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Beneficial Microbes, September 2012; 3(3): 245-247

Koen Venema

http://www.wageningenacademic.com/doi/pdf/10.3920/BM2012.x003 - Friday, November 18, 2016 5:10:47 AM - IP Address:88.249.75.107

TNO Quality of Life, Department of Biosciences, P.O. Box 360, 3700 AJ Zeist, the Netherlands

BOOK REVIEW
Lactic acid bacteria microbiological and functional aspects (4th edition)
S. Lahtinen, A.C. Ouwehand, S. Salminen and A. von Wright (editors)
List price: 162 Euro
Hardcover: 798 pages
Publisher: CRC Press (2011)
ISBN: 978-1-4398-3677-4
eISBN: 978-1-4398-3678-1

Introduction
With 798 pages this is a very comprehensive book on the
microbiological and functional aspects of lactic acid bacteria
(LAB). The fourth edition follows seven years after the
third and due to accumulation of new findings practically
all chapters have been extensively rewritten. In addition to
purely scientific chapters, the regulatory framework related
to the safety and efficacy of LAB, particularly probiotics, has
also been added. The book contains 33 chapters, roughly
divided, although not indicated as such by the editors,
into 4 parts: characterisation of LAB, application of LAB
in food fermentations, probiotics and health benefits, and
safety and efficacy.

Genetics and characterisation of lactic acid bacteria

The first part consists of nine chapters, that deal with
the different genera that make up the LAB (including
bifidobacteria), their genetics and mechanisms to prevent
or overcome bacteriophage infections. It starts with an
introductory chapter describing the current taxonomic
position of LAB, their carbohydrate fermentation pathways,
bioenergetics, solute transport and proteolytic system.
The homolactic and heterolactic fermentation are briefly
highlighted. Lactic acid or a mixture of lactic acid and
other metabolites (ethanol or acetate) are produced, upon
carbohydrate fermentation, respectively, which is what has
given these bacteria the general name LAB. Alternative fates
of pyruvate (an intermediate in carbohydrate fermentation)

are discussed as well, leading e.g. to the production of the

flavour compounds diacetyl and acetoin. The generation
of energy through ATP production and proton motive
force are discussed. Both forms of energy are used for
solute transport into the cell, to allow for accumulation of
building blocks for biomass. Also the proteolytic system,
that liberates amino acids and short peptides from (milk)
protein, is highlighted. It is interesting to see the progress
in this field, in which I started my scientific career in 1990.
The second chapter reviews the genetics of LAB. With the
sequencing of many genomes of LAB in the past decade, this
is a chapter that has been completely rewritten compared
to previous editions. Comparative genomics allowed for the
identification of a core genome and the pan-genome of LAB.
Niche-specific genes have been identified. Plasmid biology
has always been important in LAB, also as a means of gene
transfer. In addition, insertion elements, transposons and
group II introns are discussed.
Chapters 3 to 8 each deal with a genus (or multiple genera
for Chapter 6) within the LAB, where bifidobacteria have
been included as well, although they are strictly spoken not
part of LAB. Enterococci (Chapter 3), lactococci (Chapter
4), lactobacilli (Chapter 5), pediococci, Leuconostoc,
Weissella, Carnobacterium and related genera (Chapter
6), streptococci (Chapter 7) and bifidobacteria (Chapter 8)
are discussed. In 20 to 25 pages per chapter, the highlights
are given for each genera, including origin, phylogeny and
taxonomy, typical ecological niches, potential uses, genomic

ISSN 1876-2833 print, ISSN 1876-2891 online, DOI 10.3920/BM2012.x003245

http://www.wageningenacademic.com/doi/pdf/10.3920/BM2012.x003 - Friday, November 18, 2016 5:10:47 AM - IP Address:88.249.75.107

Book review

features and how these can be used for functionalities of

LAB, and metabolic capacity. For streptococci also the
classification of pathogenic streptococci is given, and
the distinction between aggressors, opportunists and
protectors are reviewed. The last chapter of this section
deals with bacteriophages and anti-phage mechanisms.
Bacteriophages are the principal cause of starter culture
failure. LAB have developed several mechanisms to
overcome this phage attack, amongst which prevention
of phage adsorption, inhibition of DNA entry, restriction/
modification systems, abortive infection, and the clustered
regularly interspaced short palindromic repeat (CRISPR)Cas system, which have been proposed to function as small
interfering RNAs, eventually promoting degradation of
phage RNA or DNA.

