Lactobacillus acidophilus

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Lactobacillus acidophilus
20101212 200110 LactobacillusAcidophilus.jpg
Lactobacillus acidophilus, Numbered ticks are 11  μm
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Bacteria
Phylum: Bacillota
Class: Bacilli
Order: Lactobacillales
Family: Lactobacillaceae
Genus: Lactobacillus
Species:
L. acidophilus
Binomial name
Lactobacillus acidophilus
(Moro 1900) Hansen & Mocquot 1970

Lactobacillus acidophilus (Neo-Latin 'acid-loving milk-bacillus') is a rod-shaped, Gram-positive, homofermentative, anaerobic microbe first isolated from infant feces in the year 1900. [1] The species is commonly found in humans, specifically the gastrointestinal tract and oral cavity as well as some speciality fermented foods such as fermented milk or yogurt, though it is not the most common species for this. The species most readily grows at low pH levels (below 5.0), and has an optimum growth temperature of 37 °C. Certain strains of L. acidophilus show strong probiotic effects, and are commercially used in dairy production. The genome of L. acidophilus has been sequenced.

Contents

L. acidophilus has antagonistic effects on the growth of Staphylococcus aureus, Escherichia coli, Salmonella typhimurium, and Clostridium perfringens . [2] Out of the four organisms, Staphylococcus aureus is the most affected. Along with S. aureus, the other Gram-positive bacteria, C. perfringens, was affected more by L. acidophilus, than the two other bacteria that are Gram-negative. L. acidophilus is found to also reduce oral plaque formation by Streptococcus mutans . [3]

History

Lactobacillus acidophilus was first isolated from the human gastrointestinal tract in 1900 by Ernst Moro with the original name Bacillus acidophilus. Over time, there have been many changes to the methods for characterizing taxonomy of organisms, leading to the genus distinction of Lactobacillus in 1929. Complication around finding the original strain arose when multiple strains of a single isolate were given a variety of names. Most studies on L. acidophilus was focused on one particular strain, Lactobacillus acidophilus NCFM. With the large amount of information discovered about L. acidophilus NCFM, the US Food and Drug Administration has adjudged the microbe to be an approved ingredient in beverages, dairy products, and other probiotic foods. [4]

Biological and biochemical features

Morphology

Lactobacillus acidophilus image taken with a scanning electron microscope (SEM). Lactobacillus acidophilus SEM.jpg
Lactobacillus acidophilus image taken with a scanning electron microscope (SEM).
A Lactobacillus acidophilus culture Lactobacillus acidophilus (259 08) Lactobacillus acidophilus (Doderlein bacillus).jpg
A Lactobacillus acidophilus culture

Lactobacillus acidophilus is an immobile rod-shaped (bacillus), gram-positive organism that ranges in size from 2-10 μm in size. L. acidophilus has one phospholipid bilayer membrane with a large cell wall consisting of peptidoglycan exterior to the membrane. The cell wall of L. acidophilus is interwoven with teichoic acids and surface proteins, with anionic and neutral polysaccharides as well as an S-layer lining the exterior of the cell. [5] The S-layer proteins of L. acidophilus have been shown to adhere to epithelial cells as well as mucus and other extracellular proteins. [6] The S-layer is made of two structural domains. The C-terminal domain is responsible for cell wall anchoring, while the N-terminal domain is responsible for interacting with the cell environment, as well as S-layer self assembly. [6] In the L. acidophilus species, the N-terminal region shows high amino acid variability along with low sequence homology (31-72%). However, the C-terminus shows low amino acid variability and high amino acid sequence homology (77-99%). [6] L. acidophilus does not have any extracellular means of motion like a flagellum or pilli, and therefore is an immobile microbe.

Lactobacillus acidophilus under microscope with dark light background. Lactobacillus acidophilus (259 09) Lactobacillus acidophilus (Doderlein bacillus).jpg
Lactobacillus acidophilus under microscope with dark light background.

Metabolism

Pathway by which glucose is converted to lactic acid as a means of energy production Lactic acid fermentation.png
Pathway by which glucose is converted to lactic acid as a means of energy production

L. acidophilus is a homofermentative anaerobic microorganism, meaning it only produces lactic acid as an end product of fermentation; and that it can only ferment hexoses (not pentoses) by way of the EMP pathway (glycolysis). [5] L. acidophilus has a slower growth time in milk than when in a host due to limited available nutrients. Because of its use as a probiotic in milk, a study done by the American Journal of Dairy Science examined the nutrient requirements of L. acidophilus in an effort to increase its low growth rate. The study found that glucose and the amino acids cysteine, glutamic acid, isoleucine, leucine, lysine, methionine, phenylalanine, threonine, tyrosine, valine, and arginine are essential nutrients to the growth of L. acidophilus, with glycine, calcium-pantothenate, and Mn2+ acting as stimulatory nutrients. [7] The study helps to explain the low growth rate of L. acidophilus in milk, as some of the amino acids necessary to L. acidophilus growth are lacking in milk. Adding amino acids with high rates of consumption to fermented milk is a possible solution to the problem. [7]

