Food Microbiology 26 (2009) 676–684

Contents lists available at ScienceDirect

Food Microbiology
journal homepage: www.elsevier.com/locate/fm

Sourdough in gluten-free bread-making: An ancient technology to solve

a novel issue?
Alice V. Moroni a, b, Fabio Dal Bello a, b, Elke K. Arendt a, *
a
Department of Food Science, Food Technology and Nutrition, National University of Ireland, Cork, Ireland
b
National Food Biotechnology Centre, National University of Ireland, Cork, Ireland

a r t i c l e i n f o a b s t r a c t

Article history: The increasing demand for high quality gluten-free (GF) bread, clean labels and natural products is
Received 1 May 2009 raising the need for new approaches in GF bread-making. Sourdough is the foremost fermentation used
Received in revised form for baking purposes and it has been proven to be ideal for improving the texture, palatability, aroma,
19 June 2009
shelf life and nutritional value of wheat and rye breads. These characteristic features derive from the
Accepted 6 July 2009
Available online 26 July 2009
complex metabolic activities of the sourdough-resident lactic acid bacteria and yeasts, e.g. acidification,
production of exopolysaccharides, proteolytic- amylolytic- and phytase activity, and production of
antimicrobial substances. These effects have been extensively studied and well described for traditional
Keywords:
Gluten-free bread baking, whereas little is known about the role of sourdough in GF baking. Yet, the microbiological and
Sourdough qualitative characterisation of local GF fermented products indicate an overlap with the microbiota of
wheat/rye fermentation and suggest that the positive metabolic activities of the sourdough microbiota
are still retained during fermentation of GF crops. Thus, the use of sourdough in GF baking may be the
new frontier for improving the quality, safety and acceptability of GF bread.
Ó 2009 Elsevier Ltd. All rights reserved.

1. Celiac disease and the sensory quality of gluten-free bread
Celiac disease is an immune-mediated enteropathy triggered by
the ingestion of gluten in genetically susceptible individuals, and it
affects about 1% of the world population (Catassi and Fasano, 2008).
Currently, a lifelong exclusion of gluten from the diet is the
cornerstone treatment for celiac disease and only gluten-free (GF)
products can be consumed by celiac. GF foods are dietary foods
either consisting of ingredients which do not contain wheat, kamut,
spelt, rye, barley, oats, or made of ingredients from such cereals,
which must have been specially processed to remove gluten.
Overall, the gluten level in GF products does not exceed 20 mg/kg
in total (Deutsch, 2009). Yet, gluten-containing products, such
as bread and pasta, represent a dietary staple for a large portion
of the world population. Thus, the development of GF products
suitable for celiac patients represents a compelling, but also very
demanding call for food technologists. In particular, some
commercially available GF breads are characterised by low quality,
exhibiting poor crumb and crust characteristics as well as poor
mouth feel and flavour (Gallagher et al., 2004, 2003a). Being mainly
* Corresponding author at: Department of Food and Nutritional Sciences,
University College Cork, Western Road, Cork, Ireland. Tel.: þ353 21 4902064; fax:
þ353 21 4270213.
E-mail address: e.arendt@ucc.ie (E.K. Arendt).
based on starch, these products have very low contents of
various nutrients and undergo fast staling (Ahlborn et al., 2005). In
most cases, commercially available GF products are not fortified
(Niewinski, 2008) and contain lower levels of some B vitamins, iron,
folate and dietary fibre when compared to their gluten-containing
counterparts (Thompson, 2000; Yazynina et al., 2008). Since clinical
evidences indicate that malabsorption of several important
nutrients is associated with celiac disease (Feighery, 1999; Robins
et al., 2008; Sabry and Okada, 1992), the rigorous consumption of
commercially available GF products may yet worsen the already
unbalanced diet of celiac patients (Mariani et al., 1999).
