Lactic acid bacteria

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Lactic acid bacteria
02-0667 1b.jpg
Lesions of Weissella confusa in the mona monkey (hematoxylin and eosin stain): A) liver: portal triads with neutrophilic infiltration (x10); A1, presence of bacterial emboli inside the vein (arrow) (x40). B) acute pneumonia: edema, congestion, and leukocyte cells exudation in the pulmonary alveoli (x10). C) encephalitis: congestion and marginalized neutrophils in nervous vessels (x10)
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Bacteria
Phylum: Bacillota
Class: Bacilli
Order: Lactobacillales
Ludwig, Schleifer & Whitman 2010
Families
Synonyms
  • "Coccales" Krasilnikov 1949
  • "Plocamobacteriales" Pribram 1933

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).

Contents

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

Streptococcus Streptococci.jpg
Streptococcus

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.[ citation needed ] 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 (LPSN) [6] and National Center for Biotechnology Information (NCBI) [7]

16S rRNA based LTP_08_2023 [8] [9] [10] 120 marker proteins based GTDB 08-RS214 [11] [12] [13]
Lactobacillales

Listeriaceae

Carnobacteriaceae 2

Aerococcaceae

Carnobacteriaceae

Catellicoccus {"Catellicoccaceae"}

Vagococcus {"Vagococcaceae"}

Enterococcus s.s {Enterococcaceae}

Enterococcus phoeniculicola

Enterococcus species-group 4

Enterococcus species-group 3

Enterococcaceae 2

Enterococcus species-group 2

Enterococcus species-group 1

Streptococcaceae

Isobaculum Collins et al. 2002

Lactobacillaceae

Lactobacillales

Listeriaceae

Carnobacteriaceae

Aerococcaceae

Vagococcus {"Vagococcaceae"}

Enterococcaceae

Catellicoccus {"Catellicoccaceae"}

Streptococcaceae

Lactobacillaceae


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] [14]

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 [15] and Alzheimer's disease. [16] Future applications of probiotics have been conjectured to include delivery systems for vaccines and immunoglobulins, and the treatment of different gastrointestinal diseases and vaginosis. [15]

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. [17]

Some LAB produce bacteriocins which limit pathogens by interfering with cell wall synthesis or causing pore formation in the cell membrane. [18] 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. [19] Nisin is utilized as a food additive in at least 50 countries. [19] 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. [20] Furthermore, LAB have the potential to reduce the abundance of mycotoxins in foods by binding to them. [20] 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 . [21]

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. [22]

Fermentation

Lactic acid transformation from malic acid.png

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. [23]

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). [24] 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. [24] 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. [25]

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. [26]

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. [27]

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. [27]

Lactic acid bacteria and dental plaque

LAB are able to synthesize levans from sucrose, and dextrans from glucose. [28] 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. [29] While the primary bacteria responsible for tooth decay is Streptococcus mutans , LAB do feature among the other most common oral bacteria that cause decay. [30]

Lactic acid bacteria genera

See also

Related Research Articles

<span class="mw-page-title-main">Mother of vinegar</span> Biofilm formed on fermenting alcoholic liquids

Mother of vinegar is a biofilm composed of a form of cellulose, yeast, and bacteria that sometimes develops on fermenting alcoholic liquids during the process that turns alcohol into acetic acid with the help of oxygen from the air and acetic acid bacteria (AAB). It is similar to the symbiotic culture of bacteria and yeast (SCOBY) mostly known from production of kombucha, but develops to a much lesser extent due to lesser availability of yeast, which is often no longer present in wine/cider at this stage, and a different population of bacteria. Mother of vinegar is often added to wine, cider, or other alcoholic liquids to produce vinegar at home, although only the bacteria is required, but historically has also been used in large scale production.

<span class="mw-page-title-main">Lactic acid</span> Group of stereoisomers

Lactic acid is an organic acid. It has the molecular formula CH3CH(OH)COOH. It is white in the solid state and it is miscible with water. When in the dissolved state, it forms a colorless solution. Production includes both artificial synthesis as well as natural sources. Lactic acid is an alpha-hydroxy acid (AHA) due to the presence of a hydroxyl group adjacent to the carboxyl group. It is used as a synthetic intermediate in many organic synthesis industries and in various biochemical industries. The conjugate base of lactic acid is called lactate. The name of the derived acyl group is lactoyl.

<i>Lactobacillus</i> Genus of bacteria

Lactobacillus is a genus of gram-positive, aerotolerant anaerobes or microaerophilic, rod-shaped, non-spore-forming bacteria. Until 2020, the genus Lactobacillus comprised over 260 phylogenetically, ecologically, and metabolically diverse species; a taxonomic revision of the genus assigned lactobacilli to 25 genera.

