International Journal of Food Microbiology 154 (2012) 87–97

Contents lists available at SciVerse ScienceDirect

International Journal of Food Microbiology

j ournal homepage: www . elsevier . com/locate/ijfoodmi cro

Review

Food fermentations: Microorganisms with technological beneficial use

a, 1 b c d e, 2
François Bourdichon , Serge Casaregola , Choreh Farrokh , Jens C. Frisvad , Monica L. Gerds , Walter P.
f g h i j k
Hammes , James Harnett , Geert Huys , Svend Laulund , Arthur Ouwehand , Ian B. Powell , Jashbhai B. Prajapati
l m n o p q
, Yasuyuki Seto , Eelko Ter Schure , Aart Van Boven , Vanessa Vankerckhoven , Annabelle Zgoda , Sandra
r d,
Tuijtelaars , Egon Bech Hansen
1Danone Research, RD128, 91 767 Palaiseau Cedex, France
2 INRA, UMR 1319 Micalis, CIRM-Levures, AgroParisTech 78850 Thiverval-Grignon, France

3 CNIEL, 42, rue de Chateaudun, 75314 Paris Cedex 09, France

4 Department of Systems Biology, Technical University of Denmark, Søltofts Plads B. 221, DK-2800 Kgs. Lyngby, Denmark

5 Cargill Texturizing Solutions, 620 Progress Avenue, Waukesha, WI, 53187-1609, United States

6 Institut für Lebensmittelwissenschaft und Biotechnologie, University of Hohenheim, Garbenstraße 21, D-7000 Stuttgart 70, Germany

7 Fonterra Co-operative Group Ltd., Private Bag 11029, 4442 Palmerston North, New Zealand

8 BCCM/LMG Bacteria Collection & Laboratory of Microbiology, Faculty of Sciences, Ghent University, K.L. Ledeganckstraat, 35, B-9000 Gent, Belgium

9 EFFCA, European Food & Feed Cultures Association, Bd. Saint Michel 77-79, B-1040, Brussels, Belgium & Chr Hansen A/S, Boge Alle 10-12, DK-2970 Horsholm, Denmark

10 Danisco Innovation, Sokeritehtaantie 20, FIN-02460 Kantvik, Finland

11 Dairy Innovation Australia, 180 Princes Highway, Werribee, Victoria 3030, Australia

12 Anand Agricultural University, Anand 388 110 Anand, Gujarat State, India

13 Milk Science Research Institute, Megmilk Snow Brand Co., Ltd., 1-1-2 Minamidai, 350-1165 Kawagoe, Saitama, Japan

14 Laboratory & Quality Services FrieslandCampina, PO Box 226, 8901 MA Leeuwarden, Netherlands

15 CSK Food Enrichment B.V., P.O. Box 225, NL-8901 BA Leeuwarden, Netherlands

16 University of Antwerp, Vaccine & Infectious Disease Institute (Vaxinfectio), Campus Drie Eiken, Universiteitsplein 1, 2610 Antwerp, Belgium

17 Groupe Lactalis, Le Fromy, 35240 Retiers, France

18 International Dairy Federation, Silver Building, Boulevard Auguste Reyers 70/B, 1030 Brussels, Belgium
or “general recognition of safety”. Authoritative lists of
microorganisms with a documented use in food have
therefore come into high demand. One such list was
article info abstract published in 2002 as a result of a joint project between the
International Dairy Federation (IDF) and the European
Article history: Microbial food
Food and Feed Cultures Association (EFFCA). The “2002
Received 9 August 2011 cultures have
Received in revised form 1 December 2011 Accepted 22 December 2011 IDF inventory” has become a de facto reference for food
directly or
Available online 31 December 2011 cultures in practical use. Howev-er, as the focus mainly
indirectly come
was on commercially available dairy cultures, there was an
under various
Keywords: unmet need for a list with a wider scope. We present an
regulatory
Lactic acid bacteria updated inventory of microorganisms used in food
Fungi frameworks in the
fermentations covering a wide range of food matrices
Starter cultures course of the last
(dairy, meat, fish, vegetables, legumes, cereals, beverages,
History of use decades. Several
and vinegar). We have also reviewed and updated the
Fermentation of those
taxonomy of the microorganisms used in food
Food microbiology regulatory
fermentations in order to bring the taxonomy in agreement
frameworks put
with the current standing in nomenclature.
emphasis on “the
history of use”, © 2011 Elsevier B
“traditional food”, All rights reserved
Contents 2.1. Definition of MFC . . . . . . . . . . . . . . . . . . . . . . .
1. Introduction . . . . . . . . . . . . . 2.2.
. . . . .Definition . .“history
. . . . . of . . . . of
. .use”
. . .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. .. ..
2. Regulatory systems and legal terms . 2.3.
. . . . US
. . regulatory
. . . . . . environment.
. . . . . . . . . .. . . .. . . .. . . .. . . .. . . .. . . .. . . .. . . .
 2.4. European regulatory environment . . . . . . o. . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
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 88 F. Bourdichon et al. / International Journal of Food Microbiology 154 (2012) 87–97

3. Scientific criteria for evaluation of MFC. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89

3.1. Taxonomy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89
3.2. Undesirable properties of MFC . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90
3.2.1. Opportunistic infections . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90
3.2.2. Toxic metabolites and virulence factors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90
3.2.3. Antibiotic resistance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90
4. Inventory of microbial species used in food fermentations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90
4.1. Bacteria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91
4.1.1. Actinobacteriaceae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91
4.1.2. Firmicutes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91
4.1.3. Proteobacteriaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
4.2. Fungi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
4.2.1. Yeasts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
4.2.2. Filamentous fungi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
5. Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94
6. Acknowledgments and disclaimer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94
Appendix A. Supplementary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94

