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''Lactobacillus acidophilus'' (
Neo-Latin Neo-LatinSidwell, Keith ''Classical Latin-Medieval Latin-Neo Latin'' in ; others, throughout. (also known as New Latin and Modern Latin) is the style of written Latin used in original literary, scholarly, and scientific works, first in Italy d ...
'acid-loving milk-bacillus') is a rod-shaped,
Gram-positive In bacteriology, gram-positive bacteria are bacteria that give a positive result in the Gram stain test, which is traditionally used to quickly classify bacteria into two broad categories according to their type of cell wall. The Gram stain is ...
,
homofermentative Fermentation is a type of anaerobic metabolism which harnesses the redox potential of the reactants to make adenosine triphosphate (ATP) and organic end products. Organic molecules, such as glucose or other sugars, are catabolized and reduced ...
,
anaerobic Anaerobic means "living, active, occurring, or existing in the absence of free oxygen", as opposed to aerobic which means "living, active, or occurring only in the presence of oxygen." Anaerobic may also refer to: *Adhesive#Anaerobic, Anaerobic ad ...
microbe first isolated from infant feces in the year 1900. 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 Probiotics are live microorganisms that provide health benefits when consumed, generally by improving or restoring the microbiota in the gut. Probiotics are considered generally safe to consume, but may cause bacteria– host interactions ...
effects, and are commercially used in dairy production. The genome of ''L. acidophilus'' has been sequenced. ''L. acidophilus'' has antagonistic effects on the growth of ''
Staphylococcus aureus ''Staphylococcus aureus'' is a Gram-positive spherically shaped bacterium, a member of the Bacillota, and is a usual member of the microbiota of the body, frequently found in the upper respiratory tract and on the skin. It is often posi ...
,
Escherichia coli ''Escherichia coli'' ( )Wells, J. C. (2000) Longman Pronunciation Dictionary. Harlow ngland Pearson Education Ltd. is a gram-negative, facultative anaerobic, rod-shaped, coliform bacterium of the genus '' Escherichia'' that is commonly fo ...
,
Salmonella typhimurium ''Salmonella enterica'' subsp. ''enterica'' is a subspecies of ''Salmonella enterica'', the rod-shaped, flagellated, aerobic, Gram-negative bacterium. Many of the pathogenic serovars of the ''S. enterica'' species are in this subspecies, includin ...
,'' and ''
Clostridium perfringens ''Clostridium perfringens'' (formerly known as ''C. welchii'', or ''Bacillus welchii'') is a Gram-positive, bacillus (rod-shaped), anaerobic, spore-forming pathogenic bacterium of the genus '' Clostridium''. ''C. perfringens'' is ever-present ...
''. 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 ''Streptococcus mutans'' is a Facultative anaerobic organism, facultatively anaerobic, gram-positive coccus (round bacteria, bacterium) commonly found in the human oral cavity and is a significant contributor to dental caries, tooth decay. The m ...
''.


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 The United States Food and Drug Administration (FDA or US FDA) is a federal agency of the Department of Health and Human Services. The FDA is responsible for protecting and promoting public health through the control and supervision of food ...
has adjudged the microbe to be an approved ingredient in beverages, dairy products, and other probiotic foods.


Biological and biochemical features


Morphology

''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 Peptidoglycan or murein is a unique large macromolecule, a polysaccharide, consisting of sugars and amino acids that forms a mesh-like layer (sacculus) that surrounds the bacterial cytoplasmic membrane. The sugar component consists of alternating ...
exterior to the membrane. The cell wall of ''L. acidophilus'' is interwoven with
teichoic acid Teichoic acids (''cf.'' Greek τεῖχος, ''teīkhos'', "wall", to be specific a fortification wall, as opposed to τοῖχος, ''toīkhos'', a regular wall) are bacterial copolymers of glycerol phosphate or ribitol phosphate and carbohyd ...
s and surface proteins, with anionic and neutral polysaccharides as well as an S-layer lining the exterior of the cell. The S-layer proteins of ''L. acidophilus'' have been shown to adhere to epithelial cells as well as mucus and other extracellular proteins. 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. 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%).''L. acidophilus'' does not have any extracellular means of motion like a flagellum or pilli, and therefore is an immobile microbe.