Over the last 20 years, cereal technologists have tried to over-
come these issues and to produce GF bread that could meet the
expectations of celiac consumers. Various additives and combina-
tions of different nutritive ingredients have been investigated and
are currently used in GF bread formulations in order to improve
both technological and nutritional quality of the bread. The incor-
poration of nutrient-dense whole grains, such as amaranth, buck-
wheat, millet, quinoa, brown rice, sorghum and teff, in GF bread
formulations has recently shown the potential of increasing the
nutritional value of these products, especially in terms of fibre,
protein and mineral contents (Alvarez-Jubete et al., 2009; Moore
et al., 2004; Pagano, 2006; Repo-Carrasco-Valencia et al., 2009;
Yazynina et al., 2008). Iron fortified amaranth-based bread with
satisfactory sensory characteristics was also recently produced by

0740-0020/$ – see front matter Ó 2009 Elsevier Ltd. All rights reserved.
doi:10.1016/j.fm.2009.07.001
 A.V. Moroni et al. / Food Microbiology 26 (2009) 676–684
Kiskini et al. (2007). Likewise, the dietary fibre content of starch-
based GF bread was successfully increased by incorporation of
prebiotics oligosaccharides (Gallagher et al., 2002; Korus et al., 2006).
From a technological point of view, gluten replacement with GF
flours is not a trivial task. In fact, gluten possesses unique visco-
elastic properties which are crucial for the water holding capacity
of the dough and the gas retention during fermentation (Arendt
et al., 2008). Furthermore, the lack of standardized baking tests for
GF batters makes the research even more challenging. Hydrocol-
loids are currently used to improve the viscoelastic properties of GF
batters (Lazaridou et al., 2007), because of their potential to form
three-dimensional polymer network in aqueous solutions (BeMil-
ler, 2008). Many studies have been performed on the exploitation of
various hydrocolloids, such as hydroxypropril-methyl-cellulose,
carbossi- or methyl-cellulose, locust bean and guar gum, xanthan
and pectins in GF bread formulations (Ahlborn et al., 2005; Cato
et al., 2002; McCarthy et al., 2005; Moore et al., 2004; Pruska-
Kedzior
˛ et al., 2008; Schober et al., 2005; Schwarzlaff et al., 1996).
According to the flours and ingredients used, specific hydrocolloids
can affect to different extents the loaf volume and crumb firmness
of the final bread, and methyl celluloses seems to be the most
effective among all (Gujral et al., 2003b; Lazaridou et al., 2007;
Schober et al., 2008). Addition of proteins, e.g. dairy powders or
eggs, was also shown to improve the quality of GF bread. Dairy
ingredients are used in bread for nutritional benefits, including
increased calcium content and protein efficiency ratio, and for
functional benefits, such as flavour and texture enhancement as
well as storage improvement (Kenny et al., 2001). Incorporation of
milk proteins in GF bread formulations resulted in a gluten-like
matrix in the GF bread, and in turn it enhanced loaf volume,
improved crumb texture and delayed staling of the bread (Ahlborn
et al., 2005; Gallagher et al., 2003b; Moore et al., 2004). However,
Schober et al. (2005) outlined that in case of sorghum bread, the
addition of skim milk powder has negative effects on bread quality.
Regardless of the effects on bread quality, the use of milk and egg
proteins has some limitations. As a matter of fact, lactose intoler-
ance in celiac patients is a common pathology (Ojetti et al., 2005),
and thus only low lactose dairy powders can be used in GF bread
formulations (Nunes et al., 2009). Furthermore, allergens from
egg belong to some of the most frequent causes of food allergic
reactions (Poulsen et al., 2001).
The use of enzymes for improving the quality of GF bread has
also been the subject of extensive research. Rice bread with
increased specific volume and very soft crumb texture was
obtained by the addition of cyclodextrin glycosyl transferase
(Gujral et al., 2003a) or glucose oxidase (Gujral and Rosell, 2004).
Protein functionality has also been modified by the use of trans-
glutaminase (TGase). TGase was added to a rice flour-based bread
with addition of milk, soybean, peas and/or egg proteins (Marco
and Rosell, 2008; Moore et al., 2006). The enzyme treatment
improved the functional properties of the enriched rice flour and
enhanced the protein network formation, thus improving the
crumb structure of the resulting breads. Recently, Renzetti et al.
(2008) investigated the effects of TGase on different GF batters.
Among all GF flours tested, buckwheat or brown rice-based breads
were the most positively affected by the enzymatic treatment.
Despite the promising results obtained so far, the incorporation
of additives in bread formulations for improving the quality of GF
bread presents several disadvantages. First of all, excessive prices:
most of the additives used represent high-cost ingredients, and the
average price of GF bread on the market is 10–20 folds higher than
that of conventional bread (http://www.glutenfree.com/baked-
goods/breads/category357). Additionally, some of these ingredi-
ents, e.g. proteins, may be cause of allergies, and concerns have
been recently addressed on the use of enzymes in foods (Gerrard
677
and Sutton, 2005). Furthermore, additives do not meet the
consumers’ requirements for natural products. Thus, an impelling
call for researchers to find an alternative technology for producing
good quality GF bread is rising. In this review we address how
sourdough can represent the tool to solve all the different problems
related to the production of high quality GF bread.