reviewing the microbial species used

in food fermentations, we also review
the regulatory systems, some of the
1. Introduction contributions to the food legal terms, and scientific criteria
fermentation. This final step is not relevant for microbial food cul-tures
Preservation of food including the use of fermentation of otherwise without ambiguity as taste and flavor (MFC). Accordingly, we have
perishable raw materials has been used by man since the Neolithic pe-riod preferences can be quite different, structured the review to cover:
(around 10000 years BC) (Prajapati and Nair, 2003). The scientific rationale and what some would consider
behind fermentation started with the identification of micro-organisms in 1665 spoilage can be regarded as desirable 1• Regulatory systems and legal terms
by Van Leeuwenhoek and Hooke (Gest, 2004). Pas-teur revoked the by others. We intend to be
“spontaneous generation theory” around 1859 by elegantly designed conservative, and the current list is
2• Scientific criteria
experimentation (Wyman, 1862; Farley and Geison, 1974). The role of a sole therefore less than exhaustive and it 3• Inventory of microbial species in
bacterium, “Bacterium” lactis (Lactococ-cus lactis), in fermented milk was cannot be considered definitive. An food fermentations.
shown around 1877 by Sir John Lister (Santer, 2010). Fermentation, from the updat-ing process following the
Latin word fervere, was defined by Louis Pasteur as “La vie sans l'air” (life scientific rationale detailed in the 2. Regulatory systems and legal
without air). From a biochem-ical point of view, fermentation is a metabolic present article will be established and terms
process of deriving ener-gy from organic compounds without the involvement of hosted by IDF. The criteria chosen
an exogenous oxidizing agent. Fermentation plays different roles in food for including species on the list are: 2.1. Definition of MFC
processing. Major roles considered are:
It is remarkable that MFC have
1• Inclusion not been defined legally. To allevi-ate
(1) Preservation of food through formation of inhibitory metabo-lites such as o Microbial species with a this, EFFCA has proposed the
organic acid (lactic acid, acetic acid, formic acid, propionic acid), ethanol, documented presence in following definition: “Microbial food
bacteriocins, etc., often in combina-tion with decrease of water activity (by fermented foods cultures are live bacteria, yeasts or
drying or use of salt) (Ross et al., 2002; Gaggia et al., 2011). molds used in food production”. MFC
preparations are formulations,
(2) Improving food safety through inhibition of pathogens (Adams and Mitchell,
2• Exclusion consisting of one or more micro-bial
2002; Adams and Nicolaides, 2008) or removal of toxic compounds (Hammes species and/or strains, including
o Lack of documentation for any
and Tichaczek, 1994). media components carried over from
desirable function in the fermen-
(3) Improving the nutritional value (van Boekel et al., 2010; Poutanen et al., 2009). tation process the fermentation and components
which are necessary for their
(4) Organoleptic quality of the food (Marilley and Casey, 2004; Smit et al., 2005; o The species is a contaminant survival, storage, standardization, and
and/or does not harbor any
Lacroix et al., 2010; Sicard and Legras, 2011). to facilitate their application in the
relevant metabolic activity
o The species is undesirable in food production process.
An authoritative list of microorganisms with a documented use in food was
food for scientifically
established as a result of a joint project between the Inter-national Dairy
documented reasons.
Federation (IDF) and the European Food and Feed Cul-tures Association
(EFFCA). This list was published in 2002 by Mogensen et al. (2002a, 2002b).
With the current review, we have undertaken the task to establish a revised and Microorganisms conferring a
updated inventory of microorganisms with a history of use in fermented foods. health benefit to the host (FAO and
We have chosen a pragmatic approach for updating the inventory by creating a WHO, 2002) are thus included if
“gross list” consisting of the 2002 inventory supplemented with ad-ditions they are part of a culture used in a
suggested by the National Committees of IDF and members of EFFCA, as well food fermentation process, whereas
as additions found by searching the scientific literature for documentation of we have decided not to include
food fermentations with emphasis on microbial associations and food matrices microbial species of probiotic strains
not initially covered. From this greatly expanded list we then critically reviewed only used in supplements or over the
the literature for each spe-cies in order to maintain only microbial species counter (OTC) products.
making desirable As part of the process of
 F. Bourdichon et al. / International Journal of Food Microbiology 154 (2012) 87–97 89

