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Professor emeritus Göran Molin, Dept. Food Technology, Engineering and Nutrition, Lund University 2015-10-29 Lactobacillus paracasei 8700:2 CONTENT Consumption of live lactic acid bacteria (probiotics) – p. 2 Functional groups and taxonomically based taxa – p. 3 Lactic acid bacteria – p. 3 The species Lactobacillus paracasei – p. 4 Taxonomic considerations – p. 4 Characteristics – p. 5 The bacterial strain Lactobacillus paracasei 8700:2 – p. 6 Health effects p. 8 Antagonistic in vitro effects – p. 8 Animal models – p. 8 Translocation – p. 8 Mitigation of Enterobacteriaceae – p. 8 Multiple sclerosis – p. 9 Cortical bone loss – p. 9 Human trial – p. 9 Anti-oxidative activity – p. 9 Cell-mediated immunity – p. 10 Common cold infections – p. 10 Safety – p. 11 References – p. 12 Lactobacillus paracasei 8700:2 For further information please contact Probi AB who owns the commercial rights http://probi.se/en 1 Professor emeritus Göran Molin, Dept. Food Technology, Engineering and Nutrition, Lund University 2015-10-29 Consumption of live lactic acid bacteria Consumption of live lactic acid bacteria (LAB), included in fermented foods, has been a regular part of the human food intake for a long time. In fact, there are archaeological signs that humankind has used this technique from the beginning of time; it was presumably invented 1.5 million years ago by the early humanoids (Leakey 1993; Leakey 1995). Thus, humans have in this way consumed large numbers of live LAB throughout their entire history. Fermentation is the simplest and often the safest way to preserve food, and before the Industrial Revolution, fermentation was applied just as much in Europe as it still is in many rural areas of the World. Thus, it could very well be that the human digestive tract evolved to adapt to a more or less daily supply of live LAB. This supply of live LAB ceased in many industrialized countries during the twentieth century, which eventually may have led to increased frequency of gastro-intestinal (GI) and immunological dysfunctions in urbanised humans. When beneficial effects of certain types of live bacteria have been discussed, these types of bacteria have been gradually called “probiotics”. The original concept of probiotics implies that the balance between beneficial and harmful bacteria in the microbiota of the GI-tract can be positively affected by eating the right type of live microorganisms (Parker 1974; Fuller 1989). However, the concept of probiotics is today used more generally for describing live bacteria that after ingestion, exercise health beneficial effects beyond conventional nutrition. It is presupposed that these health beneficial effects have been scientifically proved. Lactobacillus paracasei 8700:2 For further information please contact Probi AB who owns the commercial rights http://probi.se/en 2 Professor emeritus Göran Molin, Dept. Food Technology, Engineering and Nutrition, Lund University 2015-10-29 Functional groups and taxonomically based taxa Lactic acid bacteria The bacteria performing the conversion of carbohydrates to carboxylic acids, mainly lactic acid in traditional fermented foods, are called lactic acid bacteria (LAB). Food microbiologists used the term early, and 1919 the Danish bacteriologist Orla Jensen tried to define key features of LAB, unaware of the fact that LAB is not forming a systematically defined group based on evolutionary relationships; instead it can be regarded as a functional group used by food microbiologists, aiming at those bacteria that occur and multiply spontaneously in traditional lactic acid fermented foods. Furthermore, it is understood that LAB are harmless to human health. Already 2002, it was shown in meta-analyses of published clinical trials that different kind of LAB can be used to prevent antibiotic associated diarrhoea (D’Souza et al. 2002) and shorten the duration of acute diarrhoeal illness in children (Huang et al. 2002). From