|Beijerinck 1901 emend. Haakensen et al. 2009|
Lactobacillus is a genus of gram-positive , mostly rod-shaped bacteria fromthe Lactobacillaceae family . The word is also written " Germanized " Laktobazillus ( plural : Laktobazillen). Lactobacillus belongs to the lactic acid bacteria together with other types of bacteria ; they all produce lactic acid by fermentation .
Lactobacillus - types are important for the food industry. They are used for the production of dairy products and beer specialties such as Berliner Weisse and Leipziger Gose . As opportunistic pathogens, with the exception of immunosuppressed patients, they usually do no harm to people themselves; they are non-pathogenic .
With regard to their characteristics , the representatives of the genus Lactobacillus do not form a uniform group, therefore individual species are combined into subgroups. Within the system, it can be observed that individual species are newly added or no longer belong to the genus. This led to an expanded description of the genus in 2009.
Origin of the designation
Lactobacillus is a typical genus for a lactic acid bacterium. The representatives of the genus are gram-positive bacteria, they do not form persistent forms like endospores and in most cases are not capable of active movement. The types of Lactobacillus are predominantly rod-shaped , the cells occur individually or in chains. However, curved and helical (e.g. Lactobacillus curvatus ) variants can also occur. After the assignment of different species, which were initially classified in other genera, there are now also some cocci and bacterial forms between cocci and rods among the lactobacilli.
On solid, carbohydrate-containing nutrient media , the cells grow into colonies , which in Lactobacilli are typically quite small. In L. paralimentarius, for example , the diameter of the colonies is between 0.8 and 1.5 mm.
Growth and metabolism
As representatives of the lactic acid bacteria, lactobacilli grow anaerobically but aerotolerantly, i. that is, they grow in the presence of atmospheric oxygen, but do not need oxygen for their metabolism . They are catalase negative and oxidase negative. However, they are able to form cytochromes when cultured on nutrient media containing heme or blood . In this case they show a positive reaction in the oxidase test. In the case of Lactobacillus , however, there are also some species that absolutely do not tolerate oxygen. L. aviarius and L. ruminis are examples of these species, which are designated as obligately anaerobic . Another characteristic of lactobacilli is the need for complex growth factors and amino acids in cultivation.
The temperatures suitable for cultivation are in the range of 30–40 ° C for most species, so Lactobacillus is one of the mesophilic organisms. Some Lactobacillus species also grow well at 45 ° C. This thermophilic tendency is used to subdivide the genus and is important for use in the food industry. The optimal pH value for growth is a slightly acidic pH value (pH 5 to 6), with acidic pH values up to pH 4 also being tolerated. The lactic acid produced lowers the pH of the nutrient medium, provided it does not contain any buffering additives. Some Lactobacillus species produce up to 2.3% lactic acid in nutrient media rich in carbohydrates.
Lactic acid fermentation
Lactobacilli can use various carbohydrates for energy production in one fermentation . A characteristic of fermentation (fermentation) is that the substrates are broken down without oxygen. The typical fermentation product for lactic acid bacteria is lactic acid , which is why this metabolic pathway is called lactic acid fermentation . A distinction is made between homofermentative and heterofermentative types. Homofermentative types produce almost exclusively lactic acid from glucose through fermentation, while heterofermentative types produce not only lactic acid, but also other end products to a significant extent, mostly ethanol and carbon dioxide , sometimes also acetic acid . The heterofermentatives usually lack the enzyme aldolase .
The representatives of the genus Lactobacillus do not form a uniform group in this regard. Lactobacillus delbrueckii subsp. Belongs to the homofermentative species . bulgaricus , the typical metabolic pathway is explained there. The following table gives an overview of the lactic acid fermentation of some Lactobacillus species.
|Homofermentative species or subspecies||Heterofermentative species or subspecies|
|Lactobacillus acidophilus||Lactobacillus bifermentans|
|Lactobacillus alimentarius||Lactobacillus brevis|
|Lactobacillus casei||Lactobacillus fermentum|
|Lactobacillus delbrueckii subsp. bulgaricus||Lactobacillus oryzae|
|Lactobacillus delbrueckii subsp. delbrueckii||Lactobacillus panis|
|Lactobacillus delbrueckii subsp. lactis||Lactobacillus paracasei|
|Lactobacillus helveticus||Lactobacillus parabuchneri|
|Lactobacillus plantarum||Lactobacillus paralimentarius (formerly Lactobacillus kimchii )|
|Lactobacillus salivarius||Lactobacillus rhamnosus|
Many lactobacilli produce bacteriocins , poisonous proteins or peptides that are secreted by bacteria and kill other (competing) types of bacteria or hinder their growth. Bacteriocins produced by various Lactobacillus species include: a. Lactacin-F and Bavaricin-A. Lactobacillus plantarum forms various plantaricins (A, S, T, and Plantaricin-SIK).