2.2. Definition of “history of use” Lists of microorganisms to be a fast track for species
and microbial derived for which there is a sufficient
The concept of “history of safe use” has ingredients used in foods can body of knowledge that all
appeared recently in reg-ulations and in safety be found at the FDA web site strains within a species are
assessment guidance. One definition of “history (FDA, 2001). As a result of the assumed to be safe. This
of safe use” proposes “significant human different ways to obtain presumption may be qualified
consumption of food over several generations GRAS, the FDA lists of GRAS by some restrictions such as
and in a large, genetically diverse population for substances are not expected to the absence of specific
which there exist adequate toxicological and characteristics (for example
include all substances, nor all
allergenicity data to pro-vide reasonable the absence of trans-missible
pre-1958 natural, nutri-tional
certainty that no harm will result from antibiotic resistance, absence
substances. For a more
consumption of the food” (Health Canada, 2003). of food poisoning toxins, ab-
comprehensive US regulatory
In order to evaluate the history of safe use of a sence of surfactant activity,
update on MFC, we refer to a
microorganism, it is necessary to document not and absence of enterotoxic
recent review by Stevens and
just the occurrence of a microorganism in a activity). The QPS list covers
O'Brien Nabors (2009).
fermented food product, but also to provide only selected groups of
evidence whether the presence of the microorganisms which have
microorganism is beneficial, fortuitous, or been referred to EFSA for a
2.4. European regulatory
undesired. formal assessment of safety
environment
(Anon, 2005; Leuschner et al.,
2010). Seventy-nine species of
In the European Union, the
2.3. US regulatory environment microorgan-isms have so far
MFCs are considered
been submitted to EFSA for a
ingredients and must satisfy
In the United States, food and substances the legal requirements of safety assessment; the list is
used in food are regulat-ed according to the Food regulation EC no. 178/2002. updated annually (EFSA,
Drug and Cosmetic Act (1958), in which the Consequently, 2007, 2008, 2009, 2010). The
the
status of Generally Recognized As Safe (GRAS) responsibility for the safe use ab-sence of a particular
was introduced (FDA, 2010). Accordingly, a of microorganisms in food organism from the QPS list
GRAS substance is generally recognized, among should be ensured by food does not necessarily imply a
qualified experts, as having been adequately manufacturers. risk associated with its use.
shown to be safe under the conditions of its Individual strains may be safe,
In 2007, the European
intended use. A substance recognized for such but this cannot be ascertained
Food Safety Authority (EFSA)
use prior to 1958 is by default GRAS (like food from the existing knowledge of
introduced “Qualified
used in the EU prior to May 15, 1997, not being the taxonomic unit to which it
Presumption of Safety” (QPS)
Novel Food) (Anon, 1997, ILSI Europe Novel belongs. Another reason for a
for a premarket safety as-
Food Task Force, 2003). MFC are an integral species not being on the list
sessment of microorganisms
part of traditional fermented foods. As a could be that EFSA has not
used in food and feed
significant number of people have consumed been asked to assess the safety
production. QPS is applicable
these foods for many centuries before 1958, the of any strains of the species. A
to food and feed additives,
fermenting microor-ganisms of these products recent review (Herody et al.,
food enzymes and plant pro-
can be said to be GRAS. If a substance (mi- 2010) gives a thorough
tection products (Anon, 2005).
croorganism) is GRAS for one food usage, it is description of the European
The QPS system was proposed
not necessarily GRAS for all food uses. It is the regulatory envi-ronment for
to har-monize approaches to
use of a substance rather than the substance itself microbial food cultures.
the safety assessment of
that is GRAS, as the safety determination is
microorganisms across the
always limited to its intended conditions of
various EFSA scientific panels.
usage. When microorganisms with a safe his-tory Denmark is the nation with
The QPS approach is meant
in food are employed for a different use or at a the first national legislation
significantly higher dosage, a GRAS (since 1974) that specifically
determination for these new usages is needed. requires safety approval of
MFC. More than 80 species
There are three ways to obtain GRAS status used in 14 different food
for an MFC: categories have been approved
and published at the Danish
1. A GRAS notification where a person/company Veterinary and Food
informs FDA of a de-termination that the usage Administration web site
of a substance is GRAS and followed by the (Anon, 2009). In 2010, the
receipt of a no-objection letter from FDA regulation was changed.
2. A GRAS determination made by qualified Approval is no longer needed,
experts outside of the US government and the but a notification of a new
result is kept by the person/company behind the species or a new ap-plication is
determination still required before it can be
marketed in Denmark. This
3. GRAS due to a general recognition of safety,
topic has also recently been
based on experience from common use in food investigated by Germany
by a significant number of people before 1958. (Vogel et al., 2011).
 3. Scientific criteria for evaluation of MFC

3.1. Taxonomy

Taxonomy and systematics constitute the

basis for the regulatory frameworks for MFCs. It
is thus somewhat unfortunate that the defi-nition
of microbial species as a taxonomic unit lacks a
theoretical basis (Stackebrandt, 2007). For this
reason, we briefly outline the cur-rent status of
bacterial and fungal taxonomy.
In the third edition of Prokaryotes
(Stackebrandt, 2006), Stackeb-randt proposes a
prokaryotic species to be defined by:

1• a phylogenetic component given as “the

smallest diagnosable clus-ter of individual
organisms within which there is a parental
pattern of ancestry and descendents” (Cracraft,
1983),
and

1• a taxonomic component given as “a group of

related organisms that is distinguished from
similar groups by a constellation of significant
genotypic, phenotypic, and ecological
characteristics.” (Colwell, 1970).

In general, a polyphasic approach to

taxonomy is recommended in bacteriology
(Vandamme et al., 1996). In practice, this means
that a bacterial species is represented by a type
strain with strains showing a high degree of
phenotypic and/or genotypic similarity to the
type strain regarded as belonging to the same
species. Whilst objective measures of relatedness
have been proposed (such as percentage ge-
nome hybridization or sequence similarity), there
is no simple defini-tion of the species as a
taxonomical unit.

As a basis for the current taxonomy of

prokaryotes we have used the classification of
the International Committee on Systematics of
Prokaryotes (ICSP— http://www.the-icsp.org/)
and available publica-tions in International
Journal of Systematic and Evolutionary Microbi-
ology (IJSEM— http://ijs.sgmjournals.org/). The
Taxonomic Outline of the Bacteria and Archea
(TOBA— http://www.taxonomicoutline.org/) in
its release 7.7 of March 6, 2007, and the
amended lists of bacterial names (Skerman et al.,
1989) were used as reference.
90 F. Bourdichon et al. / International Journal of Food Microbiology 154 (2012) 87–97
http://www.doctorfungus.org/).
In fungal taxonomy different concepts to important marker of serious
define microbial species are used without3.2. Undesirable properties of underlying disease” (Husni et
reaching a final consensus between the numerousMFC al., 1997). Sporadic infections
relationships observed between phenotypic and have been reported in immuno-
molecular methods (Guarro et al., 1999; Although they have been compromised pa-tients. The
Hawksworth, 2006). Several definitions haveused since ancient times in underlying problems have
been used to describe the yeast domain. Yeasts fermentation processes without mainly been central venous
may be defined as being ascomycetous or any identified major concern, catheter (CVC) in place,
basidiomycetous fungi that reproduce vegeta- recent discovery of rare events metabolic disorders, organ
tively by budding or fission, with or withoutof adverse effects caused by failure, or invasive procedures
pseudohyphae and hy-phae, and forming sexualmicroorganisms in fermented such as dental work (Axelrod
states that are not enclosed in fruiting bodies foods raise uncertainty about et al., 1973; Liong, 2008).
(Boekhout and Robert, 2003). Phylogeneticthe level of risk, depending Infections by other bacterial
studies have now clearly shown the clustering of either on the food matrix or the species used as MFC are also
the hemiascomycetous yeasts forming a single susceptibility of the host extremely rare (Horowitz et
clade within the ascomycota, the other yeasts (Gasser, 1994; Miceli et al., al., 1987; Barton et al., 2001;
belonging to the basidiomycetes (Hibbett et al.,2011). Mofredj et al., 2007;
2007). Leuschner et al., 2010).