the taxonomic point of view, LAB means a relatively wide variety of different taxonomically based groups (taxa). The only absolute condition for organisms involved in lactic acid fermentation of food must be that the bacteria mainly produce lactic acid and that they are harmless to consume in high numbers, even for consumers with underlying sicknesses that may have weaken their immunological defence. The different kind of lactic acid producing bacteria frequently occurring in high numbers in traditional, spontaneously fermented foods belong to genera as Lactobacillus, Pediococcus, Weissella, Leuconostoc, Oenococcus, Lactococcus, and the species Streptococcus thermophilus (and similar species). The genera Lactobacillus and Pediococcus belong to the family Lactobacillaceae which also includes the relatively new genera Paralactobacillus and Sharpea. They can all be included in the trivial expression "lactobacilli”. Leuconostoc, Weissella and Oenococcus belong to the family Leuconostocaceae together with the genus Fructobacillus. Lactococcus and S. thermophilus have from the phylogenetic point of view Lactobacillus paracasei 8700:2 For further information please contact Probi AB who owns the commercial rights http://probi.se/en 3 Professor emeritus Göran Molin, Dept. Food Technology, Engineering and Nutrition, Lund University 2015-10-29 relatively little in common with Lactobacillaceae and Leuconostocaceae even if they all are included in the order of Lactobacillales. The species Lactobacillus paracasei Taxonomic considerations Lactobacillus paracasei is a bacterial species in the huge and relatively diverse genus of Lactobacillus, which comprises around 90 validly named species. Lactobacillus paracasei is a so called facultatively heterofermentative Lactobacillus, i.e. L. paracasei ferment hexoses exclusively to lactic acid, but can also ferment pentoses and/or gluconate, and then producing lactic and acetic acid (Kandler and Weiss 1986). The type strain of L. paracasei is NCDO 151 T (= NCFB 151 T; T = type strain). The systematics of the two species L. paracasei and L. casei was under discussion for several years: It started with the loss of the original type strain of L. casei; L. casei was originally described by Orla-Jensen 1916, and referred by Kandler and Weiss in Bergey’s Manual of Systematic Bacteriology (Kandler and Weiss 1986). However, when the original type strain had been lost, Hansen and Lessel (1971) designated strain ATCC 393 as the neotype of L. casei (ATCC 393T). Five subspecies were recognized within this restored species of L. casei: Lactobacillus casei subsp. casei (Hansen and Lessel, 1971; Orla-Jensen 1916), Lactobacillus casei subsp. pseudoplantarum (Abo-Elnaga and Kandler 1965; ATCC 25598T), Lactobacillus casei subsp. tolerans (AboElnaga and Kandler 1965; ATCC 25599T), Lactobacillus casei subsp. rhamnosus (Hansen 1968; ATCC 7469T) and Lactobacillus casei subsp. alactosus (Mills and Lessel. (1973; ATCC 27216T). However, Johnson (1973) and Dellaglio et al. (1975) found high DNA:DNA homology between strains designated L. casei subsp. casei and L. casei subsp. alactosus. Furthermore, in Bergey’s Manual of Sytematic Bacteriology (Kandler and Weiss, 1986) L. casei subsp. alactosus was not mentioned. Though, it was commented that L. casei subsp. casei ATCC 393T and strains of L. casei subsp. rhamnosus had low DNA:DNA homology to other L. casei strains. This was confirmed by Collins et al. (1989), and it was suggested that members of L. casei subsp. alactosus, L. casei subsp. pseudoplantarum and L. casei subsp. tolerans, and the majority of the tested L. casei subsp. casei strains should be given separate species status, and the names Lactobacillus paracasei subsp. paracasei (NCDO 151T), and L. paracasei subsp. tolerans (ATCC 25599T) were proposed (L. paracasei sp. nov.). It was also proposed that L. casei subsp. rhamnosus should be elevated to species status, as L. rhamnosus sp. nov. (ATCC 7469T; Collins et al., 