Metchnikoff concluded “as lactic fermentation serves so well to arrest putrefaction in general, why should it not be used for the same purpose within the digestive tube?” Taken as the proof of its efficacy, milk fermented with the “Bulgarian bacillus” of Metchnikoff subsequently enjoyed considerable popularity in western Europe (94). The reason for this special foregut association is likely due to the adherence of lactobacilli to the surface of the nonsecretory epithelium lining of these sites, which enables the bacteria to form a biofilm-like structure that provides a bacterial inoculum of the digesta (92). h��Y[O�H�+�تb=����H�Y�B -U-?��K� J����=�Lb'$V�B��s��9�;c�Ќ3!��%q9&9={&-fx`Br�\ Intestinal ecosystems are no exception, and although they are dynamic, they remain remarkably resistant and resilient to chaotic blooms of subpopulations and pathogens (48). Some brands of this medicine (lactobacillus tablets and capsules) need to be stored at room temperature. Lactobacillus can do in a week what takes Pediococcus a year! Specifically, there has been a general and persistent assumption that a large number of Lactobacillus species form stable and numerically significant populations in the human intestinal tract, especially in the small intestine, where they are presumed to form epithelial associations (101). Bacteria belonging to the genus Lactobacillus are members of the lactic acid bacteria (LAB), a broadly defined group characterized by the formation of lactic acid as the sole or main end product of carbohydrate metabolism. As a consequence, gut inhabitants that share long evolutionary histories with their host species are likely to possess adaptive health attributes that can be explored when these organisms are used as probiotics. As noted above, most Lactobacillus species found in the human intestinal tract do not appear to be true inhabitants, and it remains unclear how autochthonous species satisfy their fastidious nutritional requirements in regions distal to host nutrient absorption. Gut bacteria can enhance host immune functions and the mucosal barrier, and they provide protection against incoming microbes (97). They can be found in plants or material of plant origin, silage, fermented food (yogurt, cheese, olives, pickles, salami, etc. Lactobacilli: Preparation of fermented food (yoghurt, sauerkraut, pickles). Products Containing Lactobacillus Acidophilus. How lactobacilli facilitate rapid growth in the human intestinal tract remains dubious, as they are fastidious organisms with nutritional requirements one would consider disadvantageous in regions distal to host nutrient absorption. The bacteria that occupy a niche in the GIT are true residents or autochthonous (i.e., found where they are formed) components, as defined by Savage more than 30 years ago (80). Comparative genomic investigations to identify colonization determinants require exact knowledge about the origin of strains in order to link genome features to ecological function. Autochthonous strains have a long-term association with a particular host, and they form stable populations of a characteristic size in a particular region of the gut (80). Although Metchnikoff's theories focused on LAB that were introduced into the digestive tract through the consumption of fermented food, he argued that each bacterium was “able to take its place in the intestinal flora of man” (59). I acknowledge various inspiring discussions with Gerald Tannock (University of Otago, Dunedin, New Zealand). Around 5% of the sequences obtained from the colons of non-IBD patients accounted for lactobacilli (Table 1). First, Bulgarian peasants, assumed to have a long life expectancy, consumed large amounts of fermented milk products (97). These significant numbers certainly do imply that L. reuteri does inhabit all these different sites. In monogastric animals, the largest numbers of bacteria reside in the distal gut (colon), reaching densities of around 1011 microbes per gram of luminal contents (90). One could also assume that the different anatomical and physiological conditions present throughout the gut would account for distinct bacterial traits to be required for colonization. Accordingly, strains of L. reuteri are especially successful in the prevention of colitis in several animal models (54, 67). However, these species still lack pathways for the synthesis of most amino acids, nucleotides, and vitamins. Like lactobacilli in conventional animals, Lactobacillus reuteri colonizes Lactobacillus-free mice throughout the gut and stably maintains cell numbers of around 109 cells per gram in the forestomach, around 107 cells per gram in the jejunum, and around 108 cells per gram in the cecum (102, 103, 105). 