The
lactic acid bacteria (LAB) comprise a
cladeA clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...
of
Gram-positiveGram-positive bacteria are those that are stained dark blue or violet by Gram staining. This is in contrast to Gram-negative bacteria, which cannot retain the crystal violet stain, instead taking up the counterstain and appearing red or pink...
, low-
GGuanine is one of the four main nucleobases found in the nucleic acids DNA and RNA, the others being adenine, cytosine, and thymine . In DNA, guanine is paired with cytosine. With the formula C5H5N5O, guanine is a derivative of purine, consisting of a fused pyrimidine-imidazole ring system with...
CCytosine is one of the four main bases found in DNA and RNA, along with adenine, guanine, and thymine . It is a pyrimidine derivative, with a heterocyclic aromatic ring and two substituents attached . The nucleoside of cytosine is cytidine...
, acid-tolerant, generally non-sporulating, non-respiring rod or cocci that are associated by their common metabolic and physiological characteristics. These
bacteriaBacteria are a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria have a wide range of shapes, ranging from spheres to rods and spirals...
, usually found in decomposing plants and lactic products, produce
lactic acidLactic acid, also known as milk acid, is a chemical compound that plays a role in various biochemical processes and was first isolated in 1780 by the Swedish chemist Carl Wilhelm Scheele. Lactic acid is a carboxylic acid with the chemical formula C3H6O3...
as the major metabolic end-product of
carbohydrateA carbohydrate is an organic compound with the empirical formula ; that is, consists only of carbon, hydrogen, and oxygen, with a hydrogen:oxygen atom ratio of 2:1 . However, there are exceptions to this. One common example would be deoxyribose, a component of DNA, which has the empirical...
fermentation. This trait has, throughout history, linked LAB with
food fermentationFermentation in food processing typically is the conversion of carbohydrates to alcohols and carbon dioxide or organic acids using yeasts, bacteria, or a combination thereof, under anaerobic conditions. Fermentation in simple terms is the chemical conversion of sugars into ethanol...
s, as acidification inhibits the growth of spoilage agents. Proteinaceous bacteriocins are produced by several LAB strains and provide an additional hurdle for spoilage and
pathogenA pathogen gignomai "I give birth to") or infectious agent — colloquially, a germ — is a microbe or microorganism such as a virus, bacterium, prion, or fungus that causes disease in its animal or plant host...
ic 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 evinced by their
generally recognized as safeGenerally recognized as safe is an American Food and Drug Administration designation that a chemical or substance added to food is considered safe by experts, and so is exempted from the usual Federal Food, Drug, and Cosmetic Act food additive tolerance requirements.-History:On January 1, 1958,...
(GRAS) status, due to their ubiquitous appearance in food and their contribution to the healthy microflora of human mucosal surfaces. The
generaIn biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...
that comprise the LAB are at its core
LactobacillusLactobacillus is a genus of Gram-positive facultative anaerobic or microaerophilic rod-shaped bacteria. They are a major part of the lactic acid bacteria group, named as such because most of its members convert lactose and other sugars to lactic acid. They are common and usually benign...
,
LeuconostocLeuconostoc is a genus of Gram-positive bacteria, placed within the family of Leuconostocaceae. They are generally ovoid cocci often forming chains. Leuconostoc spp. are intrinsically resistant to vancomycin and are catalase-negative . All species within this genus are heterofermentative and are...
,
PediococcusPediococcus 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...
,
LactococcusLactococcus is a genus of lactic acid bacteria that were formerly included in the genus Streptococcus Group N1. They are known as homofermentors meaning that they produce a single product, lactic acid in this case, as the major or only product of glucose fermentation...
, and
StreptococcusStreptococcus is a genus of spherical Gram-positive bacteria belonging to the phylum Firmicutes and the lactic acid bacteria group. Cellular division occurs along a single axis in these bacteria, and thus they grow in chains or pairs, hence the name — from Greek στρεπτος streptos, meaning...
as well as the more peripheral
AerococcusAerococcus is a genus in the phylum Firmicutes .-Etymology:The name Aerococcus derives from:Greek masculine gender noun aer, aeros , air; New Latin masculine gender noun coccus , a berry; New Latin masculine gender noun Aerococcus, air coccus.-Species:The genus contains 7 species, namely* A...
,
CarnobacteriumCarnobacterium is a genus of Gram-positive bacteria within the family Leuconostocaceae. C. divergens and C. maltaromaticum are found in the wild and in food products and can grow anaerobically. These species are not known to be pathogenic in humans but may cause disease in fish....
,
EnterococcusEnterococcus is a genus of lactic acid bacteria of the phylum Firmicutes. Enterococci are Gram-positive cocci that often occur in pairs or short chains, and are difficult to distinguish from streptococci on physical characteristics alone. Two species are common commensal organisms in the...