Genomics

The specialization of prokaryotic genomes is distinguishable when recognizing how the prokaryote replicates its DNA during replication. In L. acidophilus, replication begins at an origin called oriC and moves bi-directionally in the form of replication forks. [5] The DNA is synthesized continuously on the leading strand and in discontinuous Okazaki fragments on the lagging strand with help from the DNA polymerase III enzyme. [8] An RNA primer is needed to initiate the DNA synthesis on the leading and lagging strands. DNA polymerase III follows the RNA primer with the synthesis of DNA in the 5' to 3' direction. [8] L. acidophilus consists of a small genome with a low guanine-cytosine content, approximately 30%. [5] A study comparing 46 genomes of varying strains of L. acidophilus found the genome size ranged from 1.95 Mb to 2.09 Mb, with an average size of 1.98 Mb. [1] The average number of coding sequences in the genome was 1780, with the strains isolated from fermented foods and commercial probiotics having more coding sequences on average than those isolated from humans. [1] L. acidophilus has an open state pan-genome (all of the genes within a species), meaning that the pan-genome size increased as the number of genomes sequenced increased. The core-genome (the genes shared by all individuals of a species) consist of around 1117 genes in the case of L. acidophilus. [1] Genetic analysis also revealed that all L. acidophilus strains contained at least 15 families of glycosyl hydrolases, which are the key enzymes in carbohydrate metabolism. Each of the 15 GH families were involved in metabolizing common carbohydrates, such as glucose, galactose, fructose, sucrose, starch, and maltose. Genes encoding antibiotic resistance by means of antibiotic efflux, antibiotic target alteration, and antibiotic target protection were present in all L. acidophilus strains, providing protection against 18 different classes of antibiotic across all strains. Fluoroquinolone, glycopeptide, lincosamide, macrolide and tetracycline were the five classes of antibiotic to which L. acidophilus displayed the highest level of tolerance, with more than 300 genes relevant to these classes. [1]

Environment

Columnar epithelial cells from a mammal's intestinal tract. L. acidophilus easily adheres to and commonly grows on this cell type Epithelial Tissues Simple Columnar Epithelium (41006485014).jpg
Columnar epithelial cells from a mammal's intestinal tract. L. acidophilus easily adheres to and commonly grows on this cell type

L. acidophilus grows naturally in the oral, intestinal, and vaginal cavities of mammals. [9] Nearly all Lactobacillus species have special mechanisms for heat resistance which involves enhancing the activity of chaperones. Chaperones are highly conserved stress proteins that allow for enhanced resistance to elevated temperatures, ribosome stability, temperature sensing, and control of ribosomal function at high temperatures. [10] This ability to function at high temperatures is extremely important to cell yield during the fermentation process, and genetic testing on L. acidophilus in order to increase its temperature tolerance is currently being done. [1] When being considered as a probiotic, it is important for L. acidophilus to have traits suitable for life in the gastrointestinal tract. Tolerance of low pH and high toxicity levels are often required. These traits vary and are strain specific. Mechanisms by which these tolerances are expressed include differences in cell wall structure, along with other changes is protein expression. [9] Changes in salt concentration have been shown to affect L. acidophilus viability, but only after exposure to higher salt concentrations. In another experiment highlighted by the American Dairy Science Association, viable cell counts only showed a significant reduction after exposure to NaCl concentrations of 7.5% or higher. [11] Cells were also observed to distinctly elongate when grown in conditions of 10% NaCl concentration or higher. [11] L. acidophilus is also very well suited for living in a dairy medium, as fermented milk is the ideal method of delivery for introducing L. acidophilus into a gut microbiome. [7] The viability of L. acidophilus cells encapsulated by spray drying technology stored at refrigerated condition (4 °C) is higher than the viability of cells stored at room temperature (25 °C). [12]

Quorum sensing

Quorum sensing among cells is the process among which cell signaling can lead to coordinated activities which can ultimately help bacteria control gene expression in a consecutive sequence. This is accomplished via detection of small autoinducers which are secreted in response to increasing cell-population density. [13] In Lactobacillus acidophilus, which can be found in the gastrointestinal tract, quorum sensing is important for bacterial interaction when considering biofilm formation and toxin secretion. [14]  In L. acidophilus, along with many other bacteria, the luxS-mediated quorum sensing is involved in the regulation of behavior. In monoculture, the production of luxS increased during the exponential growth phase and started to plateau as it progressed to the stationary phase. Up-regulation of luxS can occur when L. acidophilus is placed in co-cultivation with another Lactobacillus species. [13]