Yeasts used to be commonly identified3.2.1. Opportunistic infections Infections with the

phenotypically, but they are now identified from Commensal bacteria have commonly used yeast and
diagnostic sequences (Daniel and Meyer, 2003).been described to cause mold species are rare events as
Techniques using molecular biology are seen as infections in patients with well (Enache-Angoulvant and
an alternative to tradi-tional methods since they underlying disease (Berg and Hennequin, 2005). Most of the
analyze the genome independently of theGarlington, 1979; Berg, 1985, infections are due to
physiological characteristics, which may vary1995). Owing to its natural opportunistic pathogens not
within the species (Boekhout and Robert, 2003;presence in different sites of recog-nized as MFC and affect
Fernández-Espinar et al., 2006; Kurtzman et al.,the human body and in immuno-compromised patients
2011). Molecular techniques are morefermented food products, the and hospital-ized patients
reproducible and faster than the conventional genus Lactobacillus has gained (Winer-Muram, 1988; Jacques
methods based on physiological andparticular attention. and Casaregola, 2008; Miceli
morphological characteristics. Furthermore,Lactobacillus infections occur et al., 2011).
these techniques prevent misclassification ofat a very low rate in the
species on the basis of their sexuality. In some generally healthy population—
cases, ribosomal D1/D2 sequence comparisonestimated 0.5/1 million per 3.2.2. Toxic metabolites and
cannot discriminate be-tween species, and moreyear (Borriello et al., 2003; virulence factors
discriminating sequences have to be used in Bernardeau et al., 2006). As Biogenic amine formation
parallel (Jacques and Casaregola, 2008). Overall,stated in two reviews of in fermented foods by lactic
a combination of prov-en loci such as ACT1, Lactobacillus infections: acid bacte-ria (LAB) has
RPB1 and RPB2, and Elongation Factor genes“Underlying disease or immu- recently been reviewed (Spano
are suitable, if they are included in a multilocus nosuppression are common et al., 2010). Following food
analysis. Genomic studies have greatly helped features in these cases, poisoning outbreaks (Sumner
the search for yeast identification markerswhereas infection in et al., 1985), metabolic
(Casaregola et al., 2011; Aguileta et al., 2008). previously healthy humans is pathways have been elucidated
extremely rare” (Aguirre and
(Straub et al., 1995) and
The variability in the fungal kingdom is evenCollins, 1993). “Lactobacillus
screening procedures proposed
wider considering molds: estimations are bacteraemia is rarely fatal per
to limit the level of production
currently rated around 100000 species. It is se but serves as an
(Bover-Cid and Holzapfel,
thought that there are between 700000 to 1.5
1999; Bover-Cid et al., 2000).
million species that are yet to be identified and
classified (McLaughlin et al., 2009). Recently, a The presence of mycotoxin
comprehensive monograph on all the genera of genes also raises safety
anamorphic fungi (hyphomycetes, fungi concerns, al-though the level
imperfecti, deuteromycetes, asexual fungi) was of expression within fermented
published (Seifert et al., 2011). This book, food is very unlikely to cause
together with the Dictionary of the Fungi (Kirk any health hazard
et al., 2008), gives an overview of the taxonomic (Barbesgaard et al., 1992).
sta-tus of all genera of filamentous fungi. Within fungi, the potential for
antibiotic production is also an
As for the current taxonomy of fungi, we undesired property.
have used the references and documentation The occurrence of
provided by the International Commission on the virulence traits should not be
Taxon-omy of Fungi (ICTF) on their website ( present in micro-organisms
http://www.fungaltaxonomy.org/) and the used in food fermentation. A
Mycobank initiative (Crous et al., 2004), as well specific risk assessment should
as expert groups on invasive fungal infections
be conducted on strains
and taxonomic issues (Mycoses Study Group—
presenting these undesirable
proper-ties, even if they belong to a species with
a long history of use (Semedo et al., 2003a,
2003b).

3.2.3. Antibiotic resistance

The emergence and spread of antibiotic
resistance is a major glob-al health concern. The
on-going Codex ad hoc intergovernmental task
force on antimicrobial resistance is focused on
the non-human use of antimicrobials.
Microorganisms intentionally added to food and
feed for technological purposes have not been
shown to aggravate the problem of spreading
antibiotic resistant pathogens (Anon, 2001).
Intrinsic resistance or resistance that is
caused by mutation in an indigenous gene not
associated with mobile elements would repre-
sent a very low risk of dissemination (Saarela et
al., 2007). Acquired antibiotic resistance genes,
especially when associated with mobile genetic
elements (plasmids, transposons), can be
transferred to path-ogens or other commensals
along the food chain, from within the product
until consumption (FEEDAP, 2005, 2008;
Nawaz et al., 2011).
The role of MFC in the spread of antibiotic
resistance has been assessed in fermented foods
(Nawaz et al., 2011) as well as more spe-cifically
for probiotic food products (Saarela et al., 2007;
Mater et al., 2008; Vankerckhoven et al., 2008).
Results of such studies confirm the role of a
reservoir of antibiotic resistance genes from the
food microbiota, without identifying any major
health concerns to date.
It is considered that strains carrying acquired
antibiotic resistance genes might act as a
reservoir of transmissible antimicrobial resis-
tance determinants (FEEDAP, 2005, 2008). Gene
transfer of antibiotic resistance between
microorganisms in the food and feed chain is
thus considered to be a topic of surveillance for
the safety demonstration of microorganisms
(FAO and WHO, 2001, 2002; Borriello et al.,
2003; Gueimonde et al., 2005).

4. Inventory of microbial species used in

food fermentations

The “2002 IDF Inventory” listed 82 bacterial

species and 31 species of yeast and molds
whereas the present “Inventory of MFC” contains
195 bacterial species and 69 species of yeasts
and molds. The over-view of the distribution of
species over the relevant taxonomic units
F. Bourdichon et al. / International Journal of Food Microbiology 154 (2012) 87–97 91

is given in Table 1 for bacteria and Tables 2 andTable 2

3 for fungi. We pub-lish the complete current Fungal diversity in the 2011 update of
microorganisms with beneficial use.
“Inventory of Microbial Food Cultures” as
accompanying material to the present paper.