,
OenococcusOenococcus is a genus of Gram-positive bacteria, placed within the family of Leuconostocaceae. The only species in the genus used to be Oenococcus oeni . In 2006, the species Oenococcus kitaharae was identified...
,
SporolactobacillusSporolactobacillus is a genus of anaerobic endospore-forming Gram-positive motile rod-shaped lactic acid bacteria.Members of this genus are catalase-negative, do not reduce nitrates to nitrites, and do not form indole...
,
Tetragenococcus,
Vagococcus, and
Weisella; these belong to the order
LactobacillalesThe Lactobacillales are an order of Gram-positive bacteria that comprise the lactic acid bacteria. They are widespread in nature, and are found in soil, water, plants and animals. They are widely used in the production of fermented foods, including dairy products such as yogurt, cheese, butter,...
.
Characteristics
The lactic acid bacteria (LAB) are rod-shaped
bacilliBacilli refers to a taxonomic class of bacteria. It includes two orders, Bacillales and Lactobacillales, which contain several well-known pathogens like Bacillus anthracis .-Ambiguity:...
or
coccusCoccus can be used to describe any bacterium that has a spherical shape. It is one of the three distinct types of bacteria shapes, the other two being bacillus and spirillum cells...
. LAB are characterized by an increased tolerance to a lower
pHIn chemistry, pH is a measure of the acidity or basicity of an aqueous solution. Pure water is said to be neutral, with a pH close to 7.0 at . Solutions with a pH less than 7 are said to be acidic and solutions with a pH greater than 7 are basic or alkaline...
range. This aspect partially enables LAB to outcompete other bacteria in a natural
fermentationFermentation is the process of extracting energy from the oxidation of organic compounds, such as carbohydrates, using an endogenous electron acceptor, which is usually an organic compound. In contrast, respiration is where electrons are donated to an exogenous electron acceptor, such as oxygen,...
, as they can withstand the increased acidity from organic acid production (e.g.,
lactic acidLactic acid, also known as milk acid, is a chemical compound that plays a role in various biochemical processes and was first isolated in 1780 by the Swedish chemist Carl Wilhelm Scheele. Lactic acid is a carboxylic acid with the chemical formula C3H6O3...
). Laboratory media used for LAB typically includes a
carbohydrateA carbohydrate is an organic compound with the empirical formula ; that is, consists only of carbon, hydrogen, and oxygen, with a hydrogen:oxygen atom ratio of 2:1 . However, there are exceptions to this. One common example would be deoxyribose, a component of DNA, which has the empirical...
source as most species are incapable of respiration. LAB are
catalaseCatalase is a common enzyme found in nearly all living organisms that are exposed to oxygen, where it catalyzes the decomposition of hydrogen peroxide to water and oxygen...
negative. LAB are amongst the most important groups of microorganisms used in the food industry.
Metabolism
There are two main
hexoseIn organic chemistry, a hexose is a monosaccharide with six carbon atoms, having the chemical formula C6H12O6. Hexoses are classified by functional group, with aldohexoses having an aldehyde at position 1, and ketohexoses having a ketone at position 2....
fermentation
pathwaysIn biochemistry, metabolic pathways are series of chemical reactions occurring within a cell. In each pathway, a principal chemical is modified by a series of chemical reactions. Enzymes catalyze these reactions, and often require dietary minerals, vitamins, and other cofactors in order to function...
that are used to classify LAB genera. Under conditions of excess
glucoseGlucose is a simple sugar and an important carbohydrate in biology. Cells use it as the primary source of energy and a metabolic intermediate...
and limited oxygen, homolactic LAB catabolize one mole of glucose in the
Embden-Meyerhof-ParnasGlycolysis is the metabolic pathway that converts glucose C6H12O6, into pyruvate, CH3COCOO− + H+...
(EMP) pathway to yield two
moleThe mole is a unit of measurement used in chemistry to express amounts of a chemical substance, defined as an amount of a substance that contains as many elementary entities as there are atoms in 12 grams of pure carbon-12 , the isotope of carbon with atomic weight 12. This corresponds to a value...
s of pyruvate. Intracellular
redoxRedox reactions describe all chemical reactions in which atoms have their oxidation state changed....
balance is maintained through the oxidation of NADH, concomitant with pyruvate reduction to lactic acid. This process yields two moles
ATPAdenosine-5'-triphosphate is a multifunctional nucleoside triphosphate used in cells as a coenzyme. It is often called the "molecular unit of currency" of intracellular energy transfer. ATP transports chemical energy within cells for metabolism...
per mole of glucose consumed. Representative homolactic LAB genera include
Lactococcus,
Enterococcus,
Streptococcus,
Pediococcus, and group I lactobacilli.
