Lactobacillus Delbrueckii Subsp.

Bul-
garicus
Related terms:

Lactose, Peptide, Bacterium, Fermentation, Microorganism, Lactic Acid Bacteri-

um, Bifidobacterium, Lactobacillus, Lactobacillus acidophilus, Streptococcus ther-
mophilus

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LACTOBACILLUS | Lactobacillus del-

brueckii ssp. bulgaricus
P. Teixeira, in Encyclopedia of Food Microbiology (Second Edition), 2014

Taxonomy
Lactobacillus delbrueckii subsp. bulgaricus (termed Lactobacillus bulgaricus hereafter;
type strain: ATCC 11842, CCUG 41390, CIP 101027, DSM, 20081, IFO 13953, JCM
1002, LMG 6901, LMG 13551, NCIMB 11778, NCTC 12712, VKM B-1923, WDCM
00102, P.A. Hansen Lb14, S. Orla-Jensen 14), one of the three subspecies of L.
delbrueckii, is an aerobic to anaerobic homofermentative bacterium (i.e., it converts
hexoses into lactic acid via the Emden–Meyerhof pathway) normally isolated from
yogurt and cheese. Carbohydrates fermented by L. bulgaricus (90% or more strains)
are fructose, glucose, and lactose. d (−) lactic acid is the major product of fermen-
tation; however, secondary products, such as acetaldehyde, acetone, acetoin, and
diacetyl, also can be produced in very low concentrations.

In lactic acid bacteria that do not possess superoxide dismutase, the dismutation of
superoxide normally is catalyzed by internally accumulated manganese. Lactobacillus
bulgaricus, however, has a low capacity to scavenge O2− because it does not have
superoxide dismutase or high levels of Mn (II) and it is sensitive to O2 (the ability to
grow aerobically must be distinguished from the ability to survive exposure to O2).
Cells are rod shaped with rounded ends, 0.5–0.8 × 2–9 μm. They are usually
separate or in short chains (Figure 1), but long chains can be observed in late
stationary-phase cultures (Figure 2).

Figure 1. Acridine orange staining of Lactobacillus bulgaricus at early stationary phase

of growth (×100).

Figure 2. Acridine orange staining of Lactobacillus bulgaricus at late stationary phase

of growth (×100).

The cells are generally short but sometimes long, straight, and often arranged in
palisades. Internal granulations are observed with the Gram reaction or methylene
blue stain, especially when cells become older. In addition to age, variability of L.
bulgaricus cellular morphology depends on the composition of the growth medium
and oxygen tension. Additional physiological and biochemical characteristics are
presented in Table 1.

Table 1. Physiological and biochemical characteristics of Lactobacillus bulgaricus

G+C DNA Chro- Pepti- Tei- Anti- Lac- Elec- Opti- Mini- Maxi- Opti- NH3
con- melt- mo- do- choic acid
genic group
tic mum
tophoretic growth
mumdgrowth
motility (°C)
mumbgrowth
-LDH (°C)
mum pH
(°C)
from
tent ing some size
gly-(Mbp) acid argi-
(mol tem- can iso- nine
%) pera- type mer
ture
(°C)a
49–51 91.7 1.8 Glyc- E D 1.70 40–50 22 62 5.5–5.8No
erol
 Lys--
d-
-Asp

a Approximate value determined by differential scanning calorimetry; individual

strains vary.
b LDH, lactic acid dehydrogenase.

Phylogenetically, L. bulgaricus is closely related to Lactobacillus acidophilus, Lacto-

bacillus helveticus, Lactobacillus johnsonii, and Lactobacillus gasseri; the guanine and
cytosine (G+C) ratio of L. bulgaricus is higher than that found among these related
species because of a significantly higher G+C content at the third codon position
than the overall G+C content. This suggests that an evolutionary change toward
higher overall G+C content is occurring in L. bulgaricus. Unique features revealed
by genome sequencing of L. bulgaricus (e.g., high number of pseudogenes, high
ratio of RNA genes to genome size, lack of transcriptional regulators) provide strong
evidence that the L. bulgaricus genome is in an active state of size reduction, possibly
through the loss of genes necessary for a plant-associated environment but no
longer necessary for a milk-associated environment.

Fatty acid composition has been used to group and classify these microorganisms.
As shown in Table 2, lipid compositions are different in the different strains. Hexade-
canoic (16:0), hexadecenoic (16:1), octadecenoic (18:1), and lactobacillic (19:0) acids
are the major fatty acids present that are common to the three L. bulgaricus strains. In
addition to the strain, variability can occur as a result of different growth conditions
(medium composition, temperature), phase of growth, and even the methodology
used for lipid extraction.

