Beruflich Dokumente
Kultur Dokumente
Food Microbiology
journal homepage: www.elsevier.com/locate/fm
a b s t r a c t
Keywords:
Microbial ecology
Lactic acid bacteria
Yeasts
Sourdough
Bakery
Sourdough is a specic and stressful ecosystem inhabited by yeasts and lactic acid bacteria (LAB), mainly
heterofermentative lactobacilli. On the basis of their inocula, three types of sourdough fermentation
processes can be distinguished, namely backslopped ones, those initiated with starter cultures, and those
initiated with a starter culture followed by backslopping. Typical sourdough LAB species are Lactobacillus
fermentum, Lactobacillus paralimentarius, Lactobacillus plantarum, and Lactobacillus sanfranciscensis.
Typical sourdough yeast species are Candida humilis, Kazachstania exigua, and Saccharomyces cerevisiae.
Whereas region specicity is claimed in the case of artisan backslopped sourdoughs, no clear-cut relationship between a typical sourdough and its associated microbiota can be found, as this is dependent on
the sampling, isolation, and identication procedures. Both simple and very complex consortia may
occur. Moreover, a series of intrinsic and extrinsic factors may inuence the composition of the sourdough microbiota. For instance, an inuence of the our (type, quality status, etc.) and the process parameters (temperature, pH, dough yield, backslopping practices, etc.) occurs. In this way, the presence of
Lb. sanfranciscensis during sourdough fermentation depends on specic environmental and technological
factors. Also, Triticum durum seems to select for obligately heterofermentative LAB species. Finally, there
are indications that the sourdough LAB are of intestinal origin.
2013 Elsevier Ltd. All rights reserved.
1. Introduction
The sourdough ecosystem consists of a mixture of our (fractions) and water that is fermented by yeasts and lactic acid bacteria
(LAB) (Corsetti and Settanni, 2007; De Vuyst and Neysens, 2005; De
Vuyst and Vancanneyt, 2007; De Vuyst et al., 2009). From a
microbiological point of view, it is a specic and stressful ecosystem
that is characterized by a low pH, high carbohydrate concentrations, oxygen limitation, and higher cell counts of LAB [108 colony
forming units (CFU)/g] compared to yeasts (107 CFU/g). The major
metabolic activities of the sourdough microbiota are acidication
(LAB), avor formation (LAB and yeasts), and leavening (yeasts and
heterofermentative LAB species). Sourdough provides the production of breads with technological advantages regarding dough
processing and bread texture, avor, and shelf-life. Also, nutritional
properties (sourdough is rich in nutrients), health-promoting effects (owing to endogenous enzymatic modications of the cereal
matrix and/or specic activities of certain LAB species that are, for
instance, reected in gluten-free products), and artisan features
(such as a natural status, traditional value, and gastronomic quality)
contribute to the success of sourdough use. Sourdoughs are produced worldwide (Table 1).
2. Types of sourdough fermentation processes
On the basis of their inocula, three types of sourdough
fermentation processes can be distinguished (Corsetti and Settanni,
2007; De Vuyst and Neysens, 2005; De Vuyst and Vancanneyt,
2007; De Vuyst et al., 2009). A rst type represents spontaneous
sourdough fermentation processes based on backslopping, i.e.,
repeated cyclic re-inoculation of a new batch of our and water
from a previous one derived from the so-called mother dough,
traditionally carried out at room temperature. Thanks to craftsmanship and maintenance of mother doughs for years, a huge variety of bakery products produced from backslopped sourdoughs
exists. These sourdoughs harbor mixtures of distinctive yeasts and
LAB species and/or strains, which actually represent a large diversity of natural sourdough starters. Apparently, they also reect
0740-0020/$ e see front matter 2013 Elsevier Ltd. All rights reserved.
http://dx.doi.org/10.1016/j.fm.2013.06.002
Please cite this article in press as: De Vuyst, L., et al., Microbial ecology of sourdough fermentations: Diverse or uniform?, Food Microbiology
(2013), http://dx.doi.org/10.1016/j.fm.2013.06.002
Country
Reference(s)
Belgium
Wheat
Molecular
LAB: Lb. acidifarinae, Lb. brevis, Lb. buchneri, Lb. crustorum, Lb. hammesii, Lb. helveticus, Lb. nantensis, Lb.
parabuchneri, Lb. paracasei, Lb. paralimentarius, Lb. plantarum, Lb. pontis, Lb. rossiae, Lb. sakei, Lb.
sanfranciscensis, Lb. spicheri, Leuc. mesenteroides, P. pentosaceus, W. cibaria, W. confusa
Yeasts: S. cerevisiae, K. barnettii, K. unispora, T. delbrueckii, W. anomalus
LAB: Lb. brevis, Lb. hammesii, Lb. nantensis, Lb. paralimentarius, Lb. plantarum
Yeasts: S. cerevisiae, W. anomalus
Rye
Molecular
Wheat-rye
Molecular
Spelt
Molecular
Wheat rye spelt
Molecular
Denmark
Estonia
Ethiopia
Rye
Phenotypical
Rye e use of a starter (LAB: Lb. helveticus, Lb. panis, Lb. pontis, Lb.
vaginalis)
Molecular
Teff (enjera)
Phenotypical
Teff
Molecular
Teff (enjera)
Phenotypical
LAB: E. faecalis, Lb. brevis, Lb. fermentum, Lb. plantarum, Leuc. mesenteroides, P. cerevisiae
Gashe, 1985
Nigatu, 2000
Desiye and Abegaz, 2013
LAB: E. mundtii, Lb. brevis, Lb. curvatus, Lb. fermentum, Lb. paracasei, Lb. paralimentarius, Lb. plantarum, Lb.
pontis, Lb. rossiae, Lb. sanfranciscensis, P. acidilactici, P. pentosaceus, W. confusa
Yeasts: S. cerevisiae, T. delbrueckii
LAB: Lb. brevis, Lb. curvatus, Lb. hammesii, Lb. nantensis, Lb. paralimentarius, Lb. plantarum, Lb. pontis, Lb.
rossiae, Lb. sakei, Lb. sanfranciscensis, P. pentosaceus, W. cibaria
Yeasts: S. cerevisiae
LAB: Lb. brevis, Lb. hammesii, Lb. namurensis, Lb. paralimentarius, Lb. plantarum
Yeasts: W. anomalus
Finland
Rye
Phenotypical molecular
LAB: E. casseliavus, Lb. fermentum, Lc. piscium, Lc. plantarum, Lc. rafnolactis, Leuc. mesenteroides,
P. acidilactici, P. pentosaceus
Yeasts: C. humilis, C. tropicalis, K. exigua, P. norvegensis, S. cerevisiae
LAB: Lb. acidophilus, Lb. casei, Lb. plantarum
Yeasts: C. humilis, C. stellata, K. exigua, K. unispora, S. cerevisiae, W. anomalus
France
Wheat
Phenotypical
Wheat
Phenotypical
Wheat
Molecular
Wheat
Molecular
LAB: Lb. acidophilus, Lb. brevis, Lb. casei, Lb. curvatus, Lb. delbrueckii subsp. delbrueckii, Lb. plantarum, Leuc.
mesenteroides subsp. dextranicum, Leuc. mesenteroides subsp. mesenteroides, P. pentosaceus
LAB: Lb. brevis, Lb. curvatus, Lb. plantarum, Lb. rhamnosus, Leuc. mesenteroides, P. pentosaceus
Yeasts: K. exigua, S. cerevisiae, T. delbrueckii, W. anomalus
LAB: Lb. frumenti, Lb. hammesii, Lb. nantensis, Lb. panis, Lb. paralimentarius, Lb. pontis, Lb. sanfranciscensis, Lb.
spicheri, Leuc. mesenteroides subsp. mesenteroides
LAB: Lb. frumenti, Lb. hammesii, Lb. nantensis, Lb. panis, Lb. paralimentarius, Lb. sanfranciscensis, Lb. spicheri,
Leuc. mensenteroides subsp. mensenteroides
China
Wheat (laboratory)
Molecular
Spelt (laboratory)
Molecular
Wheat (laboratory)
Molecular microarray
Spelt (laboratory)
Molecular microarray
Rye (laboratory)
Molecular
Wheat
Molecular
Please cite this article in press as: De Vuyst, L., et al., Microbial ecology of sourdough fermentations: Diverse or uniform?, Food Microbiology
(2013), http://dx.doi.org/10.1016/j.fm.2013.06.002
Table 1
Non-exhaustive overview of the species diversity of lactic acid bacteria (LAB) and/or yeasts in sourdoughs made from various ours and of different geographical origins. The methods of identication (phenotypical or molecular)
of the LAB and yeast species are indicated. This table is an extended version of the one published in Huys et al. (2013).
LAB: E. hirae, Lb. brevis, Lb. casei/paracasei, Lb. curvatus, Lb. paraplantarum, Lb. pentosus, Lb. plantarum, Lb.
sakei, Lb. sanfranciscensis, Lc. lactis, Leuc. citreum, Leuc. mesenteroides, P. pentosaceus, W. cibaria, W. confusa
LAB: Lb. acetotolerans, Lb. amylolyticus, Lb. frumenti, Lb. panis
LAB: Lb. brevis, Lb. buchneri, Lb. casei, Lb. delbrueckii, Lb. fermentum, Lb. plantarum
Spicher, 1959
LAB: Lb. acidophilus, Lb. alimentarius, Lb. brevis, Lb. buchneri, Lb. casei, Lb. farciminis, Lb. fermentum, Lb.
fructivorans, Lb. plantarum, Lb. sanfranciscensis
Yeasts: K. exigua, P. kudriavzevii, S. cerevisiae
LAB: Lb. acidophilus, Lb. brevis, Lb. buchneri, Lb. casei, Lb. farciminis, Lb. fermentum, Lb. fructivorans, Lb.
plantarum, Lb. sanfranciscensis, Pediococcus spp.