2. Sourdough bread
Sourdough is a mixture of flour and water fermented with lactic
acid bacteria (LAB) and yeasts, which determine its characteristics
in terms of acid production, aroma and leavening (Hammes and
Gänzle, 1998; Vogel et al., 1999). Sourdough is a unique food
ecosystem: it selects for LAB and yeasts which are adapted to the
environment, and hosts highly specific microbial communities
(De Vuyst and Vancanneyt, 2007; Gobbetti et al., 2008). The use of
sourdough has a long tradition in the production of wheat and rye
breads. In particular, acidification induced by LAB is a prerequisite
for baking rye bread, since it promotes the solubilisation of rye
pentosans, which determine the structure of rye bread, and it also
inhibits amylase activity (De Vuyst and Vancanneyt, 2007). When
used in optimised proportions, sourdough can improve volume,
texture, flavour, nutritional value of bread and increase the shelf life
by retarding the staling process and by protecting bread from
mould and bacterial spoilage (De Vuyst and Vancanneyt, 2007;
Gobbetti et al., 2008). These positive effects are associated with the
metabolic activities of sourdough-resident microorganisms, such as
lactic acid fermentation, proteolysis, exopolysaccharides (EPS)
production and synthesis of volatile and antimicrobial compounds
(Corsetti and Settanni, 2007; Arendt et al., 2007).
The positive contribution of sourdough could be exploited for
the production of high quality GF bread from various GF cereals and
pseudocereals, with reduced need for expensive additives (Table 1).
Yet, sourdough bread is still a niche product consumed mostly in
central and eastern Europe, and thus its acceptability could repre-
sent an issue for those consumers who are not used to it. However,
this problem may be overcome by the selection of appropriate
starter cultures and process parameters for obtaining GF sourdough
breads with sensorial characteristics that best meet the consumers’
requirements. Unfortunately, only few attempts have been made
for producing and characterising GF sourdoughs and the functional
properties of the breads thereof (Di Cagno et al., 2008; Edema and
Sanni, 2008; Moore et al., 2007, 2008; Sanni et al., 1998; Schober
et al., 2007; Vogelmann et al., 2009).
3. Ecology of GF sourdoughs in cereals products
With the advent of novel molecular biological tools, the
microbial diversity of sourdough has been well described (De Vuyst
Table 1
Quality defects of GF bread and possible improvements due to sourdough fermen-
tation (adapted from Gobbetti et al., 2008).
GF bread: challenges Sourdough in bread: positive effects
Dry crumbling texture Improved dough softening
Low loaf volume Increase of bread volume
Improved gas retention
Poor mouth feel Improved palatability
Poor flavour Release of flavour compounds
Short shelf-life Decrease bread staling
Antiropiness and antifungal activity
Low nutritional value Improved mineral bioavailability
Decreased glycemic index
Prebiotic exopolysaccharides
High costs – expensive Reduced need for expensive ingredients,
ingredients e.g. hydrocolloids, enzymes, chemical preservatives
678 A.V. Moroni et al. / Food Microbiology 26 (2009) 676–684
and Vancanneyt, 2007). Microbiological studies have revealed that
more than 70 species of LAB and more than 25 species of yeasts
occur in wheat and rye sourdoughs. Species belonging to the genus
Lactobacillus are among the most adapted to the sourdough
fermentation, but also species of the genera Leuconostoc,
Pediococcus, and Weissella are found in association with sourdough
(De Vuyst and Vancanneyt, 2007; Hammes et al., 2005). Overall,
wheat and rye microbial communities are comparable, thus sug-
gesting that the process parameters rather than the raw materials
are the key factors determining the establishment of a particular
microbial community (De Vuyst and Vancanneyt, 2007; Gänzle and
Schwab, 2009; Scheirlinck et al., 2008). Indeed, Meroth et al.
(2003a) showed that a variation in the fermentation temperature
from 30 to 40  C strongly influenced the competitiveness of LAB in
rye sourdough, and unique communities were detected under the
different incubation conditions.
