

Trichococcus Scheff et al. 1984 emend. Liu et al. 2002

 
 
 

Lactobacillus algidus Kato et al. 2000

 
 

Lactobacillus species group 2

 
 

Lactobacillus Beijerinck 1901 emend. Cai et al. 2012

 
 

Leuconostocaceae

 
 

Lactobacillus species group 3

 
 

Lactobacillus species group 4

 
 

Lactobacillus species group 5

 
 

Lactobacillus species group 6

 
 

Pediococcus Claussen 1903

 

Lactobacillus species group 7

 
 

Carnobacterium Collins et al. 1987

 
 

Isobaculum melis Collins et al. 2002

 
 

Carnobacteriaceae 2 [incl. various Carnobacterium sp.]

 
 

Desemzia incerta (Steinhaus 1941) Stackebrandt et al. 1999

Enterococcaceae & Streptococcaceae

(続く)

Lactobacillales part 2 (続き)[編集]

Lactobacillales part 2  

Vagococcus fessus Hoyles et al. 2000

 
 

Vagococcus Collins et al. 1990

 
 

Catellicoccus marimammalium Lawson et al. 2006

 
 

Enterococcus (ex Thiercelin and Jouhaud 1903) Schleifer and Kilpper-Bälz 1984

 
 

Enterococcus phoeniculicola Law-Brown and Meyers 2003

 
 

Enterococcus species group 2 [incl. Melissococcus plutonius & Tetragenococcus]

 
 

Enterococcus species group 3

 
 

Enterococcus species group 4

 
 

Enterococcus raffinosus Collins et al. 1989

 

Enterococcus avium (ex Nowlan and Deibel 1967) Collins et al. 1984

 
 

Enterococcus pallens Tyrrell et al. 2002

 
 

Enterococcus hermanniensis Koort et al. 2004

 
 

Pilibacter termitis Higashiguchi et al. 2006

 

Streptococcaceae

脚注[編集]

유산균목(Lactobacillales)은 간균강에 속하는 세균 목이다.^[1] 젖산균을 포함하고 있는 그람-양성균 박테리아로 후벽균문으로 분류한다. 자연에 광범위하게 분포하며, 토양과 물 그리고 식물과 동물에게서 발견된다. 요구르트와 치즈, 버터, 우락유, 케피르 그리고 크므즈 등의 발효 음식을 만드는 데 널리 이용된다. 또한 젖산균은 와인 생산 과정의 젖산 발효(유산 발효), 사워크라우트를 만들기 위해 소금에 절인 양배추를 발효시키는 데에도 쓰인다.

하위 과

• 아이로콕쿠스과 (Aerococcaceae)
• 카르노박테리움과 (Carnobacteriaceae)
• 엔테로콕쿠스과 (Enterococcaceae)
• 유산균과 (Lactobacillaceae)
• 레우코노스톡과 (Leuconostocaceae)
• 연쇄상구균과 또는 스트렙토콕쿠스과 (Streptococcaceae)

각주