1989). Lactobacillus paracasei 8700:2 For further information please contact Probi AB who owns the commercial rights http://probi.se/en 4 Professor emeritus Göran Molin, Dept. Food Technology, Engineering and Nutrition, Lund University 2015-10-29 However, the relevance of the type strain of L. casei subsp. casei (ATCC 393T), that had been selected as the neotype strain by Hansen and Lessel (1971), was questioned by Dellaglio et al. (1991); instead they suggested a new type strain for L. casei subsp. casei (ATCC 334), and suggested a rejection of the species name L. paracasei. But these suggestions were not approved by the Judicial Commission of the International Committee on Systematic Bacteriology (Wayne, 1994). However, Dicks et al. (1996) showed that the type strain, L. casei subsp. casei ATCC 393T, exhibited high DNA:DNA homology to the reference strain, L. rhamnosus ATCC 15820, that is the former type strain of ”Lactobacterium zeae” (Kuznetsov 1959). On the basis of this, and results showing that L. casei subsp. casei ATCC 393T was separated from authentic L. casei and L. paracasei strains, Dicks et al. (1996) proposed that L. casei subsp. casei ATCC 393T and L. rhamnosus ATCC 15820 should be reclassified as members of Lactobacillus zeae nom. rev. (ATCC 15820T), that strain ATCC 334 should be designated the neotype strain of L. casei subsp. casei, and that the name L. paracasei should be rejected. Comparative sequence analyses of the genes coding for 16S rRNA have later shown that the type strains of L. zeae, L. casei (ATCC 393T), L. paracasei and L. rhamnosus were all different, and the variation in the 16S ribosomal DNA (rDNA) sequences was situated between the positions 69 and 100 (Escherichia coli numbering; Mori et al., 1997). The Judicial Commission of the International Committee on Systematics of Bacteria has come to the conclusion that Lactobacillus paracasei should remain as name of the species, with the type strain Lactobacillus paracasei subsp. paracasei NCDO 151T (Tindall, 2008). Well-known probiotic strains as L. casei “Shirota” (Yakult, in the product Yakult) and L. casei “defensis” (Danone, in the product Actimel) should thus presumably be designated L. paracasei instead of L. casei. Characteristics L. paracasei differs from many other Lactobacillus spp. in the following points: 1) L. paracasei grows very well in cheese during ripening. 2) L. paracasei is relatively resistant to heat. 3) L. paracasei has comparably high proteolytic activity. The species L. paracasei is frequently present on human gastro-intestinal (GI) mucosa of healthy individuals (Molin et al. 1993; Ahrné et al. 1998), but is also often dominating the spontaneous, secondary bacterial-flora in semi-dry cheese, especially if the cheese has been manufactured with pasteurised milk (Antonsson 1991; Antonsson et al. 2001; Antonsson et al. 2003). It has been shown that different strains of heat killed, whole cells of L. paracasei were more efficient in triggering the production of the regulatory Lactobacillus paracasei 8700:2 For further information please contact Probi AB who owns the commercial rights http://probi.se/en 5 Professor emeritus Göran Molin, Dept. Food Technology, Engineering and Nutrition, Lund University 2015-10-29 cytokin IL-12 in human blood mononuclear cells (monocytes) than were cells of Lactobacillus plantarum or Lactobacillus rhamnosus (Hessle et al. 1999). Different strains of L. paracasei has for long been used as probiotics in a wide range of different probiotic products, marketed in many countries. The most well known strains are L. paracasei strain “Shirota” (labelled “casei” by the manufacturer, Yakult) and L. paracasei strain “Immunitas/Defencis” (labelled “casei” by the manufacturer, Danone). These particular strains of L. paracasei have shown to have health beneficial effects. The bacterial strain Lactobacillus paracasei 8700:2 The L. paracasei strain 8700:2 (= DSM 13434) (Antonsson et al. 2002) has been isolated from