Fermented food products produced from rice, maize, sorghum,
and other GF cereals can be found in tropical climates (Hammes
et al., 2005; Nout and Sarkar, 1999). Several GF cereal fermentations
have been characterised on microbial and biochemical level,
showing that LAB and yeasts are the dominating organisms of these
fermentations (Table 2). However, the most common species found
in traditional (wheat and rye) sourdough fermentations, namely
Lactobacillus sanfranciscensis, Lactobacillus panis, Lactobacillus pon-
tis and Lactobacillus spicheri, are not present in fermentations
carried out in tropical climates (Gänzle and Schwab, 2009). Instead,
the more thermophilic Lactobacillus fermentum and Lactobacillus
reuteri are commonly found in these fermentations. Unfortunately,
there is a major drawback when analysing such studies, since often
these works lack information about the process parameters, and
bacterial/yeasts species were sometimes identified only by physi-
ological criteria. Moreover, in the case of maize and sorghum, most
of the research has been focused on traditional products, often
obtained by local producers, hence being their microbiological
composition affected by the flours used and by the environment
where the fermentations were carried out.
Recently, Vogelmann and co-workers (Vogelmann et al., 2009)
investigated the adaptability of a great variety of LAB and yeasts to
sourdoughs prepared from the cereals wheat, rye, oat, barley, rice,
maize and millet, from the pseudocereals amaranth, quinoa, and
Table 2
Microbiota of gluten-free sourdough and fermentations for traditional products.
Raw material Dominant microbiotaa
Product
Sorghum
Sudanese kisra L. fermentum, L. reuteri, L. vaginalis, L. helveticus, L. pontis,
P. pentosaceus, I. orientalis
b
Sudanese khamir P. pentosaceus, L. brevis, Lact. lactis, L. cellobiosus, C. parapsilosis,
C. orvegnsis, R. glutinis
Botswana-fermented L. plantarum, L. casei/paracasei, L. buchneri, L. reuteri, L. perolens
sorghum porridge
Rice
Sourdough L. paracasei, L. paralimentarius, L. perolens, L. spicheri, S. cerevisiae
Sourdough L. gallinarum, L. plantarum, L. helveticus, L. fermentum, L. kimchii,
L. pontis, I. orientalis, S. cerevisiae
Maize
Kenkey L. fermentum, L. reuteri, P. pentosaceus, C. krusei, S. cerevisiae
Mexican pozol L. delbrueckii, L. casei, L. fermentum, L. plantarum, Streptococcus spp.,
Leuconostoc spp., Weisella spp.
Sourdough L. brevis, L. casei, L. fermentum, L. plantarum, Lc. mesenteroides,
L. dextranicum, P. acidilactici, C. halbicans, S. cerevisiae, S. pombe
Sourdough L. fermentum, L. paralimentarius, L. helveticus, L. pontis,
S. cerevisiae, I. orientalis
a
I., Issatchenkia; L., Lactobacillus; Lact., Lactococcus; P., Pediococcus; R., Rhodotorula; S., Saccharomyces.
b
Addition of spices.
buckwheat as well as from cassava. In general, the dominant strains
belonged to the species L. fermentum, Lactobacillus helveticus,
Lactobacillus paralimentarius, Lactobacillus plantarum, L. pontis, L.
spicheri, Issatchenkia orientalis and Saccharomyces cerevisiae.
However, even if each substrate was started with the same starter
mixture and fermented under the same technological conditions,
competitiveness of some strains was found to be substrate-specific.
For example, L. paralimentarius was highly competitive in pseudo-
cereals, whereas L. helveticus dominated in cereals, and some
species, such as L. spicheri, dominated only few substrates. Thus, not
only the technological parameters, but also the type and quality of
the substrates and the interactions between the starter’s micro-
organisms and the natural microbiota were suggested to strongly
affect the establishment of the dominant microbiota of sourdoughs
(Vogelmann et al., 2009). The results of this study indicate the need
of developing and designing starter mixtures/fermentation condi-
tion specific for the different substrates and products. The species
isolated from the GF sourdoughs must be classified with regard to
their metabolic performances – production of acids, aroma
compounds, EPS and antimicrobial compounds – both qualitatively
and quantitatively, i.e., specific compounds have to be produced in
active concentrations. Only with the availability of such informa-
tion it will be possible to produce GF sourdoughs bread of constant
and high quality. Nonetheless, considering the diversity of LAB and
yeasts isolated from GF cereal fermentations, and the overlap with
the microbiota of wheat and rye sourdoughs, the statement the
entire metabolic activity of LAB and yeasts can be employed for the
improvement of bread quality can be extended to include GF baked
goods (Gänzle et al., 2008).