Table 2. Fatty acid composition of lipids from different Lactobacillus bulgaricus strains
determined by gas–liquid chromatography

Fatty acid content (%)

Strain 12:0 14:0 15:0 16:0 16:1 17:1 18:0 18:1 19:0
origin
Commer- tra 3.2 0.70 18.8 13.8 1.0 1.1 35.8 25.5
cial
(NCS1)
NCFB NDb ND ND 6.3 30.4 ND 0.72 35.4 21.2
1489
State 2.9 10.4 ND 23.0 13.8 ND 0.6 27.1 22.2
Universi-
ty,
Ul-
trecht-9LB

NCS1, Commercial culture of L. bulgaricus used in commercially in the production

of yoghurt and Italian cheese; NCFB, National Collection of Food Bacteria, United
Kingdom.
a The presence of a compound in an amount less than 0.5% is denoted tr.

b ND, not detected.

> Read full chapter

The Benefits of Yogurt, Cultures, and

Fermentation
M. Freitas, in The Microbiota in Gastrointestinal Pathophysiology, 2017

Abstract
Yogurt is a popular fermented milk product containing both Lactobacillus bulgaricus
and Streptococcus thermophilus cultures. Most yogurts contain several nutrients
that are lacking in the American diet, such as calcium, vitamin D, and potassium.
Both intervention and observational studies have associated yogurt consumption
with several benefits to human health, including increased digestibility of lactose
and assimilation of certain nutrients, enhancement of bone mineral acquisition,
weight management, heart health, metabolic health, and digestive and immune
health. Additional and special beneficial cultures can also be added to conventional
yogurt and provide additional health benefits. This chapter is primarily centered on
the benefits of regular yogurt consumption without added cultures. It describes
some of the most relevant studies on yogurt and discusses the plausible mechanisms
of action associated both with interactions at the gastrointestinal epithelium and
with the gut microbiota.

> Read full chapter

CHEESES | Starter Cultures Employed

in Cheese-making
G. Stanley, in Encyclopedia of Food Sciences and Nutrition (Second Edition), 2003

Thermophilic Starters
The most important thermophilic LAB are Streptococcus salivarius ssp. thermophilus
(Sc. thermophilus) and species of Lactobacillus (Lb): bulgaricus, helveticus and lactis.
They are used in those technologies where a temperature > 40 °C is used, such as
yogurt, Mozzarella, Emmenthal, and Gruyere. Generally, these fermentations use
blends of strains of Sc. thermophilus and a Lactobacillus (particulary Lb. bulgaricus),
there being an associative growth relationship between them: Sc. thermophilus pro-
duces formate that stimulates Lb. bulgaricus, and Lb. bulgaricus produces amino acids
that stimulate Sc. thermophilus.

Some differentiating characteristics of mesophilic and thermophilic acidifying

starter cultures are given in Table 1.

Table 1. Some differentiating characteristics of starter lactic acid bacteria

Type Species Shape Growth at

Lac- Cit- Galac- Ammo- Per- Per- Impor-
tate rate tose nia cent- cent- tant
iso- metab- metab- from age age meta-
mer olism olism argi- lactic salt bolic
nine acid inhibi- prod-
pro- tion ucts
duced
in milk

10 °C 40 °C 45 °C

Lac- Cocci +
Mesophilic + − l (+) − + + 0.8 4.0–6.5Lac-
tate
to-
coc-
cus
lac-
tis
ssp.
lac-
tis
Lacto- Cocci + + − l (+) + + + 0.8 4.0–6.5 Lac-
tate,
coc- di-
acetyl,
cus CO2
lactis
ssp.
lactis
(−cit+)
Lacto- Cocci + − − l (+) − + − 0.8 2.0–4.0 Lactate
coc-
cus
lactis
ssp.
cre-
moris
Leu- Cocci + − − d (−) + + − 0.2 2.0–4.0 Lac-
tate,
conos- di-
acetyl,
toc CO2
mesen-
teroides ssp. cremoris
Ther- Streptococcus
Cocci − thermophilus
+ + l (+) − − − 1.2 &- Lac-
mophilic lt;2.0 tate,
ac-
etalde-
hyde
Lacto- Rods − + + d (−) − − − 1.8 &- Lac-
lt;2.0 tate,
bacil- ac-
etalde-
lus hyde
bul-
gari-
cus