Application of lactic acid bacteria in food fermentation

This subject is composed of Chapters 10, 12-14 and 17.
Chapter 10 reviews LAB in vegetable fermentations, and
includes examples of sauerkraut, cucumber, olive, kimchi
and vegetable juices. Furthermore, the functionality
of LAB from fermented vegetables is discussed, with a
focus on kimchi. Interesting, a section is included on
possible negative effects of fermented vegetables, such as
the presence of nitrate and nitrite and biogenic amines.
Chapter 12 deals with cereal based products. Lactic
acid fermentation in wet milling is discussed, as well as
sourdough fermentation. Furthermore, there is emphasis on
traditional LAB-fermented cereal-based foods other than
bread, which are staple foods in parts of Africa, Asia and
South America. This is further reviewed for African food
fermentations in Chapter 14, where continent and regional
specific products are introduced. Examples discussed in
more detail are maize, sorghum and millet, starchy root crop
and animal protein fermentations. Chapter 13 is on meat
fermentation. The biodiversity of LAB in meat fermentation
is discussed, as well as the functionality of LAB in these
fermentations. Their main role is in flavour development
and biopreservation, through acidification and bacteriocin
production. In terms of safety, biogenic amine production
and the occurrence of antibiotic resistant isolates are
reviewed. Chapter 17 deals with grape fermentation. This is
an example of LAB being contaminants in food processing,
as LAB fermentation leads to spoilage of wine. Negative
effects are production of a multitude of toxic compounds
as well as exopolysaccharides. For whatever reason, there
is no chapter on dairy fermentation. That is an omission,
as the interest in LAB fermentations originally stems from
this area. It is also where my own scientific roots are...

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Use of lactic acid bacteria as probiotics: health benefits

Chapters 11, 15, 16, 18-21 and 23-30 deal with the beneficial
effects of LAB, or probiotics. Chapter 11 starts with the
current challenges for probiotics in food. After a brief
section on the history of use of LAB, challenges with respect
to maintaining fermentation cultures, probiotic origin and
diversity, assuring survival and activity of probiotics and
dealing with the endogenous microbiota are discussed
elegantly. Also challenges to establish benefits of probiotics
for humans and safety issue are addressed. Chapter 15 deals
with antimicrobial components produced by LAB. The
chapter reviews the various antimicrobial produced by LAB,
such as organic acids, reuterin and reutericyclin, antifungal
peptides and bacteriocins. As I did my PhD on the latter,
this chapter is close to my heart. The various bacteriocins
produced by LAB are elegantly reviewed, as well as their
targets and mode of action, and appearance of resistance
or adaptation to these bacteriocins. Applications are also
briefly mentioned. It is unfortunate that the amino acid
sequences of the bacteriocins given in Table 15.2 are hardly
readable due to the shading of the amino acid residues.
Chapter 18 describes the challenges of LAB in foods and
supplements. Factors affecting stability are discussed, and
include freeze-drying, water activity, oxygen concentration
and chemical composition of the (food) matrix. Methods
for determining stability are also reviewed. Chapter 16 is
the first of a number of chapters describing (potential)
health benefits of probiotics, and describes the possibility
(or perhaps probability) of LAB to influence atherosclerosis,
either directly or through modulation of the gut microbiota.
Mechanisms could be through long-term prevention or
alleviation of inflammation, e.g. through restoration of
gut barrier function, thereby reducing translocation of
endotoxins or other proinflammatory agents. Chapter
19 describes LAB in the gut. The focus is on lactobacilli
and streptococci. The latter are dubbed the forgotten
LAB in the gastro-intestinal tract (GIT) by the authors,
as there is relatively little known about these bacteria in
the gut. Streptococci are primarily of importance in the
small intestine. The authors discuss whether lactobacilli
are true residents in the gut or transient passengers from
food. However, they also describe successful adaptation of
lactobacilli in the gut, so at least some seem true residents.
Chapter 20 describes LAB in oral health. Although most
LAB in the oral cavity are linked to dental caries and
periodontal disease as elegantly reviewed in the chapter,
some probiotics are being studied for their potential benefit
to inhibit or displace the pathogenic microorganisms.
The effect of probiotics on immune function is reviewed
in Chapter 22. This is one of the biggest chapters in the
book, but the subject would warrant almost a complete
book on its own. Effects of probiotics on the mucosal and
systemic immunity have been described. They can be (this

Beneficial Microbes 3(3)