LAB: Lb. brevis, Lb. casei, Lb. farciminis, Lb. hilgardii, Lb. homohiochii, Lb. plantarum (spontaneous)
Lb. brevis, Lb. hilgardii, Lb. sanfranciscensis, W. viridescens (masa madre)
LAB: Lb. amylovorus, Lb. frumenti, Lb. pontis, Lb. reuteri (type II)
LAB: Lb. curvatus, Lb. fermentum, Lb. gallinarum, Lb. kimchii, Lb. plantarum, Lb. pontis
Yeasts: P. kudriavzevii, S. cerevisiae
LAB: E. faecium, Lb. casei, Lb. fermentum, Lb. plantarum, Lb. sanfranciscensis, P. pentosaceus, W. confusa
LAB: Lb. fermentum, Lb. helveticus, Lb. paralimentarius, Lb. plantarum, Lb. spicheri
Yeasts: C. glabrata, S. cerevisiae
Spicher, 1984
Spicher, 1987
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(2013), http://dx.doi.org/10.1016/j.fm.2013.06.002
Germany
Wheat
Molecular
Wheat (mother sponge)
Phenotypical molecular
Wheat
Phenotypical
Rye (sourdough starter)
Phenotypical
Country
Greece
Italy
Reference(s)
Millet e use of a starter (LAB: Lb. acetotolerans, Lb. brevis, Lb. casei,
Lb. curvatus, Lb. fermentum, Lb. helveticus, Lb. paracasei, Lb.
paralimentarius, Lb. plantarum, Lb. pontis, Lb. sanfranciscensis, Lb.
spicheri, Leuc. paramesenteroides, Lc. lactis e Yeasts: C. humilis,
P. anomala, P. kudriavzevii, S. cerevisiae, Torulaspora sp.)
Oat e use of a starter (LAB: Lb. acetotolerans, Lb. brevis, Lb. casei, Lb.
curvatus, Lb. fermentum, Lb. helveticus, Lb. paracasei, Lb.
paralimentarius, Lb. plantarum, Lb. pontis, Lb. sanfranciscensis, Lb.
spicheri, Leuc. paramesenteroides, Lc. lactis e Yeasts: C. humilis,
P. anomala, P. kudriavzevii, S. cerevisiae, Torulaspora sp.)
Quinoa e use of a starter (LAB: Lb. acetotolerans, Lb. brevis, Lb. casei,
Lb. curvatus, Lb. fermentum, Lb. helveticus, Lb. paracasei, Lb.
paralimentarius, Lb. plantarum, Lb. pontis, Lb. sanfranciscensis, Lb.
spicheri, Leuc. paramesenteroides, Lc. lactis e Yeasts: C. humilis,
P. anomala, P. kudriavzevii, S. cerevisiae, Torulaspora sp.)
Wheat e use of a starter (LAB: Lb. acetotolerans, Lb. brevis, Lb. casei,
Lb. curvatus, Lb. fermentum, Lb. helveticus, Lb. paracasei, Lb.
paralimentarius, Lb. plantarum, Lb. pontis, Lb. sanfranciscensis, Lb.
spicheri, Leuc. paramesenteroides, Lc. lactis e Yeasts: C. humilis,
P. anomala, P. kudriavzevii, S. cerevisiae, Torulaspora sp.)
Wheat, rye (laboratory e use of a starter; LAB: Lb. fermentum, Lb.
helveticus, Lb. johnsonii, Lb. reuteri, Lb. pontis, Lb. sanfranciscensis e
Yeasts: C. humilis, P. kudriavzevii, S. cerevisiae) Molecular
Wheat, rye, rye bran (laboratory e use of a starter; LAB: Lb.
fermentum, Lb. helveticus, Lb. johnsonii, Lb. reuteri, Lb. pontis, Lb.
sanfranciscensis e Yeasts: C. humilis, P. kudriavzevii, S. cerevisiae)
Molecular
Maize
Phenotypical
Wheat
Phenotypical molecular
Wheat
Phenotypical molecular
Wheat (panettone, brioche)
Phenotypical
Wheat (Umbria)
Phenotypical
Pizza from Naples
Phenotypical
Wheat (Molise, Umbria, Venise)
Molecular
Mother sponges (Lombardy)
Phenotypical molecular
Wheat (Apulia)
Phenotypical molecular
LAB: Lb. fermentum, Lb. helveticus, Lb. paralimentarius, Lb. plantarum, Lb. pontis
Yeasts: P. kudriavzevii, S. cerevisiae
Obiri-Danso, 1994
LAB: Lb. fermentum, Lb. plantarum, Lb. sanfranciscensis, Leuc. mesenteroides, Pediococcus spp.
Yeasts: C. humilis, C. stellata, K. exigua, S. cerevisiae
LAB: Lb. farciminis, Lb. plantarum, Lb. sanfranciscensis
Yeasts: K. exigua, P. kudriavzevii, S. cerevisiae, W. anomalus
LAB: Lb. plantarum, Lb. sakei, Leuc. gelidum, Leuc. mesenteroides
Sicilian sourdoughs
Phenotypical molecular
Wheat (Southern Italy)
Molecular
Wheat
Molecular
Wheat (Molise)
Phenotypical molecular
Wheat (panettone, pandoro, cornetto, colomba)
Phenotypical molecular
LAB: Lb. alimentarius, Lb. brevis, Lb. farciminis, Lb. fermentum, Lb. fructivorans, Lb. plantarum, Lb.
sanfranciscensis, W. confusa
Yeasts: C. humilis
Ghana
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(2013), http://dx.doi.org/10.1016/j.fm.2013.06.002
Table 1 (continued )
Veneto sourdoughs
Molecular
Wheat (Panettone from Central Italy)
Molecular
Wheat (Cornetto, Basilicata)
Phenotypical molecular
Wheat
Phenotypical molecular
Wheat (Marche)
Phenotypical molecular
Wheat
Molecular
Wheat e use of a starter (LAB: Lb. sanfranciscensis)
Molecular
Barley e use of a starter (LAB: Lb. brevis, Lb. curvatus, Lb. helveticus,
Lb. paralimentarius, Lb. plantarum, Lb. sanfranciscensis, Leuc. citreum,
Leuc. pseudomesenteroides, W. confusa, W. kimchii e Yeasts:
C. humilis, S. cerevisiae)
Molecular
Barley wheat e use of a starter (LAB: Lb. brevis, Lb. curvatus, Lb.
helveticus, Lb. paralimentarius, Lb. plantarum, Lb. sanfranciscensis,
Leuc. citreum, Leuc. pseudomesenteroides, W. confusa, W. kimchii e
Yeasts: C. humilis, S. cerevisiae)
Molecular
Emmer e use of a starter (LAB: Lb. plantarum, W. confusa)
Molecular
Spelt e use of a starter (LAB: Lb. brevis, Lb. plantarum, W. confusa)
Molecular
Wheat
Molecular
Wheat e use of a starter (LAB: Lb. plantarum)
Molecular
Wheat e use of a starter (LAB: Lb. plantarum, Lb. sanfranciscensis)
Molecular
Wheat germ e use of a starter (LAB: Lb. plantarum, Lb. rossiae,
P. pentosaceus, W. confusa)
Molecular
LAB: Lb. brevis, Lb. casei, Lb. graminis, Lb. paracasei, Lb. plantarum, Leuc. mesenteroides
LAB: Lb. alimentarius, Lb. brevis, Lb. casei, Lb. farciminis, Lb. pentosus, Lb. plantarum, Lb. sakei, Lb.
sanfranciscensis, Lb. zeae, Leuc. citreum, P. pentosaceus, W. confusa
LAB: Lb. alimentarius, Lb. brevis, Lb. curvatus/graminis/sakei, Lb. fermentum, Lb. paralimentarius, Lb. plantarum,
Lb. rossiae, Lb. sanfranciscensis, Leuc. citreum, P. pentosaceus, W. cibaria
Yeasts: C. humilis, K. exigua, P. kudriavzevii, S. cerevisiae, T. delbrueckii
Yeasts: C. humilis, S. cerevisiae
LAB: Lb. brevis, Lb. farciminis, Lb. plantarum, Lb. sanfranciscensis, Leuc. mesenteroides
Yeasts: C. humilis, S. cerevisiae
LAB: Lb. curvatus, Lb. paraplantarum, Lb. pentosus, Lb. plantarum, Leuc. mensenteroides, W. cibaria
LAB: Lb. brevis, Lb. paralimentarius, Lb. plantarum, Lb. sanfranciscensis, Lc. lactis, Leuc. durionis, Leuc. fructosus,
P. pentosaceus, W. cibaria
Yeasts: C. humilis, S. cerevisiae
LAB: E. faecalis, Lb. brevis, Lb. casei, Lb. curvatus, Lb. fermentum, Lb. gallinarum, Lb. helveticus, Lb. paracasei
subsp. paracasei, Lb. paralimentarius, Lb. plantarum, Lb. rhamnosus, Lb. sakei, Lb. salivarius, Lb. sanfranciscensis,
Leuc. citreum, Leuc. pseudomesenteroides, W. cibaria, W. confusa
Yeasts: C. humilis, S. cerevisiae
LAB: Lb. sanfranciscensis, W. confusa
LAB: Lb. paralimentarius, Lb. plantarum, Lb. rossiae, Lb. sanfranciscensis, Lc. lactis subsp. lactis, P. pentosaceus,
W. cibaria, W. cibaria/confusa
LAB: Lb. brevis, Lb. plantarum
Yeasts: S. cerevisiae
Please cite this article in press as: De Vuyst, L., et al., Microbial ecology of sourdough fermentations: Diverse or uniform?, Food Microbiology
(2013), http://dx.doi.org/10.1016/j.fm.2013.06.002
Wheat (Molise)
Phenotypical molecular
Home-made sourdoughs
Phenotypical molecular
Wheat (Ferrara)
Phenotypical molecular
Wheat (panettone, pandoro, cornetto, colomba)
Phenotypical molecular
Wheat (Sicily)
Molecular
Wheat (Molise)
Phenotypical molecular
Wheat (Altamura, Apulia)
Phenotypical molecular
Wheat (Sardinia)
Molecular
Wheat (Abruzzo)
Physiological molecular
Country
Reference(s)
Wheat (Campania)
Molecular
LAB: Lactobacillus sp., Lb. casei, Lb. coryniformis, Lb. curvatus, Lb. plantarum, Lb. sakei/curvatus, Lb.