Heterofermentative LAB use the
pentose phosphate pathwayThe pentose phosphate pathway is a process that generates NADPH and pentoses . There are two distinct phases in the pathway. The first is the oxidative phase, in which NADPH is generated, and the second is the non-oxidative synthesis of 5-carbon sugars...
, alternatively referred to as the pentose phosphoketolase pathway. One mole
Glucose-6-phosphateGlucose 6-phosphate is glucose sugar phosphorylated on carbon 6. This compound is very common in cells as the vast majority of glucose entering a cell will become phosphorylated in this way....
is initially dehydrogenated to 6-phosphogluconate and subsequently decarboxylated to yield one mole of CO
2. 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
ethanolEthanol, also called ethyl alcohol, pure alcohol, grain alcohol, or drinking alcohol, is a volatile, flammable, colorless liquid. It is a psychoactive drug and one of the oldest recreational drugs. Best known as the type of alcohol found in alcoholic beverages, it is also used in thermometers, as a...
via
acetyl-CoAAcetyl coenzyme A or acetyl-CoA is an important molecule in metabolism, used in many biochemical reactions. Its main function is to convey the carbon atoms within the acetyl group to the citric acid cycle to be oxidized for energy production. In chemical structure, acetyl-CoA is the thioester...
and
acetaldehydeAcetaldehyde is an organic chemical compound with the formula CH3CHO or MeCHO. It is one of the most important aldehydes, occurring widely in nature and being produced on a large scale industrially. Acetaldehyde occurs naturally in coffee, bread, and ripe fruit, and is produced by plants as part...
intermediates. In theory, end-products (including ATP) are produced in equimolar quantities from the
catabolismCatabolism is the set of metabolic pathways that break down molecules into smaller units and release energy. In catabolism, large molecules such as polysaccharides, lipids, nucleic acids and proteins are broken down into smaller units such as monosaccharides, fatty acids, nucleotides, and amino...
of one mole of glucose. Obligate heterofermentative LAB include
Leuconostoc,
Oenococcus,
Weissella, and group III lactobacilli.
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. In practice throughout history, streptococci were segregated primarily based on
serologySerology is the scientific study of blood serum and other bodily fluids. In practice, the term usually refers to the diagnostic identification of antibodies in the serum...
, which has proven to correlate well with the current taxonomic definitions. Lactococci (formerly Lancefield group N streptococci) are used extensively as starter innocula in
dairyA dairy is a business enterprise established for the harvesting of animal milk—mostly from cows or goats, but also from buffalo, sheep, horses or camels —for human consumption. A dairy is typically located on a dedicated dairy farm or section of a multi-purpose farm that is concerned...
fermentations, with humans estimated to consume 10
18 lactococci annually. Partly due to their industrial relevance, both
Lactococcus lactis subspecies (
lactis and
cremoris) are widely used as generic LAB models for research.
L. lactis ssp.
cremoris, used in the production of hard
cheeseCheese is a generic term for a diverse group of milk-based food products. Cheese is produced throughout the world in wide-ranging flavors, textures, and forms....
s, 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
genomeIn modern molecular biology and genetics, the genome is the entirety of an organism's hereditary information. It is encoded either in DNA or, for many types of virus, in RNA. The genome includes both the genes and the non-coding sequences of the DNA/RNA....
of IL1403, which coincided with a significant shift of resources to understanding LAB
genomicsGenomics is a discipline in genetics concerning the study of the genomes of organisms. The field includes intensive efforts to determine the entire DNA sequence of organisms and fine-scale genetic mapping efforts. The field also includes studies of intragenomic phenomena such as heterosis,...
and related applications.
Bacteriophages
A broad number of food products, commodity chemicals, and
biotechnologyBiotechnology is a field of applied biology that involves the use of living organisms and bioprocesses in engineering, technology, medicine and other fields requiring bioproducts. Biotechnology also utilizes these products for manufacturing purpose...
products are manufactured industrially by large-scale bacterial fermentation of various organic substrates. Because enormous amounts of bacteria are being cultivated each day in large fermentation vats, the risk that
bacteriophageA bacteriophage is any one of a number of viruses that infect bacteria. They do this by injecting genetic material, which they carry enclosed in an outer protein capsid...
contamination rapidly brings fermentations to a halt and cause economical setbacks is a serious threat in these industries. The relationship between bacteriophages and their bacterial hosts is very important in the context of the food fermentation industry. Sources of phage contamination, measures to control their propagation and dissemination, and biotechnological defence strategies developed to restrain phages are of interest. The dairy fermentation industry has openly acknowledged the problem of phage and has been working with academia and starter culture companies to develop defence strategies and systems to curtail the propagation and evolution of phages for decades.