healthy human colonic mucosa (Ahrné et al. 1998). The strain is growing quickly in milk and thrives in cheese and in yoghurt. L. paracasei 8700:2 can be defined and identified by restriction endonculease analysis (REA) of total chromosomal DNA by the use of relatively frequently cutting restriction enzymes such as EcoRI and ClaI, and traditional agarose gel electrophoresis (Johansson et al. 1995; Vásquez et al. 2004). L. paracasei 8700:2 has a relatively close genomic similarity to “L. casei” ATCC 334 and the former type strain of L. casei subspecies pseudoplantarum (DSM 20008) (Vásquez et al. 2004). The genomic relationship between L. paracasei 8700:2 and L. paracasei 02A (= DSM 13432), and the relationships to type strains and a battery of reference strains of the L. paracasei/casei-complex have been scrutinised (Vásquez et al. 2004). L. paracasei 8700:2 (= DSM 13434), has primarily been selected on the basis of the ability to grow in Swedish semi-hard cheese during storage and beneficially contribute to the sensory quality during ripening (Antonsson et al. 2002). However, the strain has also probiotic potential as it after administration in cheese to healthy volunteers could be re-isolated from faeces of the consumers (Antonsson 2001). Irrespectively of what strain that is used as probiotics, a condition must be that the bacterium survives and remains active during the passage through the gastro-intestinal tract. The ability of L. paracasei 8700:2 to survive the passage through the human gastro-intestinal tract when administrated in cheese has been proved (Antonsson 2001). L. paracasei 8700:2 has the capability to degrade oligofructose and long-chain inulin (Makras et al. 2005). L. paracasei 8700:2 grows rapidly on both oligofructose and inulin, with lactic acid as the main metabolic end-product (Makras et al. 2005). In a comparison between the ability of six different Lactobacillus paracasei 8700:2 For further information please contact Probi AB who owns the commercial rights http://probi.se/en 6 Professor emeritus Göran Molin, Dept. Food Technology, Engineering and Nutrition, Lund University 2015-10-29 probiotic strains to grow and multiply on nine different commercially carbohydrates (saccharides or fructooligosaccharides) with purported prebiotic properties, L. paracasei 8700:2 fermented seven of the tested carbohydrates while for example Lactobacillus rhamnosus GG were unable to ferment any of the tested prebiotics (Saulnier et al. 2008). L. paracasei 8700:2 was the only one of the tested probiotic strains that was able to degrade all the testfructans. L. paracasei 8700:2 can adhere to HT-29 cells via mannose-sensitive mechanisms (Rask et al. 2013). Lactobacillus paracasei 8700:2 For further information please contact Probi AB who owns the commercial rights http://probi.se/en 7 Professor emeritus Göran Molin, Dept. Food Technology, Engineering and Nutrition, Lund University 2015-10-29 Health effects Antagonistic in vitro effects L. paracasei 8700:2 has been shown in vitro to possess strong antagonistic properties against Salmonella enterica subsp. enterica, and an “intermediate” antagonistic activity against Helicobacter pylori, Shigella sonnei and Escherichia coli (Hütt et al. 2006). Animal models Translocation Translocation, i.e. the passage of viable bacteria through the epithelial mucosa into the lamina propria and then to the mesenteric lymph nodes and possibly other tissues (Berg and Garlington, 1979), was reduced in rats with colitis by treatment with L. paracasei 8700:2 (Osman et al. 2004). The colitis was induced by giving the rat 5 % (w/v) dextran sulphate sodium (DSS) dissolved in drinking water for 7 days. Samples were collected on the 7th day for examination of the bacterial translocation. The total translocation to the mesenteric lymph nodes and the translocation of Enterobacteriaceae to the liver decreased significantly when the colitis rats were treated with L. paracasei 8700:2. Mitigation of Enterobacteriaceae