Lactobacillus is one of the gram-positive bacteria with a low GC content (the proportion of the nucleobases guanine and cytosine ) in the bacterial DNA . Lactobacillus species also show a high degree of variability with regard to this characteristic; the GC content is between 32 and 53 mol percent . Lactobacillus mali, for example, has a GC content of 32-34 mol percent, while a GC content of 53-56 mol percent has been determined for Lactobacillus panis . Usually the content of guanine and cytosine in the DNA does not vary so strongly within a bacterial genus, which confirms that the lactobacilli form a rather heterogeneous group.
In addition to milk and dairy products , the species of Lactobacillus occur in or on plants; some form part of the natural intestinal flora of humans and other animals. Lactobacilli have been isolated from all parts of the digestive tract of humans, including the stomach.
The species Bifidobacterium bifidum , which used to belong to the lactobacilli, occurs in the intestines of adults and (breast-fed) infants. The obligatory anaerobic bacterium is an important part of the intestinal flora. The term bifidus flora generally stands for the entirety of the different bifidobacteria species in the human intestine. Specifically, the intestinal flora of babies breastfed with breast milk is referred to here.
Lactobacillus salivarius and Lactobacillus ruminis belong to the autochthonous intestinal flora of humans. Autochthonous bacteria are regularly detectable within the respective habitat. L. salivarius and L. ruminis are therefore "permanent residents" of the human intestine. The species Lactobacillus paracasei , L. brevis , L. fermentum , L. plantarum and L. rhamnosus only occur temporarilyin the intestine.
In the stomachs of various animals such as mice, pigs and rats, lactobacilli form layers of cells that are connected to the epithelial cells of the stomach. Lactobacillus amylovorus , L. johnsonii and L. reuteri are the most common in the digestive tract of pigs . Furthermore, some lactobacilli form dense layers on the epithelium in the crop of birds, especially the species L. salivarius .
Lactobacillus reuteri forms antibiotic substances and is used commercially as a probiotic in poultry farming, for example to prevent salmonella infections. The intestinal bacterium from the animal world, for which the Swedish company BioGaia has applied for a patent, isalso used in human medicine, where it is used as an agent for dental health, as an anti-diarrheal agent for small children and as a stomach agent to combat Helicobacter pylori . The bacterium, which is used in factory farming to save synthetic antibiotics, spreads very quickly in the human digestive tract after ingestion and can stay there not only in the intestine, but also in the acidic environment of the stomach. It can be transmitted through breast milk, i.e. it infects babies and can be found in raw milk products. Since 2018, after a study on genetically modified mice, L. reuteri has been suspected of being able to migrate from the digestive tract to the liver. There it is said totriggerthe autoimmune disease systemic lupus erythematosus . Migration to other organs is normally an exclusion criterion for the use of probiotics. The immunobiologist Martin Kriegel, who uncovered the scandal, found L. reuteri not only in the liver of mice, which developed systemic lupus as a result , but also in the liver of lupus patients. The researcher recommends those affectedto include resistant starch in their diet in orderto displace L. reuteri with less dangerous types of bacteria.
Different types of Lactobacillus form the so-called Döderlein bacteria or Döderlein rods. The Döderlein bacteria are part of the natural vaginal flora in women. Fermentation causes the bacteria in the vagina to create an acidic environment, protecting the vagina from other disease-causing bacteria that cannot tolerate low pH. The species most frequently identified in various studies include Lactobacillus crispatus , L. iners , L. gasseri, and L. jensenii . Earlier it was Lactobacillus acidophilus determined as dominant species in the vaginal flora of healthy women.
Furthermore, lactobacilli can be isolated from many plant surfaces, this being the case with intact, but also with decomposing plant parts. For example, L. brevis , L. delbrueckii , L. fermentum and L. plantarum have been detected there.
Lactobacillus is a type genus of the family Lactobacillaceae and the order Lactobacillales. The family currently (2013) comprises three genera: Lactobacillus , Pediococcus and Sharpea . Genetic and cell morphological studies by Haakensen u. a. on the species initially classified as Pediococcus dextrinicus showed that it is better assigned to the Lactobacilli , in addition to the renaming of the species to Lactobacillus dextrinicus , this also led to an expanded description of the genus Lactobacillus in 2009 .
The genus Lactobacillus includes numerous (around 80) species. Currently (2013) 218 Lactobacillus species and subspecies are listed by the Leibniz Institute DSMZ - German Collection of Microorganisms and Cell Cultures GmbH in the Prokaryotic Nomenclature up-to-date (“Prokaryotic nomenclature up to date”). This compilation includes all names validly published according to the Bacteriological Code and takes into account the validation list of the International Journal of Systematic and Evolutionary Microbiology . In the case of other species, the assignment to other genera is suggested.
