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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
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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.
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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
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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
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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
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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
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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
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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
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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
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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
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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
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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
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Sciences 4: 13-20.
Lactobacillus paracasei 8700:2
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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
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Lactobacillus paracasei 8700:2
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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
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encephalomyelitis mediated by IL-10 producing regulatory T cells. PLoS ONE
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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
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Lactobacillus paracasei 8700:2
For further information please contact Probi AB who owns the commercial rights http://probi.se/en
15