The viable count of Enterobacteriaceae in the colon decreased in rats with DSS-induced colitis by treatment with L. paracasei 8700:2 (Osman et al. 2004). Also, pretreatment with L. paracasei 8700:2 in an acute liver injury model where the injury had been induced by D-galactosamine decreased the viable count of Enterobacteriaceae in the colon (Osman et al. 2005). Lactobacillus paracasei 8700:2 For further information please contact Probi AB who owns the commercial rights http://probi.se/en 8 Professor emeritus Göran Molin, Dept. Food Technology, Engineering and Nutrition, Lund University 2015-10-29 Multiple sclerosis Multiple sclerosis (MS) is a Th1 cell-mediated chronic inflammatory disease of the central nervous system. Treatment with L. paracasei 8700:2 in a mouse model for experimental autoimmune encephalomyelitis (EAE), mimicking MS, prevented and delayed the onset of the clinical signs of EAE compared to control mice (Lavasani et al. 2010). In contrast, treatment with Lactobacillus paracasei PCC 101 or Lactobacillus delbrueckii subsp. bulgaricus DSM 20081 had no effect on the disease development. L. paracasei 8700:2 induced CD4+CD25+Foxp3+ regulatory T cells in mesenteric lymph nodes and enhanced production of serum TGF-beta1 (Lavasani et al. 2010). Cortical bone loss With the aim to determine if probiotics can protect from ovariectomy-inuced bone loss, mice were given L. paracasei 8700:2 in the drinking water for 6 weeks, starting 2 weeks prior before ovariectomy (Ohlsson et al. 2014). It was found that L. paracasei 8700:2 decreased the serum levels of the resorption marker C-terminal telopeptides and the urinary fractional excretion of calcium after ovariectomy, and reduced the expression of the two inflammatory cytokines, TNF-alfa and IL-1-beta, and increased the expression of osteoprotegerin (OPG) which is a potent inhibitor of osteoclastogenesis, in cortical bone. The probiotic treatment also improved the frequency of regulatory T cells in bone marrow (Ohlsson et al. 2014). The authors conclude that “treatment with L. paracasei 8700:2 prevents ovariectomy-induced cortical bone loss”, and the findings “indicate that the probiotic treatment alter the immune status in bone resulting in attenuated bone resorptione” (Ohlsson et al. 2014). Human trial Anti-oxidative activity The impact on the antioxidative activity markers of blood in asymptomatic Helicobacter pylori-colonized persons of a product containing a mixture of L. paracasei 8700:2, Lactobacillus fermentum ME-3 and Bifidobacterium longum 46, together with Raftilose, has been evaluated in a randomized, double-blind Lactobacillus paracasei 8700:2 For further information please contact Probi AB who owns the commercial rights http://probi.se/en 9 Professor emeritus Göran Molin, Dept. Food Technology, Engineering and Nutrition, Lund University 2015-10-29 placebo-controlled study (Hütt et al. 2009). After consumption of the product for three weeks the total antioxidative status of serum increased in H. pyloripositive subjects, while the ratio between oxidized and reduced gluthione decreased. Cell-mediated immunity In a blind placebo-controlled study, the effect of a daily intake for 2 weeks of L. paracasei 8700:2 on the innate and acquired immune system was investigated in vivo (Rask et al. 2013). Blood lymphocyte subsets were quantified by flow cytometry (FACS) and the expression of activation and memory markers was determined. The strain was also examined for its capacity to be phagocytosed by human peripheral blood mononuclear cells (PBMCs). Intake of L. paracasei 8700:2 tended to expand the NKT cell population. Also, the phagocytic activity of granulocytes towards Escherichia coli was increased. (Rask et al. 2013). Common cold infections In combination with the strain Lactobacillus plantarum HEAL9 (=DSM 15312), L. paracasei 8700:2 was supplemented daily to 272 subjects for a 12week period in a randomised, parallel, double-blind placebo controlled study with the