Bacteriophage–host interaction
The first contact between an infecting phage and its bacterial host is the attachment of the phage 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 protein, host determinant, and anti-receptor. For simplicity, the RBP term will be used here. A variety of molecules have been suggested to act as host receptors for
bacteriophageA bacteriophage is any one of a number of viruses that infect bacteria. They do this by injecting genetic material, which they carry enclosed in an outer protein capsid...
s infecting LAB; among those are
polysaccharidePolysaccharides are long carbohydrate molecules, of repeated monomer units joined together by glycosidic bonds. They range in structure from linear to highly branched. Polysaccharides are often quite heterogeneous, containing slight modifications of the repeating unit. Depending on the structure,...
s, (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 range. These studies, however, also found additional phage proteins to be important for successful a phage infection. Analysis of the crystal structure of several RBPs indicated that these proteins share a common tertiary folding as well as supporting previous indications of the saccharide nature of the host receptor. The
Gram-positiveGram-positive bacteria are those that are stained dark blue or violet by Gram staining. This is in contrast to Gram-negative bacteria, which cannot retain the crystal violet stain, instead taking up the counterstain and appearing red or pink...
LAB have a thick
peptidoglycanPeptidoglycan, also known as murein, is a polymer consisting of sugars and amino acids that forms a mesh-like layer outside the plasma membrane of bacteria , forming the cell wall. The sugar component consists of alternating residues of β- linked N-acetylglucosamine and N-acetylmuramic acid...
layer, which must be traversed in order to inject the phage
genomeIn modern molecular biology and genetics, the genome is the entirety of an organism's hereditary information. It is encoded either in DNA or, for many types of virus, in RNA. The genome includes both the genes and the non-coding sequences of the DNA/RNA....
into the bacterial
cytoplasmThe cytoplasm is a small gel-like substance residing between the cell membrane holding all the cell's internal sub-structures , except for the nucleus. All the contents of the cells of prokaryote organisms are contained within the 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.
Probiotics
Probiotics are products aimed at delivering living, potentially beneficial, bacterial cells to the gut
ecosystemAn ecosystem is a biological environment consisting of all the organisms living in a particular area, as well as all the nonliving , physical components of the environment with which the organisms interact, such as air, soil, water and sunlight....
of humans and other animals, whereas prebiotics are non-digestible
carbohydrateA carbohydrate is an organic compound with the empirical formula ; that is, consists only of carbon, hydrogen, and oxygen, with a hydrogen:oxygen atom ratio of 2:1 . However, there are exceptions to this. One common example would be deoxyribose, a component of DNA, which has the empirical...
s delivered in food to the large bowel to provide fermentable substrates for selected bacteria. Strains of LAB are the most common microbes employed as probiotics. Two principal kinds of probiotic bacteria, members of the genera
LactobacillusLactobacillus is a genus of Gram-positive facultative anaerobic or microaerophilic rod-shaped bacteria. They are a major part of the lactic acid bacteria group, named as such because most of its members convert lactose and other sugars to lactic acid. They are common and usually benign...
and
BifidobacteriumBifidobacterium is a genus of Gram-positive, non-motile, often branched anaerobic bacteria. They are ubiquitous, endosymbiotic inhabitants of the gastrointestinal tract, vagina and mouth of mammals and other animals. Bifidobacteria are one of the major genera of bacteria that make up the colon...
, have been studied in detail.
Most probiotic strains belong to the genus
LactobacillusLactobacillus is a genus of Gram-positive facultative anaerobic or microaerophilic rod-shaped bacteria. They are a major part of the lactic acid bacteria group, named as such because most of its members convert lactose and other sugars to lactic acid. They are common and usually benign...
. Probiotics have been evaluated in research studies in animals and humans with respect to antibiotic-associated diarrhoea, travellers' diarrhoea, pediatric diarrhoea,
inflammatory bowel diseaseIn medicine, inflammatory bowel disease is a group of inflammatory conditions of the colon and small intestine. The major types of IBD are Crohn's disease and ulcerative colitis.-Classification:...
, and
irritable bowel syndromeIrritable bowel syndrome is a diagnosis of exclusion. It is a functional bowel disorder characterized by chronic abdominal pain, discomfort, bloating, and alteration of bowel habits in the absence of any detectable organic cause. In some cases, the symptoms are relieved by bowel movements...
. It is possible that, in the future, probiotics will be used for different gastrointestinal diseases, vaginosis, or as delivery systems for vaccines, immunoglobulins, and other therapies.
Exopolysaccharides
The quest to find food ingredients with valuable bioactive properties has encouraged interest in
exopolysaccharideExopolysaccharides are high-molecular-weight polymers that are composed of sugar residues and are secreted by a microorganism into the surrounding environment. Microorganisms synthesize a wide spectrum of multifunctional polysaccharides including intracellular polysaccharides, structural...
s from LAB.
Functional foodFunctional food is a food where a new ingredient has been added to a food and the new product has a new function ....
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.
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