Vaginal microbiota

Lactobacillus acidophilus is relatively rare in the vaginal microbiome; [15] [16] [17] it is more common in the gut. [16] Other species in the genus are more common, including Lactobacillus crispatus , Lactobacillus gasseri , Lactobacillus jensenii , and Lactobacillus iners . [18] [19] [20] [21] In experiments, L. acidophilus seemed to decrease Candida albicans ’ ability to adhere to vaginal epithelial cells; however, L. acidophilus’ use in preventing yeast infections is unclear because this species of Lactobacilli has also been found not to have a very strong ability to adhere to (and thereby colonize) the vaginal cells. [22]

Therapeutic uses

A capsule containing L. acidophilus used for vaginal health Vaginal capsule 2.jpg
A capsule containing L. acidophilus used for vaginal health

Research has shown that the presence of L. acidophilus can produce a variety of probiotic effects in humans, such as acting as a barrier against pathogens, assisting in lactose digestion, enhancing immune response, and reducing cholesterol level. L. acidophilus must exist in concentrations of 10^5 - 10^6 c.f.u (colony-forming units) per mL in order for these effects to be seen. [8] A study conducted at the Wake Forest School of Medicine examined the effects of L. acidophilus on the structure and composition of the gut microbiome of mice with respect to the age of the mice. The research established the importance of the interactions between microbes within a gut microbial environment on the overall health of the organism, and the data showed that mice supplemented with L. acidophilus had reduced proteobacteria levels, and increased levels of other probiotic bacteria when compared to other mice of similar age. [23] Another study conducted at Maranatha Christian University studied the impact of L. acidophilus cell free supernatants (a liquid medium containing the metabolites produced by microbial growth) [24] on the growth pattern of Salmonella typhi, the microbe associated with Typhoid fever. The study showed that the presence of L. acidophilus metabolites significantly inhibited the growth curves displayed by S. typhi, [25] supporting the idea that L. acidophilus presence has a positive impact on the species makeup of a gut microbial community, providing the organism with intestinal health benefits. The innate immune system of L. acidophilus also produces antimicrobial peptides. [26] The group of short peptides found there have shown antimicrobial properties such as their strength against viruses and other cell types, including cancer cells. [27] There is also some evidence supporting the use of a symbiotic gel (containing L. acidophilus) in treating gastrointestinal symptoms in patients who had received a hemodialysis treatment. This gel also reduced the occurrence of vomit, heartburn, and stomachaches. Further study concerning this subject is needed to draw firm conclusions. [28]

Dairy industry usage

An example of fermented milk, a dairy product L. acidophilus is commonly added to for probiotic effects Katugen 002.jpg
An example of fermented milk, a dairy product L. acidophilus is commonly added to for probiotic effects

As stated in a journal from the American Dairy Science Association, "Lactobacillus acidophilus is a commercial strain and probiotic that is widely used in the dairy industry to obtain high-quality fermentation products." [7] Increased levels of beneficial bacteria, and decreased levels of pathogenic bacteria within the intestine due to the consumption of fermented milk containing strains of L. acidophilus has a range of probiotic effects. Reduced serum cholesterol levels, stimulated immune response, and improved lactic acid digestion are all probiotic effects associated with intestinal L. acidophilus presence. L. acidophilus was also effective in reducing Streptococcus mutans levels in saliva, as well as decreasing risk factors associated with the development of nonalcoholic fatty liver disease. [7] The strain of L. acidophilus that has been most widely researched, and is most widely used as a probiotic and is referred to as NCFM. [1]

The most common species of Lactobacillus for use in the production of yoghurt is Lactobacillus delbrueckii subsp. bulgaricus.

Side effects

Although probiotics are generally safe, when they are used by oral administration there is a small risk of passage of viable bacteria from the gastrointestinal tract to the blood stream (bacteremia), which can cause adverse health consequences. [29] Some people, such as those with a compromised immune system, short bowel syndrome, central venous catheters, cardiac valve disease and premature infants, may be at higher risk for adverse events.

See also

Related Research Articles

<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.

<span class="mw-page-title-main">Lactobacillus delbrueckii subsp. bulgaricus</span> Subspecies of bacteria, used in yogurt

Lactobacillus Bulgaricus 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.