intention to clarify if the treatment product could reduce the risk of common cold episodes, number of days with common cold symptoms, frequency and severity of symptoms, and cellular immune response in common cold infections (Berggren et al. 2010). It was shown that the incidence of acquiring common cold episodes was reduced, and so was the number of days with common cold symptoms. Also, the pharyngeal symptoms were reduced, and the proliferation of B lymphocytes was counteracted (Berggren et al. 2010). In another randomized, double blind and placebo controlled study with the same mixture of L. plantarum HEAL9 and L. paracasei 8700:2 (ProbiDefendum), including totally 310 subjects with increased risk for common cold infection (Busch et al. 2013). It was concluded from the study results that “the daily intake of the probiotic dietary supplement ProbiDefendum over a period of 12 weeks efficiently alleviated symptoms of common cold and the duration of cold episodes” (Busch et al. 2013). Lactobacillus paracasei 8700:2 For further information please contact Probi AB who owns the commercial rights http://probi.se/en 10 Professor emeritus Göran Molin, Dept. Food Technology, Engineering and Nutrition, Lund University 2015-10-29 Safety The species L. paracasei is present in cheese as it multiplies spontaneously during the ripening process and often reaches numbers around 107 CFU per g of cheese. Thus the species has been eaten alive in high numbers by innumerable numbers of humans over an immense period of time without any empirically found hazards. The strain L. paracasei 8700:2 has been evaluated in the EU funded PROSAFE project (Vankerckhoven et al. 2008). The identity of the strain was confirmed and no acquired antibiotic resistance could be detected (PRO SAFE report on strain Lactobacillus paracasei 8700:2). Lactobacillus paracasei 8700:2 For further information please contact Probi AB who owns the commercial rights http://probi.se/en 11 Professor emeritus Göran Molin, Dept. Food Technology, Engineering and Nutrition, Lund University 2015-10-29 References Abo-Elnaga, I.G. and Kandler, O. (1965). Zur Taxonomie der Gattung Lactobacillus Beijerinck. I. Das Subgenus Streptobacterium Orla-Jensen. Zentralbl. Bacteriol. II Abt. 119: 1-36. Ahrné, S., Nobaek, S., Jeppsson, B., Adlerberth, I., Wold, A., and Molin, G. (1998). The normal Lactobacillus flora of healthy human rectal and oral mucosa, J. Appl. Microbiol., 85: 88-94. Antonsson, M. (2001). Lactobacillus in semi-hard cheese and their use as adjunct cultures. Ph.D. thesis (15 June), Division of food technology, Lund institute of technology, Lund university, Lund, Sweden. Antonsson, M., Ardö, Y. and Molin, G. (2001). A comparison between the microflora of Herrgård cheese from three different dairies. International Dairy Journal 11:285-291. Antonsson, M., Ardö, Y., Nilsson, B.F. and Molin, G. (2002). Screening and selection of Lactobacillus strains for use as adjunct cultures in production of semi-hard cheese. Journal of Dairy Research 69 457-472. Antonsson, M, Molin, G. & Ardö, Y. (2003). Lactobacillus strains isolated from Danbo cheese as adjunct cultures in a cheese model system. International Journal of Food Microbiology 85: 159-169. Berg, R.D. and Garlington, A.W. (1979). Translocation of certain indigenous bacteria from the gastrointestinal tract to the mesenteric lymph nodes and other organs in a gnotobotic mouse model, Infect. Immunol., 23, 403-411. Berggren, A., Lazou Ahrén, I., Larsson, N. and Önning, G. (2010). Randomised, double-blind and placebo-controlled study using new probiotic lactobacilli for strengthening the body immune defence against viral infections. European Journal of Nutrition DOI 10.1007/s00394-010-0127-6 Busch, R., Gruenwald, J. and Dudek, S. (2013). Randomized, double blind and placebo controlled study using a combination of two probiotic lactobacilli to alleviate symptoms and frequency of common cold. Food and Nutrition Sciences 4: 13-20. Lactobacillus paracasei 8700:2 For further information please contact Probi AB who owns the commercial rights http://probi.se/en 12 Professor emeritus Göran Molin, Dept. Food Technology, Engineering and Nutrition, Lund University 2015-10-29 Collins, M.D., Phillips, B.A. and Zanoni, P. (1989). Deoxyribonucleic acid homology studies of Lactobacillus casei, Lactobacillus paracasei sp. nov., subsp. paracasei and subsp. tolerans, and Lactobacillus rhamnosus sp. nov., comb. nov. Int. J. Syst. Bacteriol. 