sanfranciscensis, Lc. lactis, Leuconostoc sp., Leuc. lactis, Leuc. (pseudo)mesenteroides, S. thermophilus, W. cibaria
Yeasts: S. cerevisiae, M. pulcherrima
LAB: Lb. brevis, Lb. casei, Lb. paracasei, Lb. plantarum
Wheat (Molise)
Phenotypical molecular
Wheat (Pane di Altamura PDO)
Wheat (Pane di Laterza)
Wheat (Pane di Matera PGI)
Wheat (Pane Casereccio, Calabria)
Wheat (Pane Casereccio, Molise)
Wheat (Pane Casereccio, Genzano, PGI)
Wheat (Bozza Pratese)
Wheat (Pane di Altopascio Tradizionale)
Wheat (Pane di Terni)
Wheat (Bastone di Padova)
Wheat (Coppia Ferrarese, PGI)
Wheat (Pane Casereccio Marchigiano)
Wheat (Pane di Cappelli)
Wheat (Moddizzosu)
Wheat (Pane Carasau)
Wheat
Wheat
Wheat
Wheat
Wheat (laboratory)
Iran
Ireland
Mexico
Minervini
Minervini
Minervini
Minervini
Minervini
Minervini
Minervini
Minervini
Minervini
Minervini
Minervini
Minervini
Minervini
Minervini
Minervini
et
et
et
et
et
et
et
et
et
et
et
et
et
et
et
al.,
al.,
al.,
al.,
al.,
al.,
al.,
al.,
al.,
al.,
al.,
al.,
al.,
al.,
al.,
2012a
2012a
2012a
2012a
2012a
2012a
2012a
2012a
2012a
2012a
2012a
2012a
2012a
2012a
2012a
Minervini
Minervini
Minervini
Minervini
et
et
et
et
al.,
al.,
al.,
al.,
2012a
2012a
2012a
2012b
LAB: Lb. amylovorus, Lb. brevis, Lb. fermentum, Lb. frumenti, Lb. paralimentarius, Lb. plantarum, Lb.
sanfranciscensis, Leuc. argentinum, W. cibaria
Yeasts: -
LAB: Lb. amylovorus, Lb. brevis, Lb. fermentum, Lb. frumenti, Lb. paralimentarius, Lb. plantarum, Lb. pontis, Lb.
reuteri, Lb. sanfranciscensis, P. acidilactici
Yeasts: K. barnettii, S. cerevisiae
LAB: Lb. crispatus, Lb. fermentum, Lb. gallinarum, Lb. graminis, Lb. plantarum, Lb. sakei, Lb. vaginalis, Leuc.
holzapfelii, P. pentosaceus, W. cibaria
Yeasts: K. barnettii
LAB: Lb. fermentum, Lb. gallinarum, Lb. pontis, Lb. vaginalis, Leuc. holzapfelii, P. pentosaceus
Yeasts: C. glabrata, S. cerevisiae
LAB: Lb. confusa, Lb. plantarum, Lc. lactis, Lc. rafnolactis, Leuc. mesenteroides
LAB: E. saccharolyticus, Lb. casei, Lb. delbrueckii, Lb. fermentum, Lb. plantarum, S. bovis
Please cite this article in press as: De Vuyst, L., et al., Microbial ecology of sourdough fermentations: Diverse or uniform?, Food Microbiology
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Table 1 (continued )
Nigeria
Pakistan
Portugal
Romania
Russia
Spain
South Korea
Sudan
Sweden
Thailand
Turkey
USA
LAB: Lb. alimentarius, Lb. casei, Lb. delbrueckii, Lb. plantarum, Lc. lactis, S. suis
LAB: Lb. brevis, Lb. buchneri, Lb. casei, Lb. plantarum, Leuc. mesenteroides, Pediococcus spp.
Yeasts: C. humilis, S. cerevisiae
LAB: Lb. buchneri, Lb. casei, Lb. delbrueckii, Lb. plantarum, Lb. sanfranciscensis, Leuc. mesenteroides,
P. pentosaceus
LAB: Lb. brevis, Lb. casei, Lb. fermentum, Lb. plantarum, Leuc. mesenteroides, P. acidilactici
Yeasts: S. cerevisiae
LAB: E. casseliavus, E. durans, E. faecium, Lb. brevis, Lb. curvatus, Lc. lactis subsp. lactis, Leuconostoc spp.,
S. constellatus, S. equines
Yeasts: S. cerevisiae, T. delbrueckii, W. anomalus
LAB: Enterococci, lactobacilli, lactococci, leuconostocs, streptococci
LAB: Lb. brevis, Lb. helveticus, Lb. plantarum, Lc. lactis subsp. lactis, W. confusa
Yeasts: S. cerevisiae, Kl. marxianus
LAB: Lb. helveticus
Yeasts: Kl. marxianus
LAB: Lb. brevis, Lb. fermentum, Lb. plantarum
LAB: Lb. brevis, Lb. plantarum
Yeasts: S. cerevisiae
LAB: Lb. brevis, Lb. cellobiosus, Lb. plantarum, Leuc. mesenteroides
Yeasts: Meyerozyma guilliermondii, S. cerevisiae
Leuc. citreum, W. koreensis
LAB: Lb. acidophilus, Lb. brevis, Lb. delbrueckii, Lb. farciminis, Lb. fermentum, Lb. plantarum, Lb. rhamnosus, Lb.
sanfranciscensis, W. viridescens
LAB: Lactobacillus sp., P. pentosaceus
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(2013), http://dx.doi.org/10.1016/j.fm.2013.06.002
Morocco
Maize (pozol)
Phenotypical
Maize (pozol)
Molecular
Sourdough ferments, traditional starter sponges
Phenotypical
Soft wheat our
Phenotypical
Maize
Molecular
Wheat
Phenotypical
Rye maize wheat
Phenotypical
Maize (broa, Northern Portugal)
Phenotypical
LAB: C., Carnobacterium; E., Enterococcus; Lb., Lactobacillus; Lc., Lactococcus; Leuc., Leuconostoc; P., Pediococcus; S., Streptococcus; W., Weissella.
Yeasts: C., Candida; D., Debaryomyces; K., Kazachstania; Kl., Kluyveromyces; M., Metschnikowia; P., Pichia; S., Saccharomyces; T., Torulaspora; W., Wickerhamomyces.
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10
amplication, inadequately standardized electrophoresis conditions, and the occurrence of heteroduplex bands for the same
species. The use of microarrays is limited to the identication of
known species because of prior probe design, during which
attention should go to the avoidance of inter-species cross-hybridization events. Finally, next-generation sequencing results are
biased by the choice and selectivity of the primers used in the case
of targeted approaches, sequencing depth and read length, and
algorithms and databases available and used for post-sequencing
data analysis.
4.3. Problematic identications
Based on the literature review of sourdough yeasts mentioned
above and comparing studies using phenotypic identication
techniques (n 19) with those using DNA-based techniques
(n 23), the incidence of C. humilis isolates has increased markedly
in the latter studies, probably at the cost of a decreased frequency of
detecting K. exigua. These two phylogenetically closely related yeast
species may be mistaken for each other when only phenotypical
identication methods are used. For instance, originally, S. exiguus
was isolated from San Francisco sourdough (Sugihara et al., 1971),
while C. milleri was reported from various sourdoughs produced in
Italy (Ottogalli et al., 1996; Valmorri et al., 2010; Venturi et al.,
2012b). Some strains of S. exiguus from the study of Sugihara
et al. (1971) that were preserved in culture collections later
served for the description of C. milleri, currently a synonym of
C. humilis. The reassignment of S. exiguus strains to C. milleri was
based on the signicantly higher guanine-plus-cytosine contents in
selected San Francisco sourdough strains compared to the type and
other reference strains of S. exiguus. However, the sourdough
isolate M14 initially reported as S. exiguus (and also deposited as
CBS 7901) was later considered as a member of C. humilis after
DNA-based analyses (Gobbetti et al., 1994a; Meroth et al., 2003a).
Actually, the yeast species complex C. humilis/C. milleri was placed
in the articial genus Candida, because no sexual reproduction
could be found. Phylogenetically, C. humilis/C. milleri belongs to the
same group as the genus Kazachstania (Kurtzman, 2003), but it has
not yet been transferred to this genus mainly because the phylogenetic reclassication of yeasts is treating sexually reproducing
taxa with priority. Anyway, C. humilis and C. milleri are indicated to
be conspecic (Daniel and Meyer, 2003; Meyer et al., 1998;
Vaughan-Martini, 1995). Moreover, a continuum of internal transcribed spacer (ITS) sequence variants between both type strains
exists, making allocation of sourdough isolates of these species to
one of them difcult. These data suggest that C. humilis and
C. milleri may be considered as one species, and because C. humilis
and K. exigua are closely related phylogenetically, it is likely that
K. exigua may be a sourdough-specic yeast species. Additionally,
K. barnettii might have been misidentied as K. exigua or
S. cerevisiae in studies only using phenotypical identication
methods.