Metabolism

''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). ''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 Glucose is a sugar with the Chemical formula#Molecular formula, molecular formula , which is often abbreviated as Glc. It is overall the most abundant monosaccharide, a subcategory of carbohydrates. It is mainly made by plants and most algae d ...
and the amino acids
cysteine Cysteine (; symbol Cys or C) is a semiessential proteinogenic amino acid with the chemical formula, formula . The thiol side chain in cysteine enables the formation of Disulfide, disulfide bonds, and often participates in enzymatic reactions as ...
,
glutamic acid Glutamic acid (symbol Glu or E; known as glutamate in its anionic form) is an α- amino acid that is used by almost all living beings in the biosynthesis of proteins. It is a non-essential nutrient for humans, meaning that the human body can ...
,
isoleucine Isoleucine (symbol Ile or I) is an α-amino acid that is used in the biosynthesis of proteins. It contains an α-amino group (which is in the protonated −NH form under biological conditions), an α-carboxylic acid group (which is in the depro ...
,
leucine Leucine (symbol Leu or L) is an essential amino acid that is used in the biosynthesis of proteins. Leucine is an α-amino acid, meaning it contains an α-amino group (which is in the protonated −NH3+ form under biological conditions), an α-Car ...
,
lysine Lysine (symbol Lys or K) is an α-amino acid that is a precursor to many proteins. Lysine contains an α-amino group (which is in the protonated form when the lysine is dissolved in water at physiological pH), an α-carboxylic acid group ( ...
,
methionine Methionine (symbol Met or M) () is an essential amino acid in humans. As the precursor of other non-essential amino acids such as cysteine and taurine, versatile compounds such as SAM-e, and the important antioxidant glutathione, methionine play ...
,
phenylalanine Phenylalanine (symbol Phe or F) is an essential α-amino acid with the chemical formula, formula . It can be viewed as a benzyl group substituent, substituted for the methyl group of alanine, or a phenyl group in place of a terminal hydrogen of ...
,
threonine Threonine (symbol Thr or T) is an amino acid that is used in the biosynthesis of proteins. It contains an α-amino group (which is in the protonated −NH form when dissolved in water), a carboxyl group (which is in the deprotonated −COO− ...
,
tyrosine -Tyrosine or tyrosine (symbol Tyr or Y) or 4-hydroxyphenylalanine is one of the 20 standard amino acids that are used by cells to synthesize proteins. It is a conditionally essential amino acid with a polar side group. The word "tyrosine" is ...
,
valine Valine (symbol Val or V) is an α-amino acid that is used in the biosynthesis of proteins. It contains an α- amino group (which is in the protonated −NH3+ form under biological conditions), an α- carboxylic acid group (which is in the deproton ...
, and
arginine Arginine is the amino acid with the formula (H2N)(HN)CN(H)(CH2)3CH(NH2)CO2H. The molecule features a guanidinium, guanidino group appended to a standard amino acid framework. At physiological pH, the carboxylic acid is deprotonated (−CO2−) a ...
are essential nutrients to the growth of ''L. acidophilus'', with
glycine Glycine (symbol Gly or G; ) is an amino acid that has a single hydrogen atom as its side chain. It is the simplest stable amino acid. Glycine is one of the proteinogenic amino acids. It is encoded by all the codons starting with GG (G ...
, calcium-pantothenate, and Mn2+ acting as stimulatory nutrients. 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.


Genomics

The specialization of
prokaryotic A prokaryote (; less commonly spelled procaryote) is a single-celled organism whose cell lacks a nucleus and other membrane-bound organelles. The word ''prokaryote'' comes from the Ancient Greek (), meaning 'before', and (), meaning 'nut' ...
genomes is distinguishable when recognizing how the prokaryote replicates its
DNA Deoxyribonucleic acid (; DNA) is a polymer composed of two polynucleotide chains that coil around each other to form a double helix. The polymer carries genetic instructions for the development, functioning, growth and reproduction of al ...
during replication. In ''L. acidophilus,'' replication begins at an origin called
oriC Oric may refer to: * Oric (computer), a series of home computers made in the UK in the 1980s * Oric Products International, the parent company that made the Oric computer * oriC, the origin of chromosomal replication in bacteria * ORIC, the Off ...
and moves bi-directionally in the form of replication forks. The DNA is synthesized continuously on the
leading strand In molecular biology, DNA replication is the biological process of producing two identical replicas of DNA from one original DNA molecule. DNA replication occurs in all living organisms, acting as the most essential part of biological inheritance ...
and in discontinuous
Okazaki fragments Okazaki fragments are short sequences of DNA nucleotides (approximately 150 to 200 base pairs long in eukaryotes) which are synthesized discontinuously and later linked together by the enzyme DNA ligase to create the lagging strand during DN ...
on the
lagging strand In molecular biology, DNA replication is the biological process of producing two identical replicas of DNA from one original DNA molecule. DNA replication occurs in all living organisms, acting as the most essential part of biological inheritance ...
with help from the
DNA polymerase III DNA polymerase III holoenzyme is the primary enzyme complex involved in prokaryotic DNA replication. It was discovered by Thomas Kornberg (son of Arthur Kornberg) and Malcolm Gefter in 1970. The complex has high processivity (i.e. the number o ...
enzyme. 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. ''L. acidophilus'' consists of a small
genome A genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding genes, other functional regions of the genome such as ...
with a low guanine-cytosine content, approximately 30%. 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. 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. ''L. acidophilus'' has an open state
pan-genome In the fields of molecular biology and genetics, a pan-genome (pangenome or supragenome) is the entire set of genes from all strains within a clade. More generally, it is the union of all the genomes of a clade. The pan-genome can be broken do ...
(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.'' 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.