Lacto- Rods − + + dl − + − 2.0 &- Lactate

lt;2.0
bacil-
lus
hel-
veti-
cus

Lacto- Rods − + + d (−) − − +/− 1.8 2.0–4.0 Lactate

coc-
cus
lactis

> Read full chapter

FERMENTED MILKS | Other Relevant

Products
B. McKevith, C. Shortt, in Encyclopedia of Food Sciences and Nutrition (Second
Edition), 2003

Yogurts
Yogurt is traditionally produced from milk and/or cream through the action of two
particular strains of LAB – Lactobacillus bulgaricus and Streptococcus thermophilus.
Codex currently defines yogurt as a milk product obtained by fermentation using
these two specific strains with or without optional additions (such as milk powder,
other LAB, and sugar). In many countries (e.g., Sweden, France, Belgium, Portugal,
Spain, USA) legislation exists allowing only L. bulgaricus and S. thermophilus to be
used to produce yogurt. In the UK yogurt can be made using both of these strains or
just one of them. In other countries (e.g., Switzerland) strains such as L. acidophilus
and bifidobacteria may be used in addition to the traditional yogurt strains. It is
likely that the new Codex standard will build on the draft definition currently used
for mild yogurt (a product made from S. thermophilus and lactobacilli other than L.
bulgaricus). In Japan no separate legislation exists for yogurt and so it is not obligatory
that the two traditional strains are used, either collectively or separately. Similarly, in
Finland there are also no compositional regulations for any milk products.

> Read full chapter

Enzymatic conversions of starch

Piotr Tomasik, Derek Horton, in Advances in Carbohydrate Chemistry and Biochem-
istry, 2012

(ii) Fermentation
Fermentation of the saccharified material may involve various microorganisms, for
instance, Lactobacillus plantarum, Streptococcus themophilus, and/or Lactobacillus
bulgaricus; fermentation conditions are 25–45 °C for 3–20 h.2662,2663 A strain of S.
cerevisiae named amylolytic nuclear petite could increase the ethanol yield by over
54%.2664 Enzymes isolated from strains of Rhizopus and Aspergillus increased the
starch consumption up to 90.5%, along with a shortened fermentation period and
decrease of microbial contamination.2665 A cocultivated Schwanniomyces occidentalis
mutant and Saccharomyces cerevisae fermented sugars liberated from sorghum
starch very efficiently. The concentration of slurry could reach 28% without any
decrease in the yield of ethanol, although the time required for total fermentation
had to be extended with increasing concentration of the slurry.2666

Solid-phase fermentation is also possible.2667 The use of Schwanniomyces castellii

provided a solid-state method for production of ethanol in an aerobic–anaerobic
process.2668 In an aerobic cycle, growth of the microorganism and hydrolysis of
starch took place, and fermentation in a subsequent anaerobic cycle produced
ethanol. Foaming needed to be controlled.2669 When corn mashes are fermented,
the distillation is facilitated when the mash contains alpha amylase and glucoamy-
lase.2670 After cloning and expressing on either A. niger or Trichoderma resei, an A.
kawachi acid-stable alpha amylase in conjunction with glucoamylase, acting at pH
4.5, readily converted granular starch into ethanol. The amount of ethanol produced
is higher and the amount of residual starch is considerably decreased.2671

A common problem with fermentation is to assure sterility of the process and

avoiding contamination of the fermenting wort with undesired, deleterious mi-
croorganisms. In the production of ethanol from amylaceus material by use of
yeast, such contamination effects could be eliminated by the addition of 0.5% boric
acid. However, the yeast had to be acclimatized to that acid.2672 When NaF is used
for inhibiting the fermentation with yeast, hydrolysis begins to predominate over
fermentation.2673 Fluoride anions can protect Mucor mucedo from developing sec-
ondary strains.51 The fertility of pure cultures of microorganisms can be controlled
by pH, as shown with Rhizopus japonicus as used for fermenting corn starch.2674