Similarly, the LAB species responsible for acidication of San
Francisco sourdough was named Lactobacillus sanfrancisco (Kline
and Sugihara, 1971; Sugihara et al., 1971), whereas that responsible for the acidication of various sourdoughs produced in Italy
was identied as Lb. brevis var. lindneri (Gobbetti et al., 1994b;
Ottogalli et al., 1996). However, both LAB species have been
shown to be synonymous and now considered members of Lb.
sanfranciscensis (Weiss and Schillinger, 1984). Although Lb. sanfranciscensis can be misidentied as Lb. brevis (De Vuyst et al.,
2002), molecular techniques exist for its unequivocal identication (Picozzi et al., 2006; Zapparoli and Torriani, 1997). Furthermore, Lb. plantarum, Lactobacillus paraplantarum, and Lactobacillus
pentosus are phylogenetically closely related, whose taxonomic
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and bakery practices applied determine the stability and persistence of the yeast and LAB communities involved in (region-specic) sourdough fermentation processes. Of course, the use of
certain raw materials, encompassing cereal types and other ingredients such as adjunct carbohydrates, salt, yoghurt, herbs, fruits,
etc., and process parameters such as dough yield and refreshment
times, may be part of specic recipes of a region and hence be
linked to local traditions of that region. Also, they may favor
particular microorganisms as a result of trophic and metabolic
cooperations and interactions and, hence, may associate specic
LAB and/or yeast species with specic sourdoughs of certain
geographical regions. As an example, the presence of the yeast
Metschnikowia pulcherrima in a sourdough destined for a sweet
baked product has been linked with its occurrence on fruits present
in the bakery environment (Palomba et al., 2011).
Whether the nature of the our mainly directs the growth of
sourdough LAB species remains controversial. Of course, the our
plays a key role in establishing stable microbial consortia within a
short time, as described above, and only species and/or strains
adapted to the sourdough environment with respect to the nutrient
availability in and physicochemical parameters of the our will
thrive in this ecosystem. Therefore, among other mechanisms,
competitiveness may explain the apparent prevalence of certain
LAB species and/or strains in specic sourdough fermentations, as
evidenced by single reports on Lb. helveticus in sorghum sourdoughs (Hamad et al., 1997), Lb. amylovorus or Lb. helveticus in rye
sourdoughs (Mller et al., 2001; Viiard et al., 2012), Lb. sakei in
amaranth sourdoughs (Sterr et al., 2009), and Lb. pontis in teff
sourdoughs (Moroni et al., 2011). Particular conditions may have
selected for these microorganisms; for instance, the presence of bglucan may select for b-glucan-degrading LAB species such as Lb.
brevis during barley fermentation (Zannini et al., 2009). However,
these LAB species are not specic for the concomitant sourdough
ecosystems. In contrast, it has been shown that the LAB species Lb.
fermentum and Lb. plantarum dominate the majority of sourdough
fermentation processes, irrespective of the type of our or the
addition of starter cultures that are not robust enough (Moroni
et al., 2010; Van der Meulen et al., 2007; Vogelmann et al., 2009).
It indicates their presence in the cereal ours and on the surfaces of
the concomitant plants. In contrast, the prevalence of Lb. sanfranciscensis depends on appropriate environmental conditions,
albeit that logically it should be present in the our to be found in
the ripe sourdough. Hence, the microbial composition of the our is
of great importance. Also, accidental contamination of the our
from the direct environment (apparatus, air, etc.) determines its
microbial load and thus that of the sourdough fermentation
(Scheirlinck et al., 2009a). Finally, it is likely to assume that sourdough LAB, such as Lactobacillus acidophilus, Lactobacillus johnsonii
as well as Lb. rossiae and Lb. sanfranciscensis, may be of intestinal
origin, owing to fertilization practices on the grain elds;
contamination with mouse and rodent feces of elds, mills, and
apparatus, or the presence of insects in the our mills; as well as to
fecal contamination of the direct sourdough environment (bakerys
hygiene) (De Angelis et al., 2006; Du Toit et al., 2001, 2003;
Ehrmann and Vogel, 2005; Groenewald et al., 2006; Park and
Itoh, 2005; Vrancken et al., 2011). In this way, it is interesting to
see that the frequent sourdough inhabitant, Lb. reuteri, may originate from rodent intestines (Su et al., 2012).
7. Inuence of our
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emmer, millet, oat, rice, and teff to perform sourdough fermentations. Besides cereals, pseudocereals such as buckwheat, amaranth,
and quinoa can be used as the raw material for sourdough
fermentation. In contrast to wheat, rye, and barley, where different
parts of the kernel can be fractionated, the whole kernel is used in
the case of teff because of its small size. However, bran fermentation may have a different impact on the resulting ripe sourdough,
given the presence of dietary ber and bioactive compounds that
are concentrated in the bran fraction (Katina et al., 2007, 2012;
Liukkonen et al., 2003; Poutanen et al., 2009; Rizzello et al., 2010;
Ross et al., 2004).
As mentioned above, the chemical composition and quality
status of the our determine the community dynamics and
metabolite kinetics of sourdough fermentation processes. With
respect to the microorganisms involved, on the basis of a survey of
87 publications, Lb. plantarum, Lb. brevis, Lb. fermentum, and Lb.
sanfranciscensis are the prevailing LAB species during sourdough
fermentation, irrespective of the our type used. In particular, Lb.
plantarum, Lb. brevis, Lb. sanfranciscensis, Lb. fermentum, and Leuc.
mesenteroides are reported in 38 publications on wheat sourdough
and Lb. plantarum, Lb. brevis, Lb. fermentum, and Lb. sanfranciscensis
are reported in 12 publications on rye sourdough.
Concerning the microbial diversity in buckwheat, amaranth, and
quinoa, no straightforward conclusions can be made due to the low
number of publications available. Overall, the presence of Lb.
plantarum, Lb. fermentum, Lb. paralimentarius, and Lb. helveticus has
been reported. In addition, Lb. fermentum and Lb. plantarum are
found in teff, spelt, rice, oat, millet, maize, cassava, barley and
amaranth sourdoughs, but are never found in emmer sourdoughs.
Also, Lb. brevis has been isolated from teff, spelt, maize, and barley
sourdoughs, but is not found in rice, oat, millet, emmer, cassava,
and amaranth sourdoughs. Isolates identied as Lb. sanfranciscensis
have been retrieved from teff and spelt sourdoughs but not from
rice, oat, millet, maize, emmer, cassava, barley, and amaranth
sourdoughs. Although Lb. helveticus has been found in rye, barley,
cassava, oat, millet, and maize sourdoughs, it has not been found in
rice, spelt, teff, and emmer sourdoughs. Lb. paralimentarius, one of
the most abundant LAB species in sourdough fermentations, has
not been found in oat, emmer, and cassava sourdoughs.
8. Inuence of process parameters
For details, the reader is referred to Tables 2 and 3.
According to the process technology applied, distinction can be
made between type 0, I, II, and III sourdoughs (Huys et al., 2013).
Type 0 sourdoughs are pre-doughs, in which LAB contaminating
the faster growing bakers yeast develop to favor avor formation.
Type I or traditional sourdoughs are usually rm doughs with a low
dough yield that are daily backslopped at ambient temperature to
keep the microorganisms in an active state. Type II sourdoughs are
industrial semi-liquid sourdoughs that are usually performed as
one-step propagation processes of long duration and with high
water content at temperatures above 30 C to enhance acidication.
They are used as dough acidiers or avor ingredients. Type III
sourdoughs are dried sourdoughs to be used as a non-living acidier supplement and avor carrier. In contrast to type I doughs,
doughs of types II and III require the addition of bakers yeast for
leavening.
Process parameters such as, but not limited to, fermentation
temperature and time, leavening temperature and time, storage
temperature and time; pH, redox potential, water activity, dough
yield; refreshment time and number of refreshment steps, mixing,
and aeration of the initial dough determine the progress and
outcome of (backslopped) sourdough fermentation processes (De
Vuyst and Neysens, 2005; Gnzle et al., 2007; Gobbetti et al.,
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13
Table 2
Lactic acid bacteria species diversity in (backslopped) bakery or household sourdoughs grouped as a function of the process parameters dough yield, temperature and
fermentation time (nd: not dened; 0: spontaneous; 1: starter culture added; RT: room temperature). Entries are ranked as a function of dough yield, temperature, and
fermentation time. Species are ranked in order of the number of sourdoughs in which they were detected, as indicated by the number between brackets; no number is
mentioned for single detections.
Dough
yield
Temperature
( C)
Fermentation
time (h)
Number of
sourdoughs
analyzed
Species diversity
Reference
nd
nd
nd
nd
nd
nd
nd
nd
nd
nd
nd
nd
7e8
8e10
9
11
13
16
18
20
20
20
20e25
20e28
14
15e17
24
15
20
12
24
18
20
24
4e5
nd
1
1
1
1
1
1
1
1
1
2
1
18
nd
20e30
16
nd
22
nd
nd
nd
nd
nd
nd
nd
22
22
22
22
22
22
5
6
10
11
12
14
1
1
1
1
1
2
nd
nd
nd
22
23
23e28
16
6
6
2
1
20
nd
nd
nd
24
24
25
14
24
3
1
2
7
nd
nd
nd
nd
nd
25
25
25
25
25
5
6
7e8
9
10
1
2
1
1
4
nd
nd
nd
nd
nd
25
25
25
25
25
11
12
15
24
48
1
2
1
3
1
nd
26
nd
nd
nd
nd
nd
nd
26
26
26
27
27
4
8
10
4
11
1
1
1
1
1
nd
27
18
nd
nd
27
28
24
nd
1
9
nd
nd
28
30
6
nd
1
4
nd
nd
nd
30
30
30
3e24
6e7
48e72
3
1
1
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14
Table 2 (continued )
Dough
yield
Temperature
( C)
Fermentation
time (h)
Number of
sourdoughs
analyzed
Species diversity
Reference
nd
30e40
15
10
nd
RT
nd
26
nd
RT
6e8
nd
nd
nd
RT
RT
RT
20
22
24
1
1
4
nd
150
150
160
160
160
160
161
168
169
195
225e250
RT
25
25
7
8
18
26
25
25
25
28
28e34
120e168
5
13
14
16
24
8
6
4
3e4
24
6e13
2
1
1
1
1
1
1
1
1
2
1
3
Lb. plantarum (13/26), Lb. sanfranciscensis (11/26), Lb. hammesii (10/26), Lb. brevis
(9/26), Lb. paralimentarius (8/26), Lb. nantensis (7/26), Lb. rossiae (4/26), Lb.