<i>Lactobacillus delbrueckii <span style="font-style:normal;">subsp.</span> bulgaricus</i> Subspecies of bacteria, used in yogurt

Lactobacillus bulgaricus is one of over 200 published species in the Lactobacillus genome complex (LGC) and is the main bacterium used for the production of yogurt. It also plays a crucial role in the ripening of some cheeses, as well as in other processes involving naturally fermented products. It is defined as homofermentive lactic acid bacteria due to lactic acid being the single end product of its carbohydrate digestion. It is also considered a probiotic.

<i>Lactococcus</i> Genus of bacteria

Lactococcus is a genus of lactic acid bacteria that were formerly included in the genus Streptococcus Group N1. They are known as homofermenters meaning that they produce a single product, lactic acid in this case, as the major or only product of glucose fermentation. Their homofermentative character can be altered by adjusting environmental conditions such as pH, glucose concentration, and nutrient limitation. They are gram-positive, catalase-negative, non-motile cocci that are found singly, in pairs, or in chains. The genus contains strains known to grow at or below 7˚C.

<i>Lactococcus lactis</i> Species of bacterium

Lactococcus lactis is a gram-positive bacterium used extensively in the production of buttermilk and cheese, but has also become famous as the first genetically modified organism to be used alive for the treatment of human disease. L. lactis cells are cocci that group in pairs and short chains, and, depending on growth conditions, appear ovoid with a typical length of 0.5 - 1.5 µm. L. lactis does not produce spores (nonsporulating) and are not motile (nonmotile). They have a homofermentative metabolism, meaning they produce lactic acid from sugars. They've also been reported to produce exclusive L-(+)-lactic acid. However, reported D-(−)-lactic acid can be produced when cultured at low pH. The capability to produce lactic acid is one of the reasons why L. lactis is one of the most important microorganisms in the dairy industry. Based on its history in food fermentation, L. lactis has generally recognized as safe (GRAS) status, with few case reports of it being an opportunistic pathogen.

<span class="mw-page-title-main">Probiotic</span> Microorganisms said to provide health benefits when consumed

Probiotics are live microorganisms promoted with claims that they provide health benefits when consumed, generally by improving or restoring the gut microbiota. Probiotics are considered generally safe to consume, but may cause bacteria-host interactions and unwanted side effects in rare cases. There is some evidence that probiotics are beneficial for some conditions, but there is little evidence for many of the health benefits claimed for them.

<span class="mw-page-title-main">Malolactic fermentation</span> Process in winemaking

Malolactic conversion is a process in winemaking in which tart-tasting malic acid, naturally present in grape must, is converted to softer-tasting lactic acid. Malolactic fermentation is most often performed as a secondary fermentation shortly after the end of the primary fermentation, but can sometimes run concurrently with it. The process is standard for most red wine production and common for some white grape varieties such as Chardonnay, where it can impart a "buttery" flavor from diacetyl, a byproduct of the reaction.

<span class="mw-page-title-main">Viili</span> Mesophilic fermented milk product

Viili (Finnish) is a mesophilic fermented milk product found in the Nordic countries, particularly Finland. Viili is similar to yoghurt or kefir, but when left unmixed, its texture is malleable, or "long". The metabolism of the bacteria used in the fermentation also gives viili a slightly different taste.

Pediococcus is a genus of gram-positive lactic acid bacteria, placed within the family of Lactobacillaceae. They usually occur in pairs or tetrads, and divide along two planes of symmetry, as do the other lactic acid cocci genera Aerococcus and Tetragenococcus. They are purely homofermentative.

<i>Lacticaseibacillus casei</i> Species of bacterium

Lacticaseibacillus casei is an organism that belongs to the largest genus in the family Lactobacillaceae, a lactic acid bacteria (LAB), that was previously classified as Lactobacillus casei. This bacteria has been identified as facultatively anaerobic or microaerophilic, acid-tolerant, non-spore-forming bacteria.