Because of the diversity within the genus, it is to be expected that reclassification will result in new species being added or no longer belonging to the genus, and that new Lactobacillus species will be described anew . In order to structure the rather heterogeneous group of lactobacilli, a division into three subgroups has proven successful:
Lactobacillus casei / Pediococcus group
Lactobacillus delbrueckii group
Lactobacillus brevis group
|Lactic acid fermentation||homofermentative||homofermentative||heterofermentative|
|Lactic fermentation products||
Lactic acid is the main product,
no gas formation from glucose
|Lactic acid is the main product,
no gas formation from glucose
|About 50% of lactic acid is formed from glucose,
and carbon dioxide (CO 2 ) and ethanol are also produced
|Influence of temperature on growth||Growth at 15 ° C, growth at 45 ° C variable||Growth at 45 ° C but not at 15 ° C||variable|
|Typical species||L. casei, L. aviarius, L. curvatus, L. mali, L. plantarum, L. ruminis, L. salivarius||L. delbrueckii, L. acidophilus, L. amylolyticus, L. crispatus, L. helveticus||L. brevis, L. buchneri, L. fermentum, L. kefiri, L. paracasei, L. rhamnosus|
A selection of the types of Lactobacillus :
Beijerinck emend in 1901. Haakensen et al. 2009
Lactobacillus delbrueckii ( Leichmann 1896) Beijerinck 1901 ( type species )
- Lactobacillus delbrueckii subsp. bulgaricus ( Orla-Jensen 1919) Weiss et al. 1984
- Lactobacillus delbrueckii subsp. delbrueckii ( Leichmann 1896) Weiss et al. 1984
- Lactobacillus delbrueckii subsp. lactis ( Orla-Jensen 1919) Weiss et al. 1984
- Lactobacillus acidophilus ( Moro 1900) Hansen et Mocquot 1970
- Lactobacillus alimentarius (ex Reuter 1970) Reuter 1983
- Lactobacillus amylolyticus Bohak et al. 1999
- Lactobacillus amylovorus Nakamura 1981
- Lactobacillus aviarius Fujisawa et al. 1985
- Lactobacillus bifermentans (ex Pette et van Beynum 1943) Kandler et al. 1983
- Lactobacillus brevis ( Orla-Jensen 1919) Bergey et al. 1934
- Lactobacillus buchneri ( Henneberg 1903) Bergey et al. 1923
- Lactobacillus casei ( Orla-Jensen 1916) Hansen et Lessel 1971
- Lactobacillus coryniformis Abo-Elnaga et Kandler 1965
- Lactobacillus crispatus ( Brygoo et Aladame 1953) Moore et Holdeman 1970 emend. Cato et al. 1983
- Lactobacillus crustorum Scheirlinck et al. 2007
- Lactobacillus curvatus ( Troili-Petersson 1903) Abo-Elnaga et Kandler 1965 emend. Klein et al. 1996
- Lactobacillus dextrinicus ( Coster et White 1964) Haakensen et al. 2009
- Lactobacillus fermentum Beijerinck 1901 emend. Dellaglio et al. 2004
- Lactobacillus gasseri Lauer et Kandler 1980
- Lactobacillus helveticus ( Orla-Jensen 1919) Bergey et al. 1925
- Lactobacillus hilgardii Douglas et Cruess 1936
- Lactobacillus iners Falsen et al. 1999
- Lactobacillus jensenii Gasser et al. 1970
- Lactobacillus johnsonii Fujisawa et al. 1992
- Lactobacillus kefiri corrig. Kandler et Kunath 1983,
- Lactobacillus lindneri (ex Henneberg 1901) Back et al. 1997
- Lactobacillus mali Carr et Davies 1970
- Lactobacillus oryzae Tohno et al. 2013
- Lactobacillus panis Wiese et al. 1996,
- Lactobacillus parabuchneri Farrow et al. 1989,
- Lactobacillus paracasei Collins et al. 1989
- Lactobacillus paralimentarius Cai et al. 1999
- Lactobacillus plantarum ( Orla-Jensen 1919) Bergey et al. 1923
- Lactobacillus reuteri Kandler et al. 1982
- Lactobacillus rhamnosus ( Hansen 1968) Collins et al. 1989
- Lactobacillus rossiae corrig. Corsetti et al. 2005
- Lactobacillus ruminis Sharpe et al. 1973
- Lactobacillus sakei corrig. Katagiri et al. 1934 emend. Klein et al. 1996
- Lactobacillus salivarius Rogosa et al. 1953 emend. Li et al. 2006
- Lactobacillus sanfranciscensis corrig. (ex Kline et Sugihara 1971) Weiss et Schillinger 1984,
- Lactobacillus selangorensis ( Leisner et al. 2000) Haakensen et al. 2011
- Lactobacillus delbrueckii ( Leichmann 1896) Beijerinck 1901 ( type species )
Lactic acid fermentation is mainly used in the food industry in the manufacture of dairy products such as cheese and yoghurt . Without lactic acid bacteria there would be practically no dairy products. But they are also involved in the production of other acidified food and feed. In addition to the Lactobacillus species, other types of bacteria are usually also involved.
Types of Lactobacillus and Pediococcus are also known to be pests in beverage production. The formation of lactic acid and other products leads to undesirable acidification and changes in taste, e.g. B. in beer , wine and fruit juices . Even with heated drinking milk ( long-life milk , pasteurized milk ), the lactobacilli are undesirable if they get back into the product through contamination after heating.
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