alimentarius (3/26), Lb. pentosus (3/26), Lb. pontis (3/26), Lb. sakei (3/26), Lb.
farciminis (2/26), Lb. namurensis (2/26), W. confusa (2/26), Lb. buchneri, Lb. curvatus,
Lb. fermentum, Lb. parabuchneri, Leuc. citreum, Leuc. mesenteroides, P. acidilactici,
P. pentosaceus
Lb. alimentarius, Lb. brevis, Lb. casei, Lb. pentosus, Lb. plantarum, Lb. sakei, Lb.
sanfranciscensis, Lb. zeae
Lb. paracasei, Lb. plantarum, Lb. sanfranciscensis, Leuc. pseudomesenteroides
Lb. curvatus, Lb. plantarum, Lb. sakei, Lb. sanfranciscensis, Leuc. citreum, S. salivarius
Lb. fermentum (3/4), Lb. alimentarius/kimchii (2/4), Lb. paralimentarius (2/4), Lb.
plantarum (2/4), Lb. sanfranciscensis (2/4), Lb. brevis, Lb. paracasei, Lb. rhamnosus,
W. confusa
Lb. spicheri (2/2), Lb. paralimentarius (2/2), Lb. paracasei (2/2), Lb. perolens
W. cibaria
Lb. plantarum, Lb. sanfranciscensis
Lb. sanfranciscensis, Leuc. mesenteroides
Lb. frumenti, Lb. hammesii, Lb. paralimentarius
Lb. pontis, Lb. sanfranciscensis, Lb. spicheri
Lb. hammesii, Lb. nantensis, Lb. panis, Lb. sanfranciscensis
Lb. plantarum, Lb. sanfranciscensis
Lb. sanfranciscensis
Lb. plantarum (2/2), Lb. sakei, Leuc. citreum, P. pentosaceus
Lb. amolyticus, Lb. acetotolerans, Lb. frumenti, Lb. panis
Lb. panis (3/3), Lb. amylovorus (2/3), Lb. reuteri (2/3), Lb. helveticus, Lb. pontis
Paramithiotis et al.,
2010
Catzeddu et al., 2006;
Scheirlinck et al., 2007c,
2008
251
500
24
40
4
48
1
1
LAB: E., Enterococcus; F., Fructobacillus; Lb., Lactobacillus; Lc., Lactococcus; Leuc., Leuconostoc; P., Pediococcus; S., Streptococcus; W., Weissella.
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Table 3
Lactic acid bacteria species diversity in end-samples of backslopped laboratory sourdoughs grouped as a function of the process parameters dough yield, temperature and
fermentation time (nd: not dened; 0: spontaneous; 1: starter culture or mother sponge added; RT: room temperature). Entries are ranked as a function of dough yield,
temperature, and fermentation time. Species are ranked in order of the number of sourdoughs in which they were detected, as indicated by the number between brackets; no
number is mentioned for single detections.
Spontaneous/
starter culture
Dough
yield
Temperature
( C)
Fermentation
time (h)
nd
28e35
24
1
1
1
0
187
150
150
160
30
25
25
30
24
5
13
8
3
1
1
2
160
30
23
1
1
1
0
161e169
200
200
200
25e28
20
20
25
3e6
12
24
12
4
1
1
2
200
25
12
1
0
1
1
0
200
200
200
200
200
25
28
28
30
30
24
24
24
12
24
1
1
6
5
200
30
24
29
1
1
0
1
1
200
200
200
200
200
35
35
37
37
37
12
24
24
24
24
2
2
1
1
3
1
1
1
0
200
200
251
275
40
40
24
35
12
24
4
24
1
1
1
2
275
35
24
0
1
0
0
300
367
400
400
37
40
23
30
42
48
24
24
1
1
1
13
1
0
0
1
1
1
1
400
400
400
400
500
500
500
30
30
37
37
34
40
46
24
48
24
24
48
48
48
1
1
1
1
1
1
1
Number of
sourdoughs
analyzed
Species diversity
Reference
2011
2011
2011
2011
2011
LAB: Lb., Lactobacillus; Lc., Lactococcus; Leuc., Leuconostoc; P., Pediococcus; W., Weissella.
the case of the Lb. amylovorus DCE 471 strain (Leroy et al., 2007).
Also, the use of oxygen as alternative external electron acceptor
may be responsible for a shift of the production from ethanol toward acetate (Gobbetti et al., 2005: Martinez-Anaya et al., 1994).
8.4. Backslopping procedure
Whereas backslopping generally selects for heterofermentative
lactobacilli, the amount of backslopped sourdough as well as the
frequency and duration of the backsloppings inuence the community dynamics and stability of the sourdough microbiota
(Minervini et al., 2012b). The amount of backslopping dough denes the initial pH and hence inuences the growth and acidication rates of the LAB species involved, as this is correlated with
the DY and hence the availability of water, as described above
(Brandt et al., 2004; Van der Meulen et al., 2007). It is anticipated
that short refreshment times select for rapidly growing, fastacidifying LAB species, at least under a given set of environmental
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Barber, S., Bguena, R., 1988. Microora of the sour dough of wheat our bread. V.
Isolation, identication and evaluation of functional properties of sourdoughs
microorganisms. Rev. Agroqum. Tecnol. Aliment. 28, 67e78.
Barber, S., Bguena, R., 1989. Microora de la masa madre panaria. XI. Evolucin de
la microora de masas madre durante el proceso de elaboracin por el sistema
de refrescos sucesivos y de sus correspondientes masa panarias. Rev. Agroquim. Tecnol. Aliment. 29, 478e491.
Barber, S., Bguena, R., Martnez-Anaya, M.A., Torner, M.J., 1983. Microora of the
sour dough of wheat our bread. I. Identication and functional properties of
microorganisms of industrial sour doughs. Rev. Agroqum. Tecnol. Aliment. 23,
552e562.
ben Omar, N., Ampe, F., 2000. Microbial community dynamics during production of
the Mexican fermented maize dough pozol. Appl. Environ. Microbiol. 66, 3664e
3673.
Berthier, F., Ehrlich, S.D., 1999. Genetic diversity within Lactobacillus sakei and
Lactobacillus curvatus and design of PCR primers for its detection using
randomly amplied polymorphic DNA. Int. J. Syst. Bacteriol. 49, 997e1007.
Bjrkroth, K.J., Schillinger, U., Geisen, R., Weiss, N., Hoste, B., Holzapfel, W.H.,
Korkeala, H.J., Vandamme, P., 2002. Taxonomic study of Weissella confusa and
description of Weissella cibaria sp nov., detected in food and clinical samples.
Int. J. Syst. Evol. Microbiol. 52, 141e148.
Boraam, F., Faid, M., Larpent, J.P., Breton, A., 1993. Lactic acid bacteria and yeasts
associated with traditional Moroccan sour-dough bread fermentation. Sci.
Aliment. 13, 501e509.
Brandt, M.J., Hammes, W.P., Gnzle, M.G., 2004. Effects of process parameters on
growth and metabolism of Lactobacillus sanfranciscensis and Candida humilis
during rye sourdough fermentation. Eur. Food Res. Technol. 218, 333e338.
Catzeddu, P., Mura, E., Parente, E., Sanna, M., Farris, G.A., 2006. Molecular characterization of lactic acid bacteria from sourdough breads produced in Sardinia
(Italy) and multivariate statistical analyses of results. Syst. Appl. Microbiol. 29,
138e144.
Chaillou, S., Daty, M., Baraige, F., Dudez, A.-M., Anglade, P., Jones, R., Alpert, C.-A.,
Champomier-Vergs, M.-C., Zagorec, M., 2009. Intraspecies genomic diversity
and natural population structure of the meat-borne lactic acid bacterium
Lactobacillus sakei. Appl. Environ. Microbiol. 75, 970e980.
Choi, H., Kim, Y.-W., Hwang, I., Kim, J., Yoon, S., 2012. Evaluation of Leuconostoc
citreum HO12 and Weissella koreensis HO20 isolated from kimchi as a starter
culture for whole wheat sourdough. Food Chem. 134, 2208e2216.
Coda, R., Nionelli, L., Rizzello, C.G., De Angelis, M., Tossut, P., Gobbetti, M., 2010.
Spelt and emmer ours: characterization of the lactic acid bacteria microbiota
and selection of mixed starters for bread making. J. Appl. Microbiol. 108, 925e
935.
Coppola, S., Pepe, O., Masi, P., Sepe, M., 1996. Characterization of leavened doughs
for pizza in Naples. Adv. Food Sci. 18, 160e162.
Corsetti, A., De Angelis, M., Dellaglio, F., Paparella, A., Fox, P.F., Settanni, L.,
Gobbetti, M., 2003. Characterization of sourdough lactic acid bacteria based on
genotypic and cell-wall protein analyses. J. Appl. Microbiol. 94, 641e654.
Corsetti, A., Lavermicocca, P., Morea, M., Baruzzi, F., Tosti, N., Gobbetti, M., 2001.
Phenotypic and molecular identication and clustering of lactic acid bacteria
and yeasts from wheat (species Triticum durum and Triticum aestivum) sourdoughs of Southern Italy. Int. J. Food Microbiol. 64, 95e104.
Corsetti, A., Settanni, L., 2007. Lactobacilli in sourdough fermentation. Food Res. Int.
40, 539e558.
Corsetti, A., Settanni, L., Lopez, C.C., Felis, G.E., Mastrangelo, M., Suzzi, G., 2007a.