4.1. Bacteria

4.1.1. Actinobacteriaceae
The genus Brachybacterium enters the list
with two species, B. ali-mentarium and B.
tyrofermentans. Both species have been
characterized as important and beneficial
components of the surface microbiota of Gruyère
and Beaufort cheese (Schubert et al., 1996).
Microbacterium enters the list with one
species, M. gubbeenense. M. gubbeenense is a
component of the traditional red smear surface
culture of surface ripened cheeses (Bockelmann
et al., 2005). The spe-cies was first proposed by
Brennan and colleagues in 2001 (Brennan et al.,
2001), and before this, M. gubbeenense isolates
would have been considered members of
Arthrobacter nicotinae, a species includ-ed in the
“2002 IDF Inventory”.

Bifidobacterium was represented with eight

species in the 2002 IDF inventory. On the one
hand, the species B. infantis disappears, as this
taxon is now transferred to B. longum as B.
longum subsp. infantis. On the other hand, the
species B. thermophilum is included on the list
as this species is reported to have food
applications (Xiao et al., 2010).
The species Brevibacterium aurantiacum,
established in 2005, has entered the list. This
species is like the two other Brevibacterium spe-
cies, B. linens and B. casei, a component of the
red smear ripening microbiota for surface
ripened cheeses (Leclercq-Perlat et al., 2007).
Corynebacterium casei and Corynebacterium
variabile are added to the list as both are
components of the surface ripening microbiota.
C. casei is a relatively “new” species
(Bockelmann et al., 2005).
Micrococcus was represented with one
species on the 2002 IDF in-ventory, M. varians.
The species was renamed and attributed to the
genus Kocuria (Stackebrandt et al., 1995). On the
current list, Micro-coccus is represented with the
two species, M. luteus and M. lylae,
 Phylum Family Genus Species
Ascomycota Cordycipitaceae Lecanicillium 1
Dipodascaceae Geotrichum 1
Yarrowia 1
Galactomyces 1
Microascaceae Scopulariopsis 1
Nectriaceae Fusarium 2
Saccharomycetaceae Candida 10
Cyberlindnera 2
Debaryomyces 1
Dekkera 1
Hanseniaspora 3
Kazachstania 2
Kluyveromyces 1
Lachancea 2
Metschnikowia 1
Pichia 4
Saccharomyces 4
Schwanniomyces 1
Starmerella 1
Trigonopsis 1
Wickerhamomyces 1
Zygosaccharomyces 1
Zygotorulaspora 1
Kluyveromyces 1
Sarcosomataceae Torulaspora 1
Schizosaccharomycetaceae Schizosaccharomyces 1
Sordariaceae Neurospora 1
Trichocomaceae Aspergillus 4
Penicillium 7
Ascomycota—species 59
Basidiomycota Cystofilobasidiaceae Cystofilobasidium 1
Guehomyces 1
Basidiomycota—species 2
Zygomycota Mucoraceae Mucor 4
Rhizopus 4
Zygomycota—species 8
Total number of species 69
and C. piscicola.inventory if meat species.
The in-clusion ofcultures had been
Table 1
used for cheese Carnobacterium included at the
Bacterial diversity in the 2011 update
of microorganisms with beneficial use.ripening and meat commonly used intime. Weissella
fermentation, meat species are used
Phylum Family
respectively fermentations for fermen-tation
Actinobacteria Bifidobacteriaceae
(Bonnarme et al., stems fromof meat, fish,
Brevibacteriaceae
2001; Garcia widening thecabbage (Kimchi),
Corynebacteriaceae
Fontan
Dermabacteraceae et al., scope of the listcassava, and cocoa
2007).
Microbacteriaceae from dairy to food(Collins et al.,
Micrococcaceae Propionibacter fermentations 1993).
ium includes one (Hammes et al.,
new subspecies of
Propionibacteriaceae
1992). Among the
P. freudenreichii
Streptomycetaceae The genusenterococci,
Actinobacteria—species subsp. globosum, Tetragenococcus Enterococcus
Firmicutes Bacillaceae and the newly was proposed infaecalis has
Carnobacteriaceae
added species P. 1990 andentered the list
Enterococcaceae
jensenii. The validated in 1993owing to its use in
species
Lactobacillaceae P. for newlydairy, meat,
arabinosum is identified speciesvegetables and
Leuconostocaceae and some speciesprobiotics
considered
synonymous with previously be-(Foulquie Moreno
P. acidipropionici
Staphylococcaceae longing toet al., 2006).
and is thus no Pediococcus and The genus
Streptococacceae
longer on the list Enterococcus. Lactobacillus was
as a separate The genusalready widely
Firmicutes—species
entity. Weissella waspresent in the
Proteobacteria Acetobacteraceae
introduced in 1993initial inventory.
Enterobacteriaceae for some speciesOwing to its wide
4.1.2. Firmicutes pre-viously use in other food
Sphingomonadaceae
The genus belonging to thematrices and the
Proteobacteria—species
Total number of species Carnobacterium is Leuconostoc new scope of the
new on the list and mesenteroides inventory, this is
is now represented species group.the genus with the
by three species, Weissella wouldlargest number of
C. divergens, C. have been in thechanges and now
maltaromaticum, 2002 IDFrepresented by 82
92 F. Bourdichon et al. / International Journal of Food Microbiology 154 (2012) 87–97

Table 3
Filamentous fungi and yeasts for beneficial use and their teleomorphs, anamorphs and most important synonyms.