1A to D). The results from feeding studies of lactobacilli indicate that the survival of lactobacilli that originate from food during gastrointestinal passage is comparable to that of probiotic strains. �q )YP��X����H�g�� ll�p�$ѯs���h�(Τ�F �AJ2���RL:���L�>�aJQgu��c��V�i��LIk�t��E-�x�%3�S�b��=���xGE�Y��������c֢N�����Y5`xK�.���:�>��ͪ��4�5�Uv�pWe��O�uYW�u�M�b���w��J�{�W��]��l:����͛ޠ����mvQ�g���Mg0�Y����ww��Ō�9ͮ�!8�����ni5����c5��U3�d֭R��mv*"�p꬈����p��� N�oS�pTa>B���*���{޿x7�yp8 ba��V����jQ�U���ߌ*ړ���F3HkEui���]=�f��o���/N[OU6MK�0C��H��$�����=-��g�+&8��.Ҍ����4��43�8��*W����.7J��\]�xvL_H/��� W�6W�Jۘ��R���\{�ԥg�S���|���1��. Dal Bello and Hertel showed that several fecal and oral isolates from three subjects isolated at the same time point were of the same randomly amplified polymorphic DNA type, suggesting that these fecal isolates originated from the oral cavity (16). It is somehow intriguing how lactobacilli could maintain a reputation as numerically important intestinal inhabitants, given that the vast majority of experimental studies conducted after 1960 clearly showed that they form marginal populations in the human gut. %PDF-1.7 %���� Most strains were detected only in one or two fecal samples from the majority of subjects and then went missing. Journal of Microbiology & Biology Education, Microbiology and Molecular Biology Reviews, Human intestinal microbiota in health and disease, Bergey's manual of systematic bacteriology. acidophilus NCFM ( Altermann, 2005) to 3 308 274 bp for Lb. Comparisons of the characteristics of germ-free animals and those of conventional animals have clearly demonstrated that the gut microbiota has considerable influence on host biochemistry, physiology, immunology, and low-level resistance to gut infections (7, 30). Furthermore, most of the Lactobacillus strains currently used as probiotics are not adequate model organisms with which to study ecological aspects of gut colonization, as they belong to species that have never been shown to form stable populations in this ecosystem. It is important to note that the majority of traditional probiotic strains are probably allochthonous to the intestinal tract, and they show very little ability to persist in the human gut. Strains that form stable populations (over several months) in the intestinal tract without having significant upstream populations would show clear characteristics of autochthonous intestinal inhabitants. Several studies revealed that the total bacterial population, as well as bacterial groups such as bifidobacteria, Bacteroides spp., and clostridia, show a high degree of temporal stability down to the species level (82, 96, 100, 112). An even more puzzling finding was that the proportion of the mutants in the cecum always mirrored that in the forestomach in individual animals (Fig. For lactobacilli to become successful in this respect, ecological and functional aspects of the strains should already be considered when candidates are screened. In addition, the Lactobacillus succession that has been observed in the crop of chicks is remarkably similar to that in the ileum (94), suggesting that the Lactobacillus microbiota in the intestinal tract of these animals consists of bacteria originating from the crop. Redress of these ecological and immunological imbalances, for instance by probiotics, has the potential to ameliorate and prevent disease (25). in the production of fermented milk products or sausages) or as probiotics.3–6 In probiotic applications, selected LAB strains mainl… Similar to other genera in the family, Lactobacillus … These sporadic strains belonged to the L. acidophilus, L. crispatus, L. gasseri, and L. plantarum species and the L. casei group (L. casei, L. paracasei, and L. rhamnosus) (96). In this respect, it is important to point out that as the probes used for the quantification of lactobacilli by FISH are not specific for lactobacilli, the real numbers of lactobacilli could be even less. Since every ecological niche can support the existence of only one type (according to the niche exclusion theory), it is extremely difficult for an organism that is accidentally or intentionally introduced into the gut to gain access (32). The titrable acidity (% lactic acid) of reference samples are AEY 1.06%, APY 1.32%, AGY 0.95%, ACY 1.19%, BEY 1.09% and BCY 1.61%. For this purpose, it appears that the utilization of autochthonous strains makes it more likely that the recombinant organisms will persist, metabolize, and produce sufficient amounts of the therapeutic compound at a desired location in the gut. These interactions comprise modulation of signal transduction pathways and gene expression in