A taxonomic survey of lactic acid bacteria isolated from wheat (Triticum
durum) kernels and non-conventional ours. Syst. Appl. Microbiol. 30, 561e
571.
Corsetti, A., Settanni, L., Valmorri, S., Mastrangelo, M., Suzzi, G., 2007b. Identication of subdominant sourdough lactic acid bacteria and their evolution during
laboratory-scale fermentations. Food Microbiol. 24, 592e600.
De Angelis, M., Siragusa, S., Berloco, M., Caputo, L., Settanni, L., Alfonsi, G.,
Amerio, M., Grandi, A., Ragni, A., Gobbetti, M., 2006. Selection of potential
probiotic lactobacillli from pig feces to be used as additives in pelleted feeding.
Res. Microbiol. 157, 792e801.
de las Rivas, B., Marcobal, A., Muoz, R., 2006. Development of a multilocus
sequence typing method for analysis of Lactobacillus plantarum strains.
Microbiol. 152, 85e93.
De Vuyst, L., Neysens, P., 2005. The sourdough microora: biodiversity and metabolic interactions. Trends Food Sci. Technol. 16, 43e56.
De Vuyst, L., Schrijvers, V., Paramithiotis, S., Hoste, B., Vancanneyt, M., Swings, J.,
Kalatzopoulos, G., Tsakalidou, E., Messens, W., 2002. The biodiversity of lactic
acid bacteria in Greek traditional sourdoughs is reected in both composition
and metabolite formation. Appl. Environ. Microbiol. 68, 6059e6069.
De Vuyst, L., Vancanneyt, M., 2007. Biodiversity and identication of sourdough
lactic acid bacteria. Food Microbiol. 24, 120e127.
De Vuyst, L., Vrancken, G., Ravyts, F., Rimaux, T., Weckx, S., 2009. Biodiversity,
ecological determinants, and metabolic exploitation of sourdough microbiota.
Food Microbiol. 26, 666e675.
Di Cagno, R., De Angelis, M., Gallo, G., Settanni, L., Berloco, M.G., Siragusa, S.,
Parente, E., Corsetti, A., Gobbetti, M., 2007. Genotypic and phenotypic diversity
of Lactobacillus rossiae strains isolated from sourdough. J. Appl. Microbiol. 103,
821e835.
Du Toit, M., Dicks, L.M.T., Holzapfel, W.H., 2001. Taxonomy of obligately homofermentative and facultatively heterofermentative lactobacilli in pig faeces. Lett.
Appl. Microbiol. 33, 199e204.
Please cite this article in press as: De Vuyst, L., et al., Microbial ecology of sourdough fermentations: Diverse or uniform?, Food Microbiology
(2013), http://dx.doi.org/10.1016/j.fm.2013.06.002
17
Hamad, S.H., Dieng, M.C., Ehrmann, M.A., Vogel, R.F., 1997. Characterization of the
bacterial ora of Sudanese sorghum our and sorghum sourdough. J. Appl.
Microbiol. 83, 764e770.
Hammes, W.P., Brandt, M.J., Francis, K.L., Rosenheim, J., Seitter, M.F.H.,
Vogelmann, A., 2005. Microbial ecology of cereal fermentations. Trends Food
Sci. Technol. 16, 4e11.
Httner, E.K., Dal Bello, F., Arendt, E.K., 2010. Identication of lactic acid bacteria
isolated from oat sourdoughs and investigation into their potential for the
improvement of oat bread quality. Eur. Food Res. Technol. 230, 849e857.
Huys, G., Daniel, H.-M., De Vuyst, L., 2013. Taxonomy and biodiversity of sourdough
yeasts and lactic acid bacteria. In: Gobbetti, M., Gnzle, M. (Eds.), Handbook on
Sourdough Biotechnology. Springer, New York, USA, pp. 105e154.
Iacumin, L., Cecchini, F., Manzano, M., Osualdini, M., Boscolo, D., Orlic, S., Comi, G.,
2009. Description of the microora of sourdoughs by culture-dependent and
culture-independent methods. Food Microbiol. 26, 128e135.
Infantes, M., Schmidt, J.L., 1992. Characterization of the yeast ora of natural
sourdoughs located in various French areas. Sci. Aliment. 12, 271e287.
Infantes, L., Tourneur, C., 1991. Survey on the lactic ora of natural sourdoughs
located in various French areas. Sci. Aliment. 11, 527e545.
Katina, K., Heini, R.-L., Autio, K., Poutanen, K., 2006. Optimization of sourdough
process for improved sensory prole and texture of wheat bread. LWT e Food
Sci. Technol. 39, 1189e1202.
Katina, K., Laitila, A., Jovonen, R., Liukkonen, K.-H., Kariluoto, S., Piironen, V.,
Landgerg, R., man, P., Poutanen, K., 2007. Bran fermentation as a means to
enhance technological properties and bioactivity of rye. Food Microbiol. 24,
175e186.
Katina, K., Juvonen, R., Laitila, A., Flander, L., Nordlund, E., Kariluoto, S., Piironen, V.,
Poutanen, K., 2012. Fermented wheat bran as a functional ingredient in baking.
Cereal Chem. 89, 126e134.
Kazanskaya, L.N., Afanasyeva, O.V., Patt, V.A., 1983. Microora of rye sours and some
specic features of its accumulation in bread baking plants of the USSR. In:
Holas, J., Kratochvil, F. (Eds.), Developments in Food Science, Progress in Cereal
Chemistry and Technology, vol. 5B. Elsevier, London, United Kingdom, pp. 759e
763.
Kitahara, M., Sakata, S., Benno, Y., 2005. Biodiversity of Lactobacillus sanfranciscensis
strains isolated from ve sourdoughs. Lett. Appl. Microbiol. 40, 353e357.
Kline, L., Sugihara, T.F., 1971. Microorganisms of the San Francisco sourdough bread
process. II. Isolation and characterization of undescribed bacterial species
responsible for the souring activity. Appl. Microbiol. 21, 459e465.
Kurtzman, C.P., 2003. Phylogenetic circumscription of Saccharomyces, Kluyveromyces and other members of the Saccharomycetaceae, and the proposal of the
new genera Lachancea, Nakaseomyces, Naumovia, Vanderwaltozyma and Zygotorulaspora. FEMS Yeast Res. 4, 233e245.
Leroy, F., De Winter, T., Moreno, M.R.F., De Vuyst, L., 2007. The bacteriocin producer
Lactobacillus amylovorus DCE 471 is a competitive starter culture for type II
sourdough fermentations. J. Sci. Food Agric. 87, 1726e1736.
Liukkonen, K.H., Katina, K., Wilhelmsson, A., Myllymaki, O., Lampi, A.M.,
Kariluoto, S., Piironen, V., Heinonen, S.M., Nurmi, T., Adlercreutz, H.,
Peltoketo, A., Pihlava, J.M., Hietaniemi, V., Poutanen, K., 2003. Process-induced
changes on bioactive compounds in whole grain rye. Proc. Nutr. Soc. 62, 117e
122.
Lombardi, A., Zilio, F., Andrighetto, C., Zampese, L., Loddo, A., 2007. Micobiological characterization of sourdoughs of Veneto region. Ind. Aliment. XLVI,
147e151.
Lnner, C., Welander, T., Molin, N., Dostlek, M., Blickstad, E., 1986. The microora in
a sour dough started spontaneously on typical Swedish rye meal. Food Microbiol. 3, 3e12.
Luangsakul, N., Keeratipibul, S., Jindamorakot, S., Tanasupawat, S., 2009. Lactic acid
bacteria and yeasts isolated from the starter doughs for Chinese steamed buns
in Thailand. LWT e Food Sci. Technol. 42, 1404e1412.
Mntynen, V.H., Korhola, M., Gudmundsson, H., Turakainen, H., Alfredsson, G.A.,
Salovaara, H., Lindstrm, K., 1999. A polyphasic study on the taxonomic position
of industrial sour dough yeasts. Syst. Appl. Microbiol. 22, 87e96.
Martinez-Anaya, M.A., Llin, M.L., Pilar Macias, M., Collar, C., 1994. Regulation of
acetic acid production by homo- and heterofermentative lactobacilli in wholewheat sourdoughs. Z. Lebensm. Unters. Forsch. 199, 186e190.
Meroth, C.B., Hammes, W.P., Hertel, C., 2003a. Identication and population dynamics of yeasts in sourdough fermentation processes by PCR-denaturing
gradient gel electrophoresis. Appl. Environ. Microbiol. 69, 7453e7461.
Meroth, C.B., Hammes, W.P., Hertel, C., 2004. Characterisation of the microbiota of
rice sourdoughs and description of Lactobacillus spicheri, sp. nov. Syst. Appl.
Microbiol. 27, 151e159.
Meroth, C.B., Walter, J., Hertel, C., Brandt, M., Hammes, W.P., 2003b. Monitoring the
bacterial population dynamics in sourdough fermentation processes by using
PCR-denaturing gradient gel electrophoresis. Appl. Environ. Microbiol. 69, 475e
482.
Meyer, S.A., Payne, R.W., Yarrow, D., 1998. Candida Berkout. In: Kurtzman, C.P.,
Fell, J.W. (Eds.), The Yeasts: a Taxonomic Study, fourth ed. Elsevier, Amsterdam,
The Netherlands, pp. 454e573.
Mihhalevski, A., Sarand, I., Viiard, E., Salumets, A., Paalme, T., 2011. Growth characterization of individual rye sourdough bacteria by isothermal microcalorimetry. J. Appl. Microbiol. 110, 529e540.
Minervini, F., De Angelis, M., Di Cagno, R., Pinto, D., Siragusa, S., Rizzello, C.G.,
Gobbetti, M., 2010. Robustness of Lactobacillus plantarum starters during daily
propagation of wheat our sourdough type I. Food Microbiol. 27, 897e908.