39: 105-108. Dellaglio, F., Bottazzi, V. and Vescovo, M. (1975). Deoxyribonucleic acid homology among Lactobacillus species of the subgenus Streptobacterium OrlaJensen. Int. J. Syst.Bacteriol. 25: 160-172. Dellaglio, F., Dicks, L.M.T., Du Toit, M. and Torriani, S. (1991). Designation of ATCC 334 in place of ATCC 393 (NCDO 161) as the neotype strain of Lactobacillus casei subsp. casei and rejection of the name Lactobacillus paracasei (Collins et al. 1989). Request for an opinion. Int. J. Syst. Bacteriol. 41: 340-342. Dicks, L.M.T., Du Plessis, E.M., Dellaglio, F. and Lauer, E. (1996). Reclassification of Lactobacillus casei subsp. casei ATCC 393 and Lactobacillus rhamnosus ATCC 15820 as Lactobacillus zeae nom. rev., designation of ATCC 334 as neotype of L.casei subsp. casei, and rejection of the name Lactobacillus paracasei. Int. J. Syst. Bacteriol. 46: 337-340. D’Souza, A.L., Rajkumar, C., Cooke, J. and Bulpitt, C.J. (2002). Probiotics in prevention of antibiotic associated diarrhoea: meta-analysis. BMJ 324: 1361. Fuller, R. (1989). Probiotics in man and animals, J. Appl. Bacteriol., 66, 365368. Huang, J., Bousvaros, A., Lee, J.W., Diaz, A. and Davidson, E.J. (2002). Efficacy of probiotic use in acute diarrhea in children. Digestive Diseases and Sciences 47: 2625-2634. Hütt, P., Andreson, H., Kullisaar, T., Vihalemm, T., Unt, E., Kals, J., Kampus, K., Zilmer, M. and Mikelsaar, M. (2009). Effects of a synbiotic product on blood antioxidative activity in subjects colonized with Helicobacter pylori. Letters in Applied Microbiology 48: 797-800. Hütt, P., Shchepetova, J., Loivukene, K., Kullisaar, T. and Mikelsaar, M. (2006). Antagonistic activity of probiotic lactobacilli and bifidobacteria against entero- and uropathogens. Journal of Applied Microbiology 100: 1324-1332. Johansson, M.-L., Quednau, M., Ahrné, S., and Molin, G. (1995). Classification of Lactobacillus plantarum by restriction endonuclease analysis of total chromosomal DNA using conventional agarose gel electrophoresis, Int. J. System. Bacteriol., 45: 670-675. Johnson, J.L. (1973). Use of nucleic acid homologies in the taxonomy of anaerobic bacteria. Int. J. Syst. Bacteriol. 23: 308-315. Lactobacillus paracasei 8700:2 For further information please contact Probi AB who owns the commercial rights http://probi.se/en 13 Professor emeritus Göran Molin, Dept. Food Technology, Engineering and Nutrition, Lund University 2015-10-29 Hansen, P.A. (1968). Type strains of Lactobacillus species. A report by the taxonomic subcommittee on lactobacilli and closely related organisms. American Type Culture Collection, Rockville, Maryland. Hansen, P.A. & Lessel, E.F. (1971). Lactobacillus casei (Orla Jensen) comb. nov. Int. J. Syst. Bacteriol. 21: 69-71. Hütt, P., Shchepetova, J., Loivukene, K., Kullisaar, T. and Mikelsaar, M. (2006). Antagonistic activity of probiotic lactobacilli and bifidobacteria against entero- and urpathogens. Journal of Applied Microbiology 100: 1324-1332. Kandler, O. and Weiss, N. (1986). Regular, nonsporing Gram-positive rods, in Bergey’s Manual of Systematic Bacteriology, Sneath, H.A., Mair, N.S., Sharpe, M.E. and Holt, J. Eds., Williams & Wilkins, Baltimore, vol. 2, pp. 1208-1234. Kuznetsov, V.D. (1959). A new species of lactic acid bacteria. Mikrobiologiya 28: 248-351. Lavasani1, S., Dzhambazov, B., Nouri, M., Fåk, F., Buske, S., Molin, G., Thorlacius, H., Alenfall, J., Jeppsson, B. & Weström, B. (2010). A novel probiotic mixture exerts a therapeutic effect on experimental autoimmune