Some Vibrio strains can be inhibited by glucose and sucrose.2675 In some in-
stances, heating and vigorous agitation,2676 and overnight steeping of the substrate
in 0.15% sulfuric acid2677 can be helpful. Apart from the common bactericides
used in the food industry, such as sorbic, benzoic, dehydroacetic (3-acetyl-2-hy-
droxy-6-methyl-1H-pyran-4-one), and 4-hydroxybenzoic acids, various microbio-
cides such as poly(hexamethylene biguanidine) and others can be employed.2678,2679
The latter method was used in the ethanol fermentation of raw starch with coim-
mobilized Aspergillus awamori, Rhizopus japonicas, and Zymomonas mobilis. Papain
and cysteine hydrochloride were also used.2680 However, in a fluidized-bed reactor
filled with immobilized Z. mobilis, hydrolyzed B-starch could be treated without
sterilization.2681

In the production of high-quality oriental wine, amylase is used to decompose

starchy material to make it more readily fermentable and, in general, to increase
the fermentation efficiency.2682 Degradation of starch with pullulanase and beta
amylase has been used in the brewing of beer.2683

> Read full chapter

LISTERIA | Properties and Occurrence

M.W. Griffiths, in Encyclopedia of Food Sciences and Nutrition (Second Edition),
2003

Interaction with Other Microorganisms

Growth of L. monocytogenes is inhibited in the presence of lactic acid bacteria,
including Lactococcus lactis, L. cremoris, Leuconostoc gelidum, and Lactobacillus
bulgaricus. The inhibition is partly due to production of bacteriocins by the lactic
acid bacteria but there is also a substantial pH effect. In most instances growth of
the pathogen is completely inhibited when the pH drops below 4.75.

Interactions between Listeria monocytogenes and other organisms have been de-
scribed. Coinoculation of milk or broth with L. monocytogenes and Pseudomonas spp.
resulted in either slight retardation of growth of the pathogen or no effect, depend-
ing on strains and temperature of incubation. However, a significant stimulation
of growth of L. monocytogenes in milks that had previously supported growth of
Pseudomonas spp. has been reported. This effect on growth is due to the production
of proteases by the pseudomonads. Accelerated growth of L. monocytogenes in whey
cultured with Penicillium camembertii compared with uncultured whey has been
reported.

> Read full chapter

Fibromyalgia
Leon Chaitow ND DO, ... Joanna Cameron, in Fibromyalgia Syndrome, 2010

Streptococcus thermophilus
(Rastall et al 2005). This is a transient (non-resident) bacterium of the human
intestine which, together with Lactobacillus bulgaricus, is a yogurt culture, also found
in some cheeses. It performs a number of useful roles; for example:

• Some strains produce natural antibiotic substances.

• They enhance the ability to digest milk and its products by producing the
enzyme lactase which is absent or deficient in almost half the adults on earth,
and in many children, especially if they are of Asian, African or Mediterranean
genetic stock.
• Because they produce lactic acid (this is the only streptococcus to produce lactic
acid, which it makes in even greater quantities than L. bulgaricus), they help
to create an environment that encourages colonization by the bifidobacteria
(they are therefore known as ‘bifidogenic’ bacteria) and L. acidophilus, as well
as helping to prevent colonization by undesirable micro-organisms.

> Read full chapter

The Gut-Brain-Skin Axis and Role of

Intestinal Dysbiosis in Acne Vulgaris:
Therapeutic Effects of Pro- and Prebiot-
ic Use
Bryan Tungland, in Human Microbiota in Health and Disease, 2018

13.4.2.2 Topical acne therapy with probiotics

Topical use of probiotic therapy in acne vulgaris has a long history, with first reports
published in 1912 by Peyri (1912), concluding that Lactobacillus bulgaricus may be
beneficial in acne and seborrhoea. However, it took until the late 20th century for
research to demonstrate through modern scientific methods that topical probiotic
therapy could be useful in acne by producing both antimicrobial activity and provide
direct anti-inflammatory properties. Some sphinglipids, such as phytosphingosine,
exert both anti-inflammatory and anti-microbial activities when used topically (-
Pavicic et al., 2007). These skin agents are low in acne vulgaris (Borodzicz et al.,
2016; Farwick et al., 2008; Pavicic et al., 2007; Yamamoto et al., 1995). Furthermore,
in winter months the sphinglipids are also low in the skin, which may account
for seasonal acne occurrence and dermatological visits during that seasonal period
(Hancox et al., 2004). In 1999 work by Di Marzio and others, which was later con-
firmed in studies by this group (Di Marzio et al., 2003, 2008), demonstrated that a
probiotic, Streptococcus thermophilus, increases ceramide sphingolipid production
when applied to skin as a cream for 7 days. Topical use of a probiotic providing
increased production of ceramide has relevance to acne as a few ceramide sphin-
golipids, particularly phytosphingosine, has been shown to be bactericidal against P.
acnes and have direct anti-inflammatory activity when used as a topical application
of 0.2%, reducing papules and pustules by 89% (Pavicic et al., 2007). Studies of the
topical use of probiotic Bifidobacterium longum strains are also in literature showing
that the probiotic can ameliorate skin inflammation by reducing the production
of substance P (Guéniche et al., 2010a,b), via mechanisms previous described. In
addition to the study by Pavici and others (2007), a more recent in vitro study
found that application of a topical Enterococcus faecalis lotion reduced inflammatory
lesions by over 50% in 8-weeks, as compared to a placebo group (Kang et al., 2009).
In further study, Al-Ghazzewi and Tester (2010) showed in another in vitro study
that the fermentable fiber, konjac glucomannan hydrolysates (GMH) use alone, or
in combination with probiotic lactic acid bacteria, can provide bactericidal activity
against P. acnes.