Current name Teleomorphic state Anamorphic state Important synonyms

Aspergillus acidus – Aspergillus acidus Aspergillus foetidus
Aspergillus niger Aspergillus niger
Aspergillus oryzae Aspergillus oryzae
Aspergillus sojae Aspergillus sojae
Candida etchellsii Candida etchellsii Torulopsis etchelsii
Candida milleri Candida milleri Candida humilis
Candida oleophila Candida oleophila Candida deformans
Candida rugosa Candida rugosa Mycoderma rugosum
Candida tropicalis Candida tropicalis Odium tropicale,
Candida kefyr
Candida versatilis Candida versatilis Torulopsis versatilis
Candida zemplinina Candida zemplinina
Candida zeylanoides Candida zeylanoides Monilia zeylanoides
Cyberlindnera jadinii Cyberlindnera jadinii Candida guillermondii Candida utilis
Hanseluna jadinii
Cyberlindnera mrakii Cyberlindnera mrakii Williopsis mrakii,
Hansenula mrakii
Cystofilobasidium infirmominiatum Cystofilobasidium infirmominiatum Cryptococcus infirmominiatus Rhodosporium infirmominatum
Debaryomyces hansenii Debaryomyces hansenii Atelosaccharomyces hudeloi Pichia hansenii
Dekkera bruxellensis Dekkera bruxellensis Brettanomyces abstinens
Fusarium domesticum Fusarium domesticum Trichothecium domesticum
Fusarium venenatum Fusarium venenatum
Galactomyces candidum Galactomyces candidum
Geotrichum candidum Geotrichum candidum Acrosporium candidum
Guehomyces pullulans Guehomyces pullulans Trichosporon fuscans
Hanseniaspora guilliermondii Hanseniaspora guilliermondii Kloeckera apiculata Hanseniaspora apuliensis
Hanseniaspora osmophila Hanseniaspora osmophila Kloeckera corticis
Hanseniaspora uvarum Hanseniaspora uvarum Kloeckeraspora uvarum Hanseniaspora apiculata
Kazachstania exigua Kazachstania exigua Candida holmii
Kazachstania unispora Kazachstania unispora Saccharomyces unisporus
Kluyveromyces lactis Kluyveromyces lactis Saccharomyces lactis
Kluyveromyces marxianus Kluyveromyces marxianus Atelosaccharomyces pseudotropicalis Saccharomyces marxianus
Lachancea fermentati Lachancea fermentati Zygosaccharomyces fermentati
Lachancea thermotolerans Lachancea thermotolerans Kluyveromyces thermotolerans
Lecanicillium lecanii Cordyceps confragosa Lecanicillium lecanii Verticillium lecanii
Metschnikowia pulcherrima Metschnikowia pulcherrima Asporomyces uvae Candida pulcherrima
Mucor hiemalis
Mucor mucedo
Mucor plumbeus
Mucor racemosus
Neurospora sitophila Neurospora sitophila Chrysonilia sitophila
Penicillium camemberti Penicillium camemberti Penicillium album,
Penicillium candidum,
Penicillium caseicola,
Penicillium rogeri
Penicillium caseifulvum Penicillium caseifulvum
Penicillium chrysogenum Penicillium chrysogenum Penicillium notatum
Penicillium commune Penicillium commune Penicillium cyclopium
Penicillium nalgiovense Penicillium nalgiovense
Penicillium roqueforti Penicillium roqueforti Penicillium aromaticum,
Penicillium gorgonzolae,
Penicillium stilton
Penicillium solitum Penicillium solitum Pemicillium casei,
Penicillium mali
Pichia fermentans Pichia fermentans Zymopichia fermentans
Pichia kluyveri Pichia kluyveri Hansenula kluyveri
Pichia kudriavzevii Pichia kudriavzevii Candida acidothermophilum Issatchenkia orientalis
Pichia membranifaciens Pichia membranifaciens Saccharomyces membranifaciens
Pichia occidentalis Pichia occidentalis Candida soli
Pichia pijperi Pichia pijperi Wickerhamomyces pijperi,
Hanseniasporia pijperi
Rhizopus microsporus Mucor microsporus
Rhizopus oligosporus
Rhizopus oryzae Rhizopus arrhizus,
Mucor arrhizus
Rhizopus stolonifer Mucor stolonifer
Saccharomyces bayanus Saccharomyces bayanus Saccharomyces uvarum
Saccharomyces cerevisiae Saccharomyces cerevisiae
Schizosaccharomyces pombe Schizosaccharomyces pombe Saccharomyces pombe
Schwanniomyces vanrijiae Schwanniomyces vanrijiae Pichia vanrijiae
Scopulariopsis flava Scopulariopsis flava Acaulium flavum
Starmerella bombicola Starmerella bombicola
Torulaspora delbrueckii Torulaspora delbrueckii Candida colliculosa Zymodebaryomyces delbrueckii
Torulopsis candida Torulopsis candida Cryptococcus candidus
Torulopsis holmii Torulopsis holmii Candida holmii
F. Bourdichon et al. / International Journal of Food Microbiology 154 (2012) 87–97 93
Table 3 (continued)
Current name Teleomorphic state Anamorphic state Important synonyms
Trigonopsis cantarellii Trigonopsis cantarellii Candida cantarellii,
Torulopsis vinacea
Wickerhamomyces anomalus Wickerhamomyces anomalus Candida beverwijkiae Saccharomyces anomalus
Yarrowia lipolytica Yarrowia lipolytica Candida deformans Saccharomycopsis lipolytica
Zygosaccharomyces rouxii Zygosaccharomyces rouxii Zygosaccharomyces japonicas,
Torulaspora rouxii
Zygotorulaspora florentina Zygotorulaspora florentina Saccharomyces florentinus,
Torulapora florentinus