<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, such as helping to ease some symptoms of irritable bowel syndrome (IBS). However, many claimed health benefits, such as treating eczema, lack substantial scientific support.

Lactiplantibacillus plantarum is a widespread member of the genus Lactiplantibacillus and commonly found in many fermented food products as well as anaerobic plant matter. L. plantarum was first isolated from saliva. Based on its ability to temporarily persist in plants, the insect intestine and in the intestinal tract of vertebrate animals, it was designated as a nomadic organism. L. plantarum is Gram positive, bacilli shaped bacterium. L. plantarum cells are rods with rounded ends, straight, generally 0.9–1.2 μm wide and 3–8 μm long, occurring singly, in pairs or in short chains. L. plantarum has one of the largest genomes known among the lactic acid bacteria and is a very flexible and versatile species. It is estimated to grow between pH 3.4 and 8.8. Lactiplantibacillus plantarum can grow in the temperature range 12 °C to 40 °C. The viable counts of the "L. plantarum" stored at refrigerated condition (4 °C) remained high, while a considerable reduction in the counts was observed stored at room temperature.

<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.

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

Lacticaseibacillus rhamnosus is a bacterium that originally was considered to be a subspecies of L. casei, but genetic research found it to be a separate species in the L. casei clade, which also includes L. paracasei and L. zeae. It is a short Gram-positive homofermentative facultative anaerobic non-spore-forming rod that often appears in chains. Some strains of L. rhamnosus bacteria are being used as probiotics, and are particularly useful in treating infections of the female urogenital tract, most particularly very difficult to treat cases of bacterial vaginosis. The species Lacticaseibacillus rhamnosus and Limosilactobacillus reuteri are commonly found in the healthy female genito-urinary tract and are helpful to regain control of dysbiotic bacterial overgrowth during an active infection. L. rhamnosus sometimes is used in dairy products such as fermented milk and as non-starter-lactic acid bacterium (NSLAB) in long-ripened cheese. While frequently considered a beneficial organism, L. rhamnosus may not be as beneficial to certain subsets of the population; in rare circumstances, especially those primarily involving weakened immune system or infants, it may cause endocarditis. Despite the rare infections caused by L. rhamnosus, the species is included in the list of bacterial species with qualified presumed safety (QPS) status of the European Food Safety Agency.

<span class="mw-page-title-main">Lactic acid bacteria</span> Order of bacteria

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).

Limosilactobacillus reuteri is a lactic acid bacterium found in a variety of natural environments, including the gastrointestinal tract of humans and other animals. It does not appear to be pathogenic and may have health effects.

<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.

Levilactobacillus brevis is a gram-positive, rod shaped species of lactic acid bacteria which is heterofermentative, creating CO2, lactic acid and acetic acid or ethanol during fermentation. L. brevis is the type species of the genus Levilactobacillus (previously L. brevis group), which comprises 24 species. It can be found in many different environments, such as fermented foods, and as normal microbiota. L. brevis is found in food such as sauerkraut and pickles. It is also one of the most common causes of beer spoilage. Ingestion has been shown to improve human immune function, and it has been patented several times. Normal gut microbiota L. brevis is found in human intestines, vagina, and feces.

Limosilactobacillus fermentum is a Gram-positive species in the heterofermentative genus Limosilactobacillus. It is associated with active dental caries lesions. It is also commonly found in fermenting animal and plant material including sourdough and cocoa fermentation. A few strains are considered probiotic or "friendly" bacteria in animals and at least one strain has been applied to treat urogenital infections in women. Some strains of lactobacilli formerly mistakenly classified as L. fermentum have since been reclassified as Limosilactobacillus reuteri. Commercialized strains of L. fermentum used as probiotics include PCC, ME-3 and CECT5716

Propionibacterium freudenreichii is a gram-positive, non-motile bacterium that plays an important role in the creation of Emmental cheese, and to some extent, Jarlsberg cheese, Leerdammer and Maasdam cheese. Its concentration in Swiss-type cheeses is higher than in any other cheese. Propionibacteria are commonly found in milk and dairy products, though they have also been extracted from soil. P. freudenreichii has a circular chromosome about 2.5 Mb long. When Emmental cheese is being produced, P. freudenreichii ferments lactate to form acetate, propionate, and carbon dioxide

(3 C3H6O3 → 2 C2H5CO2 + C2H3O2 + CO2).
<i>Bifidobacterium</i> Genus of bacteria

Bifidobacterium is a genus of gram-positive, nonmotile, often branched anaerobic bacteria. They are ubiquitous inhabitants of the gastrointestinal tract though strains have been isolated from the vagina and mouth of mammals, including humans. Bifidobacteria are one of the major genera of bacteria that make up the gastrointestinal tract microbiota in mammals. Some bifidobacteria are used as probiotics.