Please cite this article in press as: De Vuyst, L., et al., Microbial ecology of sourdough fermentations: Diverse or uniform?, Food Microbiology
(2013), http://dx.doi.org/10.1016/j.fm.2013.06.002
18
Minervini, F., Di Cagno, R., Lattanzi, A., De Angelis, M., Antonielli, L., Cardinali, G.,
Cappelle, S., Gobbetti, M., 2012a. Lactic acid bacterium and yeast microbiotas of
19 sourdoughs used for traditional/typical Italian breads: Interactions between
ingredients and microbial species diversity. Appl. Environ. Microbiol. 78, 1251e
1264.
Minervini, F., Lattanzi, A., De Angelis, M., Di Cagno, R., Gobbetti, M., 2012b. Inuence
of artisan bakery- or laboratory-propagated sourdoughs on the diversity of
lactic acid bacterium and yeast microbiotas. Appl. Environ. Microbiol. 78, 5328e
5340.
Miyashita, M., Yukphan, P., Chaipitakchonlatarn, W., Malimas, T., Sugimoto, M.,
Yoshino, M., Potacharoen, W., Tanasupawat, S., Nakagawa, Y., Kirtikara, K.,
Tanticharoen, M., Suzuki, K., 2012. 16S rRNA gene sequence analysis of lactic
acid bacteria isolated from fermented foods in Thailand. Microbiol. Cult. Collect.
28, 1e9.
Mora, D., Fortina, M.G., Parini, C., Daffonchio, D., Manachini, P.L., 2000. Genomic
subpopulations within the species Pediococcus acidilactici detected by multilocus typing analysis: relationships between pediocin AcH/PA-producing and
non-producing strains. Microbiol. 146, 2027e2038.
Moroni, A.V., Arendt, E.K., Dal Bello, F., 2011. Biodiversity of lactic acid bacteria and
yeasts in spontaneously-fermented buckwheat and teff sourdoughs. Food
Microbiol. 28, 497e502.
Moroni, A.V., Arendt, E.K., Morrissey, J.P., Dal Bello, F., 2010. Development of
buckwheat and teff sourdoughs with the use of commercial starters. Int. J. Food
Microbiol. 142, 142e148.
Mller, M.R.A., Wolfrum, G., Stolz, P., Ehrmann, M.A., Vogel, R.F., 2001. Monitoring
the growth of Lactobacillus species during a rye our fermentation. Food
Microbiol. 18, 217e227.
Neumann, H., Stephan, H., Brmmer, J.-M., 2006. Roggen als Rohstoff und Technik
der Roggensauerteigfhrung. In: Brandt, M.J., Gnzle, M.G. (Eds.), Handbuch
Sauerteig: Biologie, Biochemie, Technologie. B. Behrs Verlag GmbH & Co. KG,
Hamburg, Germany, pp. 185e283.
Neysens, P., Messens, W., De Vuyst, L., 2003. Effect of sodium chloride on growth
and bacteriocin production by Lactobacillus amylovorus DCE 471. Int. J. Food
Microbiol. 88, 29e39.
Nigatu, A., 2000. Evaluation of numerical analyses of RAPD and API 50 CH patterns
to differentiate Lactobacillus plantarum, Lact. fermentum, Lact. rhamnosus, Lact.
sake, Lact. parabuchneri, Lact. gallinarum, Lact. casei, Weissella minor and related
taxa isolated from kocho and tef. J. Appl. Microbiol. 89, 969e978.
Nigatu, A., Ahrne, S., Gashe, B.A., Molin, G., 1998. Randomly amplied polymorphic
DNA (RAPD) for discrimination of Pediococcus pentosaceus and Ped. acidilactici
and rapid grouping of Pediococcus isolates. Lett. Appl. Microbiol. 26, 412e416.
Nuraida, L., Wacher, M.C., Owens, J.D., 1995. Microbiology of pozol, a Mexican fermented maize dough. World J. Microbiol. Biotechnol. 11, 507e511.
Obiri-Danso, K., 1994. Microbiological studies on corn dough fermentation. Cereal
Chem. 71, 186e188.
Osimani, A., Zannini, E., Aquilanti, L., Mannazzu, I., Comitini, F., Clementi, F., 2009.
Lactic acid bacteria and yeasts from wheat sourdoughs of the Marche Region.
Ital. J. Food Sci. 21, 269e286.
Ottogalli, G., Galli, A., Foschino, R., 1996. Italian bakery products obtained with sour
dough: characterization of the typical microora. Adv. Food Sci. 18, 131e144.
Palomba, S., Blaiotta, G., Ventorino, V., Saccone, A., Pepe, O., 2011. Microbial characterization of sourdough for sweet baked products in the Campania region
(southern Italy) by a polyphasic approach. Ann. Microbiol. 61, 307e314.
Paramithiotis, S., Muller, M.R.A., Ehrmann, M.A., Tsakalidou, E., Seiler, H., Vogel, R.,
Kalantzopoulos, G., 2000. Polyphasic identication of wild yeast strains isolated
from Greek sourdoughs. Syst. Appl. Microbiol. 23, 156e164.
Paramithiotis, S., Tsiasiotou, S., Drosinos, E., 2010. Comparative study of spontaneously fermented sourdoughs originating from two regions of Greece: Peloponnesus and Thessaly. Eur. Food Res. Technol. 231, 883e890.
Parente, E., Ciocia, F., Ricciardi, A., Zotta, T., Felis, G.E., Torriani, S., 2010. Diversity of
stress tolerance in Lactobacillus plantarum, Lactobacillus pentosus and Lactobacillus paraplantarum: a multivariate screening study. Int. J. Food Microbiol. 144,
270e279.
Park, S.H., Itoh, K., 2005. Species-specic oligonucleotide probes for the detection
and identication of Lactobacillus isolated from mouse faeces. J. Appl. Microbiol.
99, 51e57.
Pepe, O., Blaiotta, G., Anatasio, M., Moschetti, G., Ercolini, D., Villani, F., 2004.
Technological and molecular diversity of Lactobacillus plantarum strains isolated
from naturally fermented sourdoughs. Syst. Appl. Microbiol. 27, 443e453.
Picozzi, C., Bonacina, G., Vigentini, I., Foschino, R., 2010. Genetic diversity in Italian
Lactobacillus sanfranciscensis strains assessed by multilocus sequence typing
and pulsed-eld gel electrophoresis analyses. Microbiology 156, 2035e2045.
Picozzi, C., DAnchise, F., Foschino, R., 2006. PCR detection of Lactobacillus sanfranciscensis in sourdough and Panettone baked product. Eur. Food Res. Technol.
222, 330e335.
Poutanen, K., Flander, L., Katina, K., 2009. Sourdough and cereal fermentation in a
nutritional perspective. Food Microbiol. 26, 693e699.
Pulvirenti, A., Caggia, C., Restuccia, C., Gullo, M., Giudici, P., 2001. DNA ngerprinting
methods used for identication of yeasts isolated from Sicilian sourdoughs.
Ann. Microbiol. 51, 107e120.
Pulvirenti, A., Solieri, L., Gullo, M., De Vero, L., Giudici, P., 2004. Occurrence and
dominance of yeast species in sourdough. Lett. Appl. Microbiol. 38, 113e117.
Randazzo, C.L., Heilig, H., Restuccia, C., Giudici, P., Caggia, C., 2005. Bacterial population in traditional sourdough evaluated by molecular methods. J. Appl.
Microbiol. 99, 251e258.
Ravyts, F., De Vuyst, L., 2011. Prevalence and impact of single-strain starter cultures
of lactic acid bacteria on metabolite formation in sourdough. Food Microbiol.
28, 1129e1139.
Reale, A., Di Renzo, T., Succi, M., Tremonte, P., Coppola, R., Sorrentino, E., 2011.
Identication of lactobacilli isolated in traditional ripe wheat sourdoughs by
using molecular methods. World J. Microbiol. Biotechnol. 27, 237e244.
Reale, A., Tremonte, P., Succi, M., Sorrentino, E., Coppola, R., 2005. Exploration of
lactic acid bacteria ecosystem of sourdoughs from the Molise region. Ann.
Microbiol. 55, 17e22.
Ricciardi, A., Parente, E., Piraino, P., Paraggio, M., Romano, P., 2005. Phenotypic
characterization of lactic acid bacteria from sourdoughs for Altamura bread
produced in Apulia (Southern Italy). Int. J. Food Microbiol. 98, 63e72.
Rizzello, C.G., Nionelli, L., Coda, R., De Angelis, M., Gobbetti, M., 2010. Effect of
sourdough fermentation on stabilisation, and chemical and nutritional characteristics of wheat germ. Food Chem. 119, 1079e1089.
Robert, H., Gabriel, V., Fontagn-Faucher, C., 2009. Biodiversity of lactic acid bacteria
in French wheat sourdough as determined by molecular characterization using
species-specic PCR. Int. J. Food Microbiol. 135, 53e59.
Rocha, J.M., Malcata, F.X., 1999. On the microbiological prole of traditional Portuguese sourdough. J. Food Prot. 62, 1416e1429.
Rocha, J.M., Malcata, F.X., 2012. Microbiological prole of maize and rye ours, and
sourdough used for the manufacture of traditional Portuguese bread. Food
Microbiol. 31, 72e88.
Rollan, G., Lorca, G.L., Font de Valdez, G., 2003. Arginine catabolism and acid
tolerance response in Lactobacillus reuteri isolated from sourdough. Food
Microbiol. 20, 313e319.
Rosenquist, H., Hansen, A., 2000. The microbial stability of two bakery sourdoughs
made from conventionally and organically grown rye. Food Microbiol. 17, 241e
250.
Ross, A.B., Kamal-Eldin, A., Aman, P., 2004. Dietary alkylresorcinols: absorption,
bioactivities, and possible use as biomarkers of whole-grain wheat- and ryerich foods. Nutr. Rev. 62, 81e95.