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is strain dependent) beneficial in disease resistance and

immunoinflammatory disorders. The former includes
boosting resistance to infections in the GIT, respiratory and
urogenital tract and cancer. The latter relates to reduction in
allergy or inflammatory bowel disease (IBD), or diminishing
the symptoms of irritable bowel syndrome (IBS).
Mechanisms of how probiotics may exert these multitude
of effects are reviewed briefly. Chapter 23 discusses the
clinical evidence for the GIT benefit of probiotics. There is
some overlap between other chapters that are specific for
certain disorders, e.g. with the next chapter on studies of
probiotics with infants and children. Both describe diarrhea,
Heliobacter pylori infection, necrotising enterocolitis, IBD
and IBS. Chapter 25 describes the endogenous microbiota
in the urogenital tract, as well as the use of probiotics to
conquer dysbiosis due to bacterial vaginosis, vulvovaginal
candidiasis and urinary tract infections. The potential of
LAB to aid in reduction of high blood pressure is reviewed
in Chapter 26. The mechanism is thought to occur due to
the production of bioactive, antihypertensive peptides IlePro-Pro and Val-Pro-Pro from the milk protein casein. The
next 3 chapters describe the beneficial effects of probiotics
for animals. This is usually largely overlooked in books
reviewing LAB or probiotics, and it adorns the editors
that they have included these chapters. The first (Chapter
27) is on companion animals, and essentially describes
similar disorders in pets as those that occur in humans.
The effects of probiotics on these are reviewed. The second
(Chapter 28) describes the use of probiotics in aquaculture.
Besides LAB, the chapter describes the use of Bacillus
(spores), Clostridium. tyrobutyricum and Saccharomyces
cerevisiae in commercial probiotic products, either alone
or in combination with LAB. The past decade has seen
an explosion in research on the benefits of probiotics in
aquaculture. The third chapter (Chapter 29) describes the
use of probiotics for farm animals. Besides pigs, ruminants
and poultry, the use of probiotics for other farm animals like
honeybees, foxes, minks, rabbits and horses are reviewed.
The application for each animal has a different focus, with
reduction of diarrhea and benefits post-weaning for pigs,
reduction of diarrhea and infections, and reduction in
ruminal acidosis and methane production in ruminants,
and increase in meat quality and egg production in poultry.
The last chapter in this imaginary section deals with health
effects of non-viable probiotics. Although this seems a
contradictio in terminis, as the definition of probiotics
is all about live bacteria, dead bacteria (which are no
doubt present in food products or supplements) might
also have a beneficial effect on health. For instance, the
mechanisms of LAB aiding in lactose digestion depends
on lysis of beta-galactosidase-containing cells, to release
the lactase in the GIT lumen, where it can help in digestion
of ingested lactose. Other proposed mechanisms by which
dead bacteria, or bacterial fragments may have an effect
are reviewed as well.

Beneficial Microbes 3(3)

Book review

Safety and efficacy of lactic acid bacteria

This is dealt with in Chapters 21, and 31-33. Chapter 21
describes some considerations for safety of novel probiotics.
These include the use of enterococci (some of which may
harbor pathogenicity genes), spore-forming probiotics,
antibiotic resistant strains and the potential of probiotics
to infect (especially immunocompromised) hosts. Chapter
31 continues on the safety and efficacy of probiotics. It
reviews the safety assessment for use of probiotics in
humans and as microbial feed additives. With respect to
efficacy reference is made to the health claim regulation
in the EU, and steps that are required to obtain claims on
probiotic products. Specific challenges in human trials are
proper characterisation of the strain used, viability of the
probiotic, and properly designed and powered studies to
show the efficacy of probiotics in the target group (usually
healthy human volunteers). So far, EFSA has not approved
a single dossier on probiotics, due to flaws in the evidence.
Health claim regulation in Asian countries is reviewed in
Chapter 32. This includes an overview of the differences
(and similarities) in countries like China, India, Japan,
Korea, Indonesia, Malaysia, the Philippines, Singapore,
Thailand and Taiwan. Unlike in the EU, the guidelines
for probiotic products in Asia are not harmonised and
each country has its unique regulation, with some being
restrictive and others quite elaborative. The last chapter
in the book describes the regulation of probiotics in
South America. Except for Brazil, no specific regulation
on probiotics or probiotic health claims exist in South
America. In other countries than Brazil, food regulations
are defined by the Regional Trade Agreement MERCOSUL,
but this does not yet include legislation on probiotics. It is
unfortunate that information on regulation in the USA and
Canada is missing. But perhaps I may revert the readers
attention to an upcoming paper in the 4th issue of this year
of Beneficial Microbes by Diane Hoffmann entitled Health
claim regulation in the USA and the EU of probiotics: is
there a middle way?

Conclusion
Overall this is a very extensive survey of the current
knowledge on science and technology of probiotics. Despite
some omissions such as use of LAB dairy fermentation
and the North American regulatory framework, this is
a nice reference for the current state-of-the-art on LAB
and probiotics.

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