epithelial and immune cells, and their high level of complexity makes it unlikely that they have emerged by coincidence. So, it is unlikely that inactivation of Lsp, a protein involved in adherence to the forestomach epithelium, would result in reduced population levels in the distal intestinal tract (102). Accordingly, in the era following Metchnikoff, lactobacilli were identified as one of the dominant organisms in the human gut (91). As shown below, the identification of the exact ecological status of individual Lactobacillus species in the human GIT remains a major challenge. Indeed, many researchers consider human origin as an important criterion for the selection of probiotics (21, 66, 77). Lactobacillus acidophilus … Lactobacilli could utilize simple carbohydrates that result from the degradation of complex carbohydrates by other microbes. Unlike the human stomach, which is lined with a glandular mucosa, the stomachs of pigs, mice, and rats and the crops of birds are lined, at least partly, with a nonglandular, squamous stratified epithelium (92). 0 Species such as L. acidophilus, L. casei, L. paracasei, L. rhamnosus, L. delbrueckii, L. brevis, L. johnsonii, L. plantarum, and L. fermentum have, so far, not been reported to form stable populations in the gut and are likely to be allochthonous. They are commonly used as probiotics, which are defined by the FAO/WHO as live microorganisms that when administered in adequate amounts confer a health benefit on the host. It has been shown that some lactobacilli have the ability to bind to intestinal mucus and polymers associated with the surface of enterocytes (64, 75), and putative adherence factors of lactobacilli have been identified (101). Lactic acid bacteria: microbiology and functional aspects. Many niches are likely to be determined by anatomical, immunological, and physiological characteristics of the host species. Evidence for significant in vivo association of lactobacilli with the columnar epithelium in the intestinal tract of humans is still inconclusive, and more work is needed to determine if the association with the epithelium contributes to the persistence of lactobacilli in the human gut. Although knowledge about their exact ecological function is still rudimentary, the findings suggest that bacterial adherence to the squamous stratified epithelium of the forestomach of mice is a key feature. High-throughput analysis of 16S rRNA sequences retrieved directly by PCR now allows a comprehensive view of the microbial diversity of the human GIT (25). The mechanisms underlying these interactions are likely to be specific for a particular microbe and its host and are probably influenced by other partners of the gut microbiota. Autochthonous strains that naturally persist in human subjects over long periods are tested by nature for their functionality in the gut, and they are likely to possess adaptive traits to benefit their human host. Copyright © 2021 American Society for Microbiology | Privacy Policy | Website feedback, Print ISSN: 0099-2240; Online ISSN: 1098-5336, Department of Food Science and Technology, University of Nebraska, Lincoln, Nebraska, Ecological Role of Lactobacilli in the Gastrointestinal Tract: Implications for Fundamental and Biomedical Research, Sign In to Email Alerts with your Email Address. We found only 3 reported cases of Lactobacillus … Analysis of the gut microbiota of rodents. These findings suggest that numerically dominant Lactobacillus populations present in the rodent, pig, and chicken intestinal tract are allochthonous and that they originate from the forestomach, pars esophagus, and crop, respectively. ASM journals are the most prominent publications in the field, delivering up-to-date and authoritative coverage of both basic and clinical microbiology. Lactobacilli offer exciting research opportunities, both in terms of biomedical applications and in acquiring fundamental knowledge about the functionality of gut microbes (94). Generic drug Lactobacillus Acidophilus combination available with manufacturer details. Because of the variations in physical and chemical properties in the different compartments of the GIT, specific microbial communities exist in the stomach, small intestine, and large intestine (93). Mechanistic explanations for this strain specificity are so far lacking, but it is likely that the distinct evolutionary histories of currently used probiotic strains are at least partly responsible for their different effects. Bacterial factors with similar function have recently been shown to be important for Bacteroides thetaiotaomicron in the murine gut (68). To view a list of products filtered by media type, move your cursor over the 'Media Type' … In aquaculture, one of the problem is related to water quality. The astounding degree of microbial diversity in the GIT indicates a multitude of ecological niches. ARE THE MAJORITY OF LACTOBACILLI IN THE INTESTINAL TRACT OF RODENTS, PIGS, AND CHICKENS ALLOCHTHONOUS? 