Currently, some cosmeceutical manufacturers, such as Glycologic Limited located

in Glasgow, Scotland, BIOK laboratorija (Lithuania), de Maniel (Granary House) in
Harpenden, England and Clinique in the U.K, have used probiotics or in combination
with GMH in masks, creams, facial mists or cleansing pads. These products have
been shown to benefit the skin via different means, such as producing a protective
barrier to modulate immune response, or produce anti-inflammatory mediators,
as well as produce bactericidal substances that minimize risk of P. acnes infection,
refer to company websites for further information. In 1999, Brook reported that in
patients suffering with acne and rosacea, probiotics applied topically can behave
as a protective shield to prevent skin cells from reacting to pathogenic bacteria
that can trigger an inflammatory cascade. This process, referred to as “bacterial
interference” is postulated to prevent colonization by competing with potentially
pathogenic microorganisms for binding sites, thereby competitively inhibiting in-
fection. Many probiotic bacteria are also known to produce and secrete bactericidal
substances, such as peptides and organic acids that decrease the risk of colonization
and infection by P. acnes. By example, Kang and others (2012) demonstrated in
vitro that the probiotic Lactobacillus reuteri, inhibited P. acnes by its production of
organic acids, which reduced skin pH. In addition, the oral microbe, Streptococcus
salivarius, produces a bacteriocin-like inhibitory substance (BLIS) that is capable of
retarding P. acnes growth (Bowe et al., 2006); also refer to section on S. salivarius
use in periodontal disease. As described in the periodontal section, these probiotic
microorganisms also regulate several inflammatory pathways that may impact skin
inflammatory responses (Cosseau et al., 2008). A more recent in vitro study by Lee
and others (2012) has shown that the probiotic strains, Bifidobacterium adolescentis
SPM0308 and B. longum SPM1207, isolated from healthy donor fecal samples,
reduce P. acnes viability by 84% and 75%, respectively.

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FOOD AND NUTRITIONAL ANALY-

SIS | Dairy Products
R.J. Marshall, in Encyclopedia of Analytical Science (Second Edition), 2005

Microbial methods
Any inhibitory substance in the milk will reduce the growth of test bacteria (e.g.,
Bacillus stearothermophilus, Streptococcus thermophilus, Bacillus subtilis, Sarci-
na lutea, Lactobacillus bulgaricus, Bacillus megaterium, etc.). The disk-assay plate
method uses a disk of absorbent paper that is impregnated with the milk sample to
be examined and placed on the surface of an agar plate preinoculated with a suitable
test organism. After incubation, the growth of the bacteria is seen as turbidity in the
agar. The presence of inhibitory substances such as antibiotics which diffuse into
the agar is indicated by a clear zone around the edge of the disk. The inhibitor may
be identified by adding an agent that blocks its action. For example, the effect of
 penicillin is prevented by the addition of penicillinase, while p-aminobenzoic acid
blocks sulfonamides. With such substances in the growth medium, the antibiotic no
longer prevents the test organism from growing. Commercially available methods
include Intertest, Delvotest, and TTC.

On the farm, a test kit may be used. The test organism is inoculated in tablet form
into liquid medium in a test tube with a sample of the milk. An indicator in the
buffer changes color if the bacteria are able to grow (i.e., no antibiotic is present).
The Charm analyzer can be used for several different tests (Figure 2).

Figure 2. Charm Sciences ROSA® portable incubator/analyzer (Reproduced with

permission from Charm Sciences Inc.).

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