Saeed, M., Anjum, F.M., Zahoor, T., Nawaz, H., Rehman, S.U., 2009. Isolation and
characterization of starter culture from spontaneous fermentation of sourdough. Int. J. Agric. Biol. 11, 329e332.
Sakamoto, N., Tanaka, S., Sonomoto, K., Nakayama, J., 2011. 16S rRNA
pyrosequencing-based investigation of the bacterial community in nukadoko, a
pickling bed of fermented rice bran. Int. J. Food Microbiol. 144, 352e359.
Salovaara, H., Savolainen, J., 1984. Yeast type isolated from Finnish sour rye dough
starters. Acta Aliment. Pol. 10, 241e246.
Scheirlinck, I., Van der Meulen, R., De Vuyst, L., Vandamme, P., Huys, G., 2009a.
Molecular source tracking of predominant lactic acid bacteria in traditional
Belgian sourdoughs and their production environments. J. Appl. Microbiol. 106,
1081e1092.
Scheirlinck, I., Van der Meulen, R., Van Schoor, A., Cleenwerck, I., Huys, G.,
Vandamme, P., De Vuyst, L., Vancanneyt, M., 2007a. Lactobacillus namurensis sp.
nov., isolated from a traditional Belgian sourdough. Int. J. Syst. Evol. Microbiol.
57, 223e227.
Scheirlinck, I., Van der Meulen, R., Van Schoor, A., Huys, G., Vandamme, P., De
Vuyst, L., Vancanneyt, M., 2007b. Lactobacillus crustorum sp. nov., isolated from
two traditional Belgian wheat sourdoughs. Int. J. Syst. Evol. Microbiol. 57, 1461e
1467.
Scheirlinck, I., Van der Meulen, R., Van Schoor, A., Vancanneyt, M., De Vuyst, L.,
Vandamme, P., Huys, G., 2007c. Inuence of geographical origin and our type
on diversity of lactic acid bacteria in traditional Belgian sourdoughs. Appl.
Environ. Microbiol. 73, 6262e6269.
Scheirlinck, I., Van der Meulen, R., Van Schoor, A., Vancanneyt, M., De Vuyst, L.,
Vandamme, P., Huys, G., 2008. Taxonomic structure and stability of the bacterial
community in Belgian sourdough ecosystems as assessed by culture and population ngerprinting. Appl. Environ. Microbiol. 74, 2414e2423.
Scheirlinck, I., Van der Meulen, R., Vrancken, G., De Vuyst, L., Settanni, L.,
Vandamme, P., Huys, G., 2009b. Polyphasic taxonomic characterization of
Lactobacillus rossiae isolates from Belgian and Italian sourdoughs reveals
intraspecic heterogeneity. Syst. Appl. Microbiol. 32, 151e156.
Serrazanetti, D.I., Guerzoni, M.E., Corsetti, A., Vogel, R., 2009. Metabolic impact and
potential exploitation of the stress reactions in lactobacilli. Food Microbiol. 26,
700e711.
Settanni, L., Van Sinderen, D., Rossi, J., Corsetti, A., 2005. Rapid differentiation and in
situ detection of 16 sourdough Lactobacillus species by multiplex PCR. Appl.
Environ. Microbiol. 71, 3049e3059.
Siezen, R.J., Tzeneva, V.A., Castioni, A., Wels, M., Phan, H.T.K., Rademaker, J.L.W.,
Starrenburg, M.J.C., Kleerebezem, M., Molenaar, D., Vlieg, J.E.T.V., 2010. Phenotypic and genomic diversity of Lactobacillus plantarum strains isolated from
various environmental niches. Environ. Microbiol. 12, 758e773.
Siragusa, S., Di Cagno, R., Ercolini, D., Minervini, F., Gobbetti, M., De Angelis, M.,
2009. Taxonomic structure and monitoring of the dominant population of lactic
acid bacteria during wheat our sourdough type I propagation using Lactobacillus sanfranciscensis starters. Appl. Environ. Microbiol. 75, 1099e1109.
Spicher, G., 1959. Die Mikroora des Sauerteiges. I. Mitteilung: Untersuchungen
ber die Art der in Sauerteigen anzutreffenden stbchenfrmigen Milchsurebakterien (Genus Lactobacillus Beijerinck). Zeitblatt fur Bakteriologie II Abt 113,
80e106.
Spicher, G., 1984. Weitere Unterschungen ber die Zusammensetzung und die
Variabilitt der Mikroora handelsblicher Sauerteig-Starter. Z. Lebensm.
Unters. Forsch. 178, 106e109.
Please cite this article in press as: De Vuyst, L., et al., Microbial ecology of sourdough fermentations: Diverse or uniform?, Food Microbiology
(2013), http://dx.doi.org/10.1016/j.fm.2013.06.002
19
slopping propagation revealed Lactobacillus helveticus as the dominant species. J. Appl. Microbiol. 114, 404e412.
Villani, F., Moschetti, G., Blaiotta, G., Coppola, S., 1997. Characterization of strains of
Leuconostoc mesenteroides by analysis of soluble whole-cell protein pattern,
DNA ngerprinting and restriction of ribosomal DNA. J. Appl. Microbiol. 82,
578e588.
Vogelmann, S.A., Hertel, C., 2011. Impact of ecological factors on the stability of
microbial associations in sourdough fermentation. Food Microbiol. 28, 583e
589.
Vogelmann, S.A., Seitter, M., Singer, U., Brandt, M.J., Hertel, C., 2009. Adaptability of
lactic acid bacteria and yeasts to sourdoughs prepared from cereals, pseudocereals and cassava and use of competitive strains as starters. Int. J. Food
Microbiol. 130, 205e212.
Vrancken, G., De Vuyst, L., Van der Meulen, R., Huys, G., Vandamme, P., Daniel, H.M.,
2010. Yeast species composition differs between artisan bakery and spontaneous laboratory sourdoughs. FEMS Yeast Res. 10, 471e481.
Vrancken, G., Rimaux, T., De Vuyst, L., Leroy, F., 2008. Kinetic analysis of growth and
sugar consumption by Lactobacillus fermentum IMDO 130101 reveals adaptation
to the acidic sourdough ecosystem. Int. J. Food Microbiol. 128, 55e66.
Vrancken, G., Rimaux, T., Weckx, S., Leroy, F., De Vuyst, L., 2011. Inuence of temperature and backslopping time on the microbiota of a type I propagated laboratory wheat sourdough fermentation. Appl. Environ. Microbiol. 77, 2716e
2726.
Weckx, S., Allemeersch, J., Van der Meulen, R., Vrancken, G., Huys, G., Vandamme, P.,
Van Hummelen, P., De Vuyst, L., 2011. Metatranscriptome analysis for insight
into whole-ecosystem gene expression during spontaneous wheat and spelt
sourdough fermentations. Appl. Environ. Microbiol. 77, 618e626.
Weckx, S., Van der Meulen, R., Allemeersch, J., Huys, G., Vandamme, P., Van
Hummelen, P., De Vuyst, L., 2010a. Community dynamics of sourdough fermentations as revealed by their metatranscriptome. Appl. Environ. Microbiol.
76, 5402e5408.
Weckx, S., Van der Meulen, R., Maes, D., Scheirlinck, I., Huys, G., Vandamme, P., De
Vuyst, L., 2010b. Lactic acid bacteria community dynamics and metabolite
production of rye sourdough fermentations share characteristics of wheat and
spelt sourdough fermentations. Food Microbiol. 27, 1000e1008.
Weiss, N., Schillinger, U., 1984. Lactobacillus sanfrancisco sp. nov., nom. rev. Syst.
Appl. Microbiol. 5, 230e232.
Wick, M., Stolz, P., Bcker, G., Lebeault, J.M., 2003. Inuence of several process
parameters on sourdough fermentation. Acta Biotechnol. 23, 51e61.
Zannini, E., Garofalo, C., Aquilanti, L., Santarelli, S., Silvestri, G., Clementi, F., 2009.
Microbiological and technological characterization of sourdoughs destined for
bread-making with barley our. Food Microbiol. 26, 744e753.
Zapparoli, G., De Benedictis, P., Salardi, C., Veneri, G., Torriani, S., Dellaglio, F., 1996.
Lactobacilli of sourdoughs from Verona bakery: a preliminary investigation.
Adv. Food Sci. 18, 163e166.
Zapparoli, G., Torriani, S., 1997. Rapid identication and detection of Lactobacillus
sanfrancisco in sourdough by species-specic PCR with 16S rRNA-targeted
primers. Syst. Appl. Microbiol. 20, 640e644.
Zapparoli, G., Torriani, S., Dellaglio, F., 1998. Differentiation of Lactobacillus sanfranciscensis strains by randomly amplied polymorphic DNA and pulsed-eld
gel electrophoresis. FEMS Microbiol. Lett. 166, 325e332.
Zhang, J., Liu, W., Sun, Z., Bao, Q., Wang, F., Yu, J., Chen, W., Zhang, H., 2011. Diversity
of lactic acid bacteria and yeasts in traditional sourdoughs collected from
western region in Inner Mongolia of China. Food Control 22, 767e774.
Zotta, T., Parente, E., Ricciardi, A., 2009. Viability staining and detection of metabolic
activity of sourdough lactic acid bacteria under stress conditions. World J.
Microbiol. Biotechnol. 25, 1119e1124.
Zotta, T., Piraino, P., Parente, E., Salzano, G., Ricciardi, A., 2008. Characterization of
lactic acid bacteria isolated from sourdoughs for Cornetto, a traditional bread
produced in Basilicata (Southern Italy). World J. Microbiol. Biotechnol. 24,
1785e1795.
Please cite this article in press as: De Vuyst, L., et al., Microbial ecology of sourdough fermentations: Diverse or uniform?, Food Microbiology
(2013), http://dx.doi.org/10.1016/j.fm.2013.06.002