Environment

''L. acidophilus'' grows naturally in the oral and intestinal cavities of mammals. 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. This ability to function at high temperatures is extremely important to cell yield during the fermentation process, and
genetic testing Genetic testing, also known as DNA testing, is used to identify changes in DNA sequence or chromosome structure. Genetic testing can also include measuring the results of genetic changes, such as RNA analysis as an output of gene expression, or ...
on ''L. acidophilus'' in order to increase its temperature tolerance is currently being done. 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. 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. Cells were also observed to distinctly elongate when grown in conditions of 10% NaCl concentration or higher. ''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. 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).


Quorum sensing

Quorum sensing In biology, quorum sensing or quorum signaling (QS) is the process of cell-to-cell communication that allows bacteria to detect and respond to cell population density by gene regulation, typically as a means of acclimating to environmental disadv ...
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
autoinducer In biology, an autoinducer is a signaling molecule that enables detection and response to changes in the population density of bacterial cells. Synthesized when a bacterium reproduces, autoinducers pass outside the bacterium and into the surround ...
s which are secreted in response to increasing cell-population density. In ''Lactobacillus acidophilus,'' which can be found in the gastrointestinal tract, quorum sensing is important for bacterial interaction when considering
biofilm A biofilm is a Syntrophy, syntrophic Microbial consortium, community of microorganisms in which cell (biology), cells cell adhesion, stick to each other and often also to a surface. These adherent cells become embedded within a slimy ext ...
formation and toxin secretion. 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 The enzyme S-ribosylhomocysteine lyase (EC 4.4.1.21) catalyzes the reaction :''S''-(5-deoxy-D-ribos-5-yl)-L-homocysteine = L-homocysteine + (4''S'')-4,5-dihydroxypentan-2,3-dione Nomenclature This enzyme belongs to the family of lyases, speci ...
'' 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.


Vaginal microbiota

''Lactobacillus acidophilus'' is relatively rare in the vaginal microbiome; it is more common in the gut. Other species in the genus are more common, including ''
Lactobacillus crispatus ''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 ...
'', '' Lactobacillus gasseri'', '' Lactobacillus jensenii'', and '' Lactobacillus iners''. In experiments, ''L. acidophilus'' seemed to decrease ''
Candida albicans ''Candida albicans'' is an opportunistic pathogenic yeast that is a common member of the human gut flora. It can also survive outside the human body. It is detected in the gastrointestinal tract and mouth in 40–60% of healthy adults. It is usu ...
''’ 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.


Therapeutic uses

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. A study conducted at the
Wake Forest School of Medicine Wake Forest University School of Medicine is the medical school of Wake Forest University, with two campuses located in Winston-Salem, North Carolina and Charlotte, North Carolina, United States. It is affiliated with Atrium Health Wake Forest ...
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. 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) 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'', 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. The group of short peptides found there have shown antimicrobial properties such as their strength against viruses and other cell types, including cancer cells. 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.


Dairy industry usage

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." 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. 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. 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 Bloodstream infections (BSIs) are infections of blood caused by blood-borne pathogens. The detection of microbes in the blood (most commonly accomplished by blood cultures) is always abnormal. A bloodstream infection is different from sepsis, wh ...
), which can cause adverse health consequences. 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

*
Lactic acid fermentation Lactic acid fermentation is a metabolic process by which glucose or other hexose, six-carbon sugars (also, disaccharides of six-carbon sugars, e.g. sucrose or lactose) are converted into cellular energy and the metabolite lactic acid, lactate, w ...


References


External links


''Lactobacillus''
at MedlinePlus
''Lactobacillus acidophilus''
at University of Maryland Medical Center

from the U. of Wisconsin
Global analysis of carbohydrate utilization by ''Lactobacillus acidophilus'' using cDNA microarrays

Safety and protective effect of ''Lactobacillus acidophilus'' and ''Lactobacillus casei'' used as probiotic agent in vivo

Pictures and research on Lactobacillus acidophilus

Type strain of ''Lactobacillus acidophilus'' at Bac''Dive'' – the Bacterial Diversity Metadatabase

''Lactobacillus acidophilus - Benefits & Probiotic Uses''
at Pocket Reviewer {{Authority control Probiotics Lactobacillaceae Bacteria described in 1970 Bacteria used in dairy products Gram-positive bacteria