1E). Lactobacillus is a lactic acid bacterium naturally found in the gastrointestinal tract and vagina. The lactic acid bacteria in health and disease. (104). In this review, evidence is summarized that suggests that only a small number of Lactobacillus species are true inhabitants of the mammalian intestinal tract and that most lactobacilli present are allochthonous members derived from fermented food, the oral cavity, or more proximal parts of the GIT. One would assume that allochthonous organisms are also more successful in the prevention of atopic diseases in early life because the immune system will experience novel antigenic complexes with the encounter of the bacterial strains. Lactobacilli require amino acids, peptides, nucleic acid derivatives, vitamins, salts, fatty acid esters, and fermentable carbohydrates for growth, and they have very limited abilities to utilize complex carbohydrates (39). More and more Lactobacillus genomes are becoming available, allowing systematic comparative and functional genomic studies to investigate ecological and probiotic functionality. Sequencing is a very promising tool for taxonomic and functional analysis of the Lactobacillus genome. It remains to be determined whether this also accounts for other Lactobacillus species present in the intestinal tract of rodents, pigs, and birds. Thank you for sharing this Applied and Environmental Microbiology article. However, following the succession and population dynamics of a bacterial group within the gut microbiota does permit the identification of some allochthonous bacteria: they do not persist within the ecosystem and are detectable only for a limited time. NOTE: We request your email address only to inform the recipient that it was you who recommended this article, and that it is not junk mail. Interestingly, the species that predominate in the oral cavity, such as L. acidophilus, L. gasseri, L. crispatus, L. plantarum, L. salivarius, L. brevis, L. rhamnosus, L. paracasei, and L. vaginalis, are also frequently isolated from human feces, and the species composition present in the oral cavity and in fecal samples coincides in some humans (16, 60). If you have questions about how to store this medicine (lactobacillus … Most lactobacilli present in the GIT of mice, rats, pigs, and chickens are clearly autochthonous, since they form stable populations throughout the life of the animal host, they can be cultured in large numbers, and they are present in almost all animals (62, 92). When total anaerobic culturing techniques are used, lactobacilli form a very small proportion of the cultivable human fecal microbiota and can rarely be cultured at population levels exceeding 108 CFU per gram. Consequently, lactobacilli gained a reputation as numerically dominant intestinal inhabitants, and even the advent of anaerobic culture techniques did little to correct this situation. Representation of Lactobacillus sequences in molecular-phylogenetic analysis of human gastrointestinal microbiota. However, the studies cited above show that caution is advised when particular Lactobacillus species are described as real (autochthonous) inhabitants. The autochthonous microbe-immune system relationship in healthy animals is characterized by tolerance, while the exposure to allochthonous bacteria results in a stronger immune response (8, 9). endstream endobj 76 0 obj <>>>/Lang(en-US)/Metadata 38 0 R/Outlines 70 0 R/PageLabels 71 0 R/PageLayout/OneColumn/Pages 73 0 R/Type/Catalog/ViewerPreferences<>>> endobj 77 0 obj <>/ExtGState<>/Font<>/ProcSet[/PDF/Text]/Properties<>/XObject<>>>/Rotate 0/Tabs/W/Thumb 33 0 R/TrimBox[0.0 0.0 595.276 841.89]/Type/Page>> endobj 78 0 obj <>stream Evolutionary theory predicts that in a spatially heterogeneous environment, vacant niches become occupied by organisms, and natural selection favors the emergence of ecological specialists that are highly adapted to the available niches (40). Lactobacillus bacteremia of renal origin is rarely described in the medical literature. These early findings were confirmed more recently by Tannock and coworkers (96), who followed the temporal succession of Lactobacillus strains by molecular strain typing (41, 96). Furthermore, several bacterial factors (e.g., MsrB, Dlt, and immunoglobulin A [IgA] protease) were identified as important, allowing the bacteria to overcome adverse environmental conditions generated