Candida, many additional species

have been suggested for beneficial
Leuconostoc is also a genus having expanded considerably from the two filamentous fungi: Aspergillus, use in foods, including C. etchellsii,
species present in the 2002 IDF inventory. This is mainly due to the inclusion of Guehomyces, Mucor, Neurospora, C. intermedia, C. maltosa, C.
species useful for coffee and vegetable fermen-tations, among which are also Rhizo-pus, and Zygosaccharomyces. versatilis and C. zeylanoides.
several species being proposed recently as L. holzapfelii, L. inhae, L. kimchii, The changes in taxonomy have, Teleomorphic states are not known
and L. palmae. however, also contributed to for these species. Other species
Staphylococcus is now represented by 13 species. The growth in number is changing the appearances in the recently suggested include
caused by the consideration of mostly meat fermentation processes and the role inventory. Most of the species Clavispora lusitanae,
in numerous other food matrices (Nychas and Arkoudelos, 1990). recorded as Candida in the former list Cystofilobasidium infirmominiatum,
have been transferred to other genera Dekkera bruxellensis, Hanseniaspora
Lactococcus has only been expanded with a single species L. raffi-nolactis, aor included under the teleomorphic uvarum, Kazachstania turicensis,
species occasionally involved in the ripening of cheese (Ouadghiri et al., 2005). name (Table 3). Re-cently, it has been Metschniko-wia pulcherrima, Pichia
Also Streptococcus has increased with a single species, due to the use of S. suggested by many mycologists that occidentalis, Rhodosporidium sp.,
gallolyticus subsp. macedonicus in ripening cultures for cheese (Georgalaki etonly one name should be given to Saccharomyces pastorianus,
al., 2000). any fungus, as is already done in Saccharomycopsis fibuligera,
Bacillus species have been included in the inventory due to the wid-ening of Zygomy-cota. Thus it would be Saturnisporus saitoi, Sporobolo-
scope by incorporation of new food matrices such as cocoa beans (Schwan andpreferred to refer to the most well- myces roseus, Torulaspora
Wheals, 2010) and soy beans (Kubo et al., 2011). delbrueckii, Trichosporon cutaneum,
known spe-cies as Saccharomyces
Wickerha-momyces anomalus,
cerevisiae (the teleomorphic and
Yarrowia lipolytica,
4.1.3. Proteobacteriaceae holomorphic name), rather than the
Zygosaccharomyces bailii, and
Acetobacter and Gluconacetobacter are represented by nine and eight anamorphic name Candida robusta. Z. rouxii. In the current update of the
species, respectively. They are mainly utilized in the production of vinegar, but According to present rules as guided inventory of microorganisms, we
also of importance in the fermentation of cocoa and coffee (Sengun andby the International Code of tend to be conservative and only
Karabiyikli, 2011). Botanical No-menclature Article 59,
include species with a well-
Halomonas elongata, a new species of the family Enterobacteria-ceae, was fungi in Ascomycota and
documented technological benefit.
added to the list because of its relevance in meat fermenta-tion (Hinrichsen etBasidiomycota can have two names; One example is Dekkera bruxellensis
al., 1994). one for the teleomorph and
(anamorph Brettanomyces
As a consequence of the widened scope of the inventory, the genusholomorph, which is recommended, bruxellensis), which was formerly
Zymomonas has been added to the list. It is represented by the species Z. and one for the anamorphic state. regarded as a spoiler of beer (and
mobilis, which is widely used for the fermentation of alcoholic bev-erages in wine). However, it is used for
many tropical areas of America, Africa, and Asia (Rogers et al., 1984; Escalante production of Bel-gian Lambic-
et al., 2008). 4.2.1. Yeasts Geuze beer. D. bruxellensis produces
Klebsiella mobilis, formerly Enterobacter aerogenes in the 2002 IDF Candida famata is the anamorph acetic acid that in moderate amounts
inventory, was rejected as the reference of food usage (Gassem, 1999) indicatedof Debaryomyces hansenii. Candida gives a unique taste to those beers
the species as part of the spoilage microbiota. utilis, used for single cell protein (Boekhout and Roberts, 2003). Other
production, should be called examples are Debaryomyces hansenii
4.2. Fungi Cyberlind-nera jadinii. Williopsis and Yarrowia lipolytica which are
mrakii (= Hansenula mrakii) is now very important for aroma formation
The number of recognized species with beneficial use for foods has grownalso included in the genus in Munster and Parmesan cheeses.
considerably. Contributions to the expansion come from changes in taxonomyCyberlindnera as C. mrakii. Saccharomyces cerevisiae,
and description of species to be important in natural fermentations or used as Saccharomyces unisporus has been Hanseniaspora uvarum,
inoculants (Table 3). We have added 24 eukaryotic genera: Aspergillus, transferred to Kazachstania unispora, Kluyveromyces marxianus and
Cyberlindnera, Cystofilobasi-dium, Dekkera, Guehomyces, Hanseniaspora,and Candida holmii has also been Pichia fermentans are extremely
Kazachstania, Lachancea, Lecanicillium, Metschnikowia, Mucor, Neurospora, transferred to Kazachstania as K. impor-tant for the development of the
Rhizopus, Schizosac-charomyces, Schwanniomyces, Scopulariopsis,exigua. Candida krusei is now called fine aroma of cocoa beans (Boekhout
Sporendonema, Starmerella, Torulaspora, Trigonopsis, Wickerhamomyces,Pichia kudriavzevii. Candida kefyr (= and Roberts, 2003).
Yarrowia, Zygosaccharomyces, and Zygotorulaspora. Widening the scope of Candida pseudotropicalis) is placed
food matrices covers a large number of the additions. The inclusion of wine and in Kluyveromyces marxianus.
beverages leads to the addition of the following yeast spe-cies: Cyberlindnera, Candida valida is now called Pichia 4.2.2. Filamentous fungi
Dekkera, Hanseniaspora, Lachancea, Metschniko-wia, Schizosaccharomyces, membranefaciens and finally
Relatively few filamentous fungi
Schwanniomyces, Starmerella, Trigonopsis, and Wickerhamomyces; and the Saccharomyces florentinus is now have been added to the list since the
inclusion of soy and vegetable fermentations leads to the addition of the called
last compilation. However, several
following yeast and Zygotorulaspora florentina (Table 3;
Boekhout and Robert, 2003; fungal starter cultures
Kurtzman et al., 2011). Regarding
94 F. Bourdichon et al. / International Journal of Food Microbiology 154 (2012) 87–97
Olivigni and Bullerman, 1978;
commonly used in Asia could potentially be used Engel and Prokopek, 1980; Finally, some fungi can be
in Europe, as fungi can add fiber, vitamins,Teuber and Engel, 1983; used to produce food
proteins etc. to fermented foods, or be consumed Erdogan and Sert, 2004). colorants, includ-ing
as single cell protein (SCP) (Nout, 2000, 2007).However, P. roqueforti itself Epicoccum nigrum and
Aspergillus species and other fungi found in can produce the secondary Penicillium purpurogenum, but