<span class="mw-page-title-main">Lactobacillaceae</span> Family of bacteria

The Lactobacillaceae are a family of lactic acid bacteria. It is the only family in the lactic acid bacteria which includes homofermentative and heterofermentative organisms; in the Lactobacillaceae, the pathway used for hexose fermentation is a genus-specific trait. Lactobacillaceae include the homofermentative lactobacilli Lactobacillus, Holzapfelia, Amylolactobacillus, Bombilactobacillus, Companilactobacillus, Lapidilactobacillus, Agrilactobacillus, Schleiferilactobacillus, Loigolactobacillus, Lacticaseibacillus, Latilactobacillus, Dellaglioa, Liquorilactobacillus, Ligilactobacillus, and Lactiplantibacillus; the heterofermentative lactobacilli Furfurilactobacillus, Paucilactobacillus, Limosilactobacillus, Fructilactobacillus, Acetilactobacillus, Apilactobacillus, Levilactobacillus, Secundilactobacillus, and Lentilactobacillus, which were previously classified in the genus Lactobacillus; and the heterofermentative genera Convivina, Fructobacillus, Leuconostoc, Oenococcus, and Weissella which were previously classified in the Leuconostocaceae.

<span class="mw-page-title-main">Food microbiology</span> Study of the microorganisms that inhibit, create, or contaminate food

Food microbiology is the study of the microorganisms that inhabit, create, or contaminate food. This includes the study of microorganisms causing food spoilage; pathogens that may cause disease ; microbes used to produce fermented foods such as cheese, yogurt, bread, beer, and wine; and microbes with other useful roles, such as producing probiotics.

<i>Streptococcus thermophilus</i> Species of bacterium

Streptococcus thermophilus formerly known as Streptococcus salivarius subsp. thermophilus is a gram-positive bacterium, and a fermentative facultative anaerobe, of the viridans group. It tests negative for cytochrome, oxidase, and catalase, and positive for alpha-hemolytic activity. It is non-motile and does not form endospores. S. thermophilus is fimbriated.

<span class="mw-page-title-main">Matzoon</span> Yogurt from the Caucasus

Matzoon or matsoni is a fermented milk product of Armenian origin, distributed in Armenia and Georgia. The so-called Caspian Sea yogurt circulated and commercialized in Japan is sometimes said to be the same type of yogurt as matzoon, but a comparison of microbiota and viscosity found that the two are entirely different.

<span class="mw-page-title-main">Fermented milk products</span>

Fermented milk products or fermented dairy products, also known as cultured dairy foods, cultured dairy products, or cultured milk products, are dairy foods that have been made by fermenting milk with lactic acid bacteria such as Lactobacillus, Lactococcus, and Leuconostoc. The fermentation process increases the shelf life of the product while enhancing its taste and improving the digestibility of its milk. There is evidence that fermented milk products have been produced since around 10,000 BC. A range of different Lactobacilli strains has been grown in laboratories allowing for many cultured milk products with different flavors and characteristics.

<span class="mw-page-title-main">Extracellular polymeric substance</span> Gluey polymers secreted by microorganisms to form biofilms

Extracellular polymeric substances (EPSs) are natural polymers of high molecular weight secreted by microorganisms into their environment. EPSs establish the functional and structural integrity of biofilms, and are considered the fundamental component that determines the physicochemical properties of a biofilm. EPS in the matrix of biofilms provides compositional support and protection of microbial communities from the harsh environments. Components of EPS can be of different classes of polysaccharides, lipids, nucleic acids, proteins, lipopolysaccharides, and minerals.

Pediococcus damnosus is a species of Gram-positive bacteria. The genus Pediococcus is a spherical cocci shaped bacteria with nonmotile, non spore-forming and homofermentative properties. P. damnosus is a chemo-organotrophic, catalase negative, facultative anaerobe. Strains of this species frequently grow in wine and beer, where they overproduce glucan and spoil products by increasing their viscosity. P. damnosus is a lactic acid bacteria (LAB), that can tolerate the low pH and higher ethanol levels that are found in beer. The ability to grow in beer is a strain specific characteristic of the species P. damnosus. Pediococcus damnosus LMG 28219 is a lactic acid bacterium that has proved to be capable of growing in beer.

<span class="mw-page-title-main">Dahi (curd)</span> Fermented milk product

Curd, also mosaru or dahi or Thayir or Perugu, is a traditional yogurt or fermented milk product, originating from and popular throughout the Indian subcontinent. It is usually prepared from cow's milk, and sometimes buffalo milk or goat milk. The word curd is used in Indian English to refer to homemade yogurt, while the term yogurt refers to the pasteurized commercial variety known as heat-treated fermented milk.

Postbiotics - also known as metabiotics, biogenics, or simply metabolites - are soluble factors, secreted by live bacteria, or released after bacterial lysis providing physiological benefits to the host.

References

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  12. ↑ "bac120_r214.sp_label". Genome Taxonomy Database . Retrieved 10 May 2023.
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