Limosilactobacillus mucosae is a rod shaped species of lactic acid bacteria first isolated from pig intestines. It has mucus-binding activity. The species is an obligate anaerobe, catalase-negative, doesn't form spores and is non-motile. Its type strain is S32T, and has been found to be most closely related to Limosilactobacillus reuteri.

<i>Lactobacillus johnsonii</i> Species of bacterium

Lactobacillus johnsonii is a species in the genus Lactobacillus identified in 1980 by John L. Johnson, an American microbiologist and his associates. Its type strain is ATCC 33200. It is part of the healthy vaginal microbiota and has been identified as having probiotic properties. The L. johnsonii strain La1 was one of the first cultures to be proposed as a probiotic dairy supplement in 1995 at the Nestlé Research Center, Lausanne. Although yeast and bacteria have been used in dairy products for fermenting purposes for centuries, the investigation and choice of a microorganism as a fermenting agent based on its health benefits was novel at the time. Today the probiotic culture is used in the LC1 yogurt products by Nestlé.

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

Lacticaseibacillus paracasei (commonly abbreviated as Lc. paracasei) is a gram-positive, homofermentative species of lactic acid bacteria that are commonly used in dairy product fermentation and as probiotic cultures. Lc. paracasei is a bacterium that operates by commensalism. It is commonly found in many human habitats such as human intestinal tracts and mouths as well as sewages, silages, and previously mentioned dairy products. The name includes morphology, a rod-shaped bacterium with a width of 2.0 to 4.0μm and length of 0.8 to 1.0μm.

Lactobacillus crispatus is a common, rod-shaped species of genus Lactobacillus and is a lactic acid producing bacterial species located in both the vagina, through vaginal discharge, and the vertebrate gastrointestinal tract. This species commonly found in vaginal microbiome and is thought to be beneficial to health.

<i>Lactobacillus bulgaricus</i> GLB44 Subspecies of bacterium

Lactobacillus delbrueckii subsp. bulgaricus is a bacterial subspecies traditionally isolated from European yogurts. Lactobacillus bulgaricusGLB44 differs from other L. bulgaricus strains, because it was isolated from the leaves of Galanthus nivalis in Bulgaria.

Lactobacillus kefiranofaciens is a species of slime-forming, homofermentative, rod-shaped lactic acid bacteria first isolated from kefir grains, hence its name. Its type strain is WT-2B. Its genome has been sequenced. Lactobaccillus kefiranofaciens was first identified in 1967 in Russia through studying kefir granules. Lactobaccillus kefiranofaciens is part of the Lactobacillus genus and Firmicutes phylum of bacteria. These bacterium metabolize carbohydrates and produce lactic acid, which can be useful in fermentation. Two subspecies have been identified as kefirgranum and kefiranofaciens, which share properties such as being gram-positive, facultatively anaerobic, and rod-shaped.L. kefiranofaciens is the subspecies related to kefir granules. Studies have investigated the origins and causes for variation in kefir composition and led to conflicting results. Some studies indicate the presence of L.kefiranofaciens was due to geographical location, while others indicate it was due to the different milks used.

<span class="mw-page-title-main">Human milk microbiome</span> Community of microorganisms in human milk

The human milk microbiota, also known as human milk probiotics (HMP), encompasses the microbiota–the community of microorganisms–present within the human mammary glands and breast milk. Contrary to the traditional belief that human breast milk is sterile, advancements in both microbial culture and culture-independent methods have confirmed that human milk harbors diverse communities of bacteria. These communities are distinct in composition from other microbial populations found within the human body which constitute the human microbiome.

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  22. ↑ Can Yogurt Prevent Yeast Infections? Archived 2012-03-09 at the Wayback Machine . Planned Parenthood Advocates of Arizona. 28 February 2012. Retrieved 28 February 2012.
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  24. ↑ Mani-López E, Arrioja-Bretón D, López-Malo A (January 2022). "The impacts of antimicrobial and antifungal activity of cell-free supernatants from lactic acid bacteria in vitro and foods". Comprehensive Reviews in Food Science and Food Safety. 21 (1): 604–641. doi:10.1111/1541-4337.12872. PMID   34907656. S2CID   245228355.
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  29. ↑ Durchschein F, Petritsch W, Hammer HF (February 2016). "Diet therapy for inflammatory bowel diseases: The established and the new". World Journal of Gastroenterology (Review). 22 (7): 2179–2194. doi: 10.3748/wjg.v22.i7.2179 . PMC   4734995 . PMID   26900283.
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