through high acidity or innate and adaptive host defenses (nitric oxide and IgA). Traditional mixed … However, it is important to consider the ecological characteristics of individual species and their relationship with their host in such studies. Overall, the comprehensive molecular-phylogenetic analysis of the human gut microbiota now provides clear evidence for the numerically minor proportion of lactobacilli. DGGE was performed as described by Walter et al. However, many niches are also generated through the development of complex food webs (niche construction) where the product of one microbe becomes the substrate for another (18, 48). In contrast, lactobacilli (together with clostridia, enterococci, and Escherichia coli) could be cultured with relative ease due to their higher oxygen tolerances. It has been clearly shown that gut microbes benefit their host in many aspects (4). Eckburg and coworkers (22) studied 11,831 bacterial near-full-length 16S rRNA sequences retrieved from cecal, colonial, and fecal samples (including those from biopsy samples) of three human subjects and, remarkably, found not one single Lactobacillus sequence. Lactobacillus is a type of "friendly" bacteria. In a review of 55 patients with Lactobacillus bacteremia, only 6 patients had underlying urinary tract infection as a clinical manifestation [2, 5].Furthermore, pyelonephritis from Lactobacillus is even more rarely reported. These strains might nonetheless be excellent probiotics with respect to activation of the immune system. As noted by Morelli, there is considerable doubt about the real value of the current selection criteria for probiotics, such as their tolerance to the hostile conditions of the stomach and the small intestine and their ability to adhere to intestinal surfaces of epithelial cell lines (65). This is generally believed to be due to competitive exclusion conferred by the resident gut bacteria and to individual differences between human subjects. In contrast, the response of an allochthonous organism to the gut environment is likely to be based on signals that are generic (e.g., stress response, basic metabolism) and, hence, will reveal neither much about the environment from which the organism originates nor how autochthonous lactobacilli manage to live in the gut. (104). How do lactobacilli maintain such high cell numbers in the distal GIT of these animals? Based on available data, the Lactobacillus genome varies from 1 993 564 bp for Lb. It would be fascinating to investigate the probiotic characteristics of strains proven to be autochthonous, both in relation to persistence and health benefits. Adherence to these epithelia appears to be more relevant than adherence to columnar epithelia or mucus present in the intestinal tract, and the identification of adherence mechanisms to squamous cells would therefore teach us a lot about how lactobacilli manage to colonize their mammalian hosts. The physiology and expression of phenotypic traits of an autochthonous gut organism colonizing the GIT is a dynamic entity that reflects the microbe's adaptation to the ecosystem and its specific host. In this respect, it is important to emphasize that colonization of mucus associated with tissue surfaces by members of the gastrointestinal microbiota is very limited in humans, and the numbers of bacteria obtained from washed tissue surfaces are considerably lower than those observed in studies of rodents (93). Although nutrient availability is highest proximal to sites of absorption (e.g., the stomach and the first two-thirds of the small bowel), these sites contain relatively small numbers of microbes in humans. Traditionally, gut microbiota research relied on techniques that required cultivation of the microbes (91). Still, the adherence of lactobacilli to epithelia or mucus is often considered to contribute to the persistence of lactobacilli in the human GIT (69, 101). Genetic factors shown to contribute toward ecological performance of lactobacilli in the gut of mice. Representation of Lactobacillus sequences in the molecular-phylogenetic analysis of the gastrointestinal microbiota of animals. However, as shown in Table 1, this view is not supported by recent molecular characterizations of the microbiota present at these sites. Data points of individual animals are connected by lines. On the other hand, the activation of the immune system (such as enhanced phagocytosis and adjuvant effects) observed after the administration of some probiotic strains may reflect the allochthonous nature of the bacteria, and these bacteria might be more effective for the treatment or prevention of infectious and rotavirus-caused diarrhea (53, 84).