Asian traditional fermented foods were not men-metab-olites PR-toxin, these fungi are not used
tioned in the first 2002 IDF inventory list as theyroquefortine C, mycophenolic directly for food fermentation
are not commonly used in fermented dairy acid and andrastin A in pure (Stricker et al., 1981; Mapari et
products. For instance Aspergillus oryzae and A.culture (Frisvad et al., 2004; al., 2010).
sojae are used in the production of miso and soya Nielsen et al., 2005). One of
sauce fermenta-tions. Aspergillus oryzae and A. these secondary metabolites is
niger are also used for production of sake and regarded as a mycotoxin, PR- 5. Conclusion
awamori liquors, respectively (Nout, 2000,toxin. This my-cotoxin is
2007). Aspergillus acidus is used for fermentingunstable in cheese and is The list of microorganisms
Puerh tea (Mogensen et al., 2009). converted to PR-imine (Engel
with a history of use in food
Rhizopus oligosporus is used in theand Prokopek, 1979; Siemens origi-nally included 31 genera
fermentation process of Tem-peh (Hachmeisterand Zawistowski, 1993).
in the 2002 IDF inventory, and
and Fung, 1993). Mycophenolic acid (Lafont et
was essen-tially limited to the
Fusarium domesticum was first identified asal., 1979; López-Díaz et al., microbial use in dairy
Trichothecium domesti-cum, but was later1996), roquefortine C (López- matrices. By also considering
allocated to Fusarium (Bachmann et al., 2005; Díaz et al., 1996; Finoli et al., other food matrices, we
Schroers et al., 2009; Gräfenham et al., 2011).2001) and andrastin A (Nielsen consider 62 genera in the 2011
This species has been used for cheeseet al., 2005; Fernández-Bodega update. One was rejected as its
fermentations (cheese smear). Fusarium solaniet al., 2009) have been found usage in food has not been
DSM 62416 was isolated from a Vacherinin blue cheese, but the documented and the initial
cheese, but has not been exam-inedconsequences to human health reference in the 2002 IDF
taxonomically in detail yet. Fusarium venenatum are probably minor (Larsen et inventory was inadequate. The
A 3/5 (first identified as F. graminearum) is being al., 2002). Yet another species, evolution in taxonomy, the
used extensively for mycopro-tein production in Penicillium solitum is found extension of var-ied usages in
Europe (Thrane, 2007). This strain is capable of on naturally fermented lamb other matrices, yeast
producing trichothecene mycotoxins in puremeat on the Faroe Islands, and fermentations and fungal foods
culture, but does not produce them under may be used as a starter
culture. This species does not have also resulted in a growing
industrial conditions (Thrane, 2007). number of species; from 113 to
produce any known
264 species with
Penicillium camemberti is the correct name mycotoxins (Frisvad et al.,
demonstration of food usage.
for the mold use for all white-mold cheeses 2004). On other meat products,
There are many new
(Frisvad and Samson, 2004). Even though P.Penicillium nalgiovense and
possibilities, however, and
com-mune, P. biforme, P. fuscoglaucum, and P. few strains of Penicillium
these should be explored to a
palitans are found on cheese, either as chrysogenum are used (Nout,
much greater extent.
contaminants or “green cheese mold”, they are2000; Frisvad and Samson,
not necessar-ily suitable for fermenting cheeses.2004), especially for mold-
P. commune is the wild-type “an-cestor” of P.fermented salami. However, P. Either in traditional
camemberti however (Pitt et al., 1986; Polonellinal-giovense was originally
found on cheeses from fermented foods or as new
et al., 1987; Giraud et al., 2010).
Nalzovy, and may be used for opportunities, the rationalized
use of microorganisms in our
A species closely related to P. camemberti, P.fermenting cheeses too.
diet opens new perspec-tives.
caseifulvum has an ad-vantage in not producing
In recent years,
cyclopiazonic acid, a mycotoxin often found in
Verticillium lecanii has microorganisms have been
P. camemberti (Lund et al., 1998; Frisvad and
used in fields other than the
Samson, 2004). P. caseifulvum grows naturally changed to Lecanicillium
traditional food industry:
on the surface of blue mold cheeses and has alecanii (Zare and Gams, 2001),
Lactococcus spp. is used for its
valuable aroma (Larsen, 1998). Importantand this strain has been listed
po-tential role in vaccination,
mycotoxins identified in these species include as potentially useful for cheese
and microorganisms are also
cyclopiazonic acid and rugulovasine A and B ripening (see Tables 2 and 3).
used for the specific production
(Frisvad and Samson, 2004), and cyclopiazonic
of biogenic compounds. As we
acid can be detected in white-mold cheeses (Le
did not consider fermentation
Bars, 1979; Teuber and Engel, 1983; Le Bars et
in liquid tailor-made media,
al., 1988).
species used in an industri-al
microbiology process were not
Blue-mold cheeses are always fermented
considered if no reference to
with Penicillium roque-forti, and not with the
food usage could be provided.
closely related species P. carneum, P. paneum or
P. psychrosexualis. The latter three species
Microbiological research
produce several myco-toxins (Frisvad and
mostly focuses on the
Samson, 2004; Houbraken et al., 2010) and have
pathogenic poten-tial of
often been referred to as P. roqueforti (Engel and
microorganisms, while
von Milczewski, 1977; von Krusch et al., 1977;
neglecting their positive role. Recent scientific
advances have revealed the preponderant role of
our own microbiota, our “other genome”, from
the skin, gut, and other mucosa. Though this
remains undoubtedly promising, one should not
forget that man has not yet finished
characterizing traditional fermented foods
consumed for centuries, with often numerous
isolates belong-ing to species with undefined
roles.

6. Acknowledgments and disclaimer

The authors of this paper are the members of

the IDF Task Force on the Update of the
Inventory of Microorganisms with a
Documented History of Use in Foods. The Task
Force is thankful to all National Committees of
the International Dairy Federation for their
helpful support, as well as the associations
EFFCA (European Food & Feed Cultures
Association) and EDA (European Dairy
Association).
The Task Force also took benefit from the
database on Microbial Traditional Knowledge of
India from the Bharathidasan University of
Tiruchirappalli (
http://www.bdu.ac.in/depa/science/biotech/sekar
db. htm) and the publication of a documented
series on fermented foods from the FAO:
bulletins #134—Fermented fruits and vegetables,
#138—Fermented cereals, #142—Fermented
grain legumes, seeds and nuts.

The authors also thank the following experts

for review of the in-ventory: Joelle Dupont
(MNHN, France), Jerôme Mounier (ESMISAB-
LUBEM, France), and Patrick Boyaval
(Danisco, France).

Appendix A. Supplementary

Supplementary data to this article can be

found online at doi:10.
1016/j.ijfoodmicro.2011.12.030.

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