encephalomyelitis mediated by IL-10 producing regulatory T cells. PLoS ONE 5 (2): 1-11 (www.plosone.org). Leakey, R. (1993). På spaning efter människans ursprung, Natur och Kultur, Stockholm [Swedish translation from: Origins reconsidered, in search of what makes us human, Bantam Doubleday Dell Publishing Group, Inc., New York, 1992]. Leakey, R. (1995). Hur människan blev till, Natur och Kultur, Stockholm [Swedish translation from: The origin of humankind, HarperCollins Publishers, Inc., 1994]. Makras, L., Van, G. and De Vuyst, L. (2005). Lactobacillus paracasei subsp. paracasei 8700:2 degrades inulin-type fructans exhibiting different degrees of <polymerization. Applied and Environmental Microbiology 71: 6531-6537. Mills, C.K. and Lessel, E.F. (1973). Designation and description of the type strain of Lactobacillus casei subsp. alactosus Rogosa et al. Int. J. Syst. Bacteriol. 23: 67-68. Molin, G., Jeppsson, B., Ahrné, S., Johansson, M.-L., Nobaek, S., Ståhl, M., and Bengmark, S. (1993). Numerical taxonomy of Lactobacillus spp. associated with healthy and diseased mucosa of the human intestines, J. Appl. Bacteriol. 74, 314-323. Lactobacillus paracasei 8700:2 For further information please contact Probi AB who owns the commercial rights http://probi.se/en 14 Professor emeritus Göran Molin, Dept. Food Technology, Engineering and Nutrition, Lund University 2015-10-29 Mori, K., Yamazaki, K., Ishiyama, T., Katsumata, M., Kobayashi, K., Kawai, Y., Inoue, N. and Shinano, H. (1997). Comparative sequence analyses of the genes coding for 16S rRNA of Lactobacillus casei-related taxa. Int. J. Syst. Bacterio.l 47: 54-57. Ohlsson, C., Engdahl, C., Fåk, F., Andersson, A., Windahl, S.H., Farman, H.H., Moverare-Skrtic, S., Islander, U. and Sjögren, K. (2014). Probiotics protect mice from ovariectomy-induced cortical bone loss. PLOS ONE 9 (3): e92368 www.plosone.org Osman, N., Adawi, D., Ahrne, S., Jeppsson, B. & Molin, G. (2004). Modulation of the effect of dextran sulfate sodium-induced acute colitis by the administration of different probiotic strains of Lactobacillus and Bifidobacterium. Digestive Diseases and Sciences 42: 320-327. Osman, N., Adawi, D., Ahrne, S., Jeppsson, B. & Molin, G. (2005). Probiotic strains of Lactobacillus and Bifidobacterium affect the translocation and intestinal load of Enterobacteriaceae differently after D-galactose-induced liver injury in rats. Microbial Ecology in Health and Disease 17:40-46. Parker, R.B. (1974). Probiotics, the other half of the antibiotic story, Anim. Nutr. Health, 29, 4-8. Tindall, B. J. (2008). The type strain of Lactobacillus casei is ATCC 393, ATCC 334 cannot serve as the type because it represents a different taxon, the name Lactobacillus paracasei and its subspecies names are not rejected and the revival of the name ‘Lactobacillus zeae’ contravenes Rules 51b (1) and (2) of the International Code of Nomenclature of Bacteria. Opinion 82. International Journal of Systematic and Evolutionary Microbiology 58: 1764–1765. Vankerckhoven, V., Huys, G., vancanneyt, M., Vael, C., Klare, I., Romond, MB., Entenza, J.M., Moreillon, P., Wind, R.D., Knol, J., Wiertz, E., Pot, B., Vaughan, E.E., Kahlmeter, G. and Goossens, H. (2008). Biosafety assessment of probiotics used for human consumption: recommendations from the EUPROSAFE project. Trends in Food Science & Technology 19: 102-114. Wayne, L.G. (1994). Actions of the Judicial Commission of the International Committee on Systematic Bacteriology on requests for opinions published between January 1985 and July 1993. International Journal of Systematic Bacteriology 44: 177-178. Vásquez, A., Molin, G., Pettersson, B., Antonsson, M. and Ahrné, S. (2005). DNA-based classification and sequence heterogeneities in the 16S rRNA genes of Lactobacillus casei/paracasei and related species. Systematic and Applied Microbiology 28: 430-441. Lactobacillus paracasei 8700:2 For further information please contact Probi AB who owns the commercial rights http://probi.se/en 15