4. Starch hydrolysis
Starch retrogradation is one of the key events involved in bread
staling, and this becomes even more evident in GF breads, which
are mainly based on starch (Zobel and Kulp, 1996). Sourdough
fermentation retards wheat bread staling (Barber et al., 1992), and it
has been shown to reduce the glycemic index of rye bread (Katina
et al., 2005; Scazzina et al., 2009). Corsetti et al. (1998, 2000)
showed that in sourdough bread staling is influenced by the
biological acidification and also by the proteolytic and amylolytic
Fermentation Reference
starter/spontaneous
Spontaneous Hamad et al., 1992, 1997;
Mohammed et al., 1991
Spontaneous Gassem, 1999
Spontaneous Sekwati-Monang, Gänzle,
unpublished
Spontaneous Meroth et al., 2003b
Starter Vogelmann et al., 2009
Spontaneous Jespersen et al., 1994; Olsen et al., 1995;
Hayford et al., 1999
Spontaneous Ampe et al., 1999; Ben Omar and Ampe, 2000;
Escalante et al., 2001
Spontaneous Sanni et al., 1998; Edema and Sanni, 2008
Starter Vogelmann et al., 2009
 A.V. Moroni et al. / Food Microbiology 26 (2009) 676–684
activities of the selected starters. Even though most LAB do not
possess amylolytic activities, amylolytic LAB have been isolated
from cereal fermentations in tropical climates (Gänzle et al., 2008;
Sanni et al., 2002; Tou et al., 2007). Recently, Songré-Ouattara et al.
(2009) suggested that amylolytic LAB can be used as starter culture
for starch hydrolysis in pearl millet gruels, if starch is pre-gelati-
nised by cooking. Noticeably, milling malted grains produces flours
containing starch more susceptible to enzymatic hydrolysis than
flours from non-malted grains. Hugo (Hugo et al., 2003) showed
that malting, boiling and also fermentation of sorghum flour could
decrease crumb firmness, dryness and grittiness of sorghum-wheat
composite bread, and in particular, fermentation resulted in highest
loaf volume. Thus, the exploitation of amylolytic LAB for producing
GF sourdough bread represents a possibility to control bread
staling. To achieve optimal results, malted GF grains should be
included in the GF bread formulation.
5. Phytase activity
Phytic acid is the major storage form of phosphorous in grains
and it strongly binds metallic cations of Ca, Fe, K, Mg, Mn and Zn,
making them insoluble and thus unavailable for nutrition (Bohn
et al., 2008). Phytic acid is therefore considered an antinutritional
factor, especially for celiac patients who suffer micronutrient
deficiencies. During bread-making phytic acid is degraded by
phytases, which are influenced by temperature, pH, fermentation
time and additives (Lopez et al., 2001). To this regard, sourdough
fermentation can influence mineral bioavailability. Fermentation
creates optimal pH conditions for the activation of endogenous
phytases and for increased mineral solubility, and some sourdough-
associated LAB may exert phytase activity (De Angelis et al., 2003).
In wheat and especially in whole wheat bread, sourdough
fermentation was shown to enhance phytate hydrolysis and
mineral solubility in respect to conventional yeast fermentation
(Lopez et al., 2001). Recently, L. sanfranciscensis CB1 has been
characterised for its high level of phytase activity by De Angelis and
co-workers (De Angelis et al., 2003). When the strain was used for
starting sourdough fermentation with wheat flour, a major decrease
of over 50% of the phytate level was found in comparison to the
unstarted dough. Unfortunately, no work has been published on
phytase activity in GF sourdough breads. Yet, phytase activity has
been investigated during fermentation of some GF crops. In particular,
fermentation of sorghum (Osman, 2004) and pearl millet was shown
to induce a decrease in the phytic acid content, and two phytase-
positive strains, i.e., L. plantarum and L. fermentum, were isolated from
fermenting pearl millet (Songré-Ouattara et al., 2009).
6. Proteolysis
Protein degradation during sourdough fermentation is among
the key phenomena that affect the overall quality of sourdough
bread (Gänzle et al., 2008). The proteolytic events occurring in
wheat or rye sourdoughs have been exhaustively illustrated by
Gänzle et al. (2008). Overall, LAB play a minor role in protein
hydrolysis (Wieser et al., 2008). Most sourdough-associated LAB,
and in particular L. sanfranciscensis, do not possess extracellular
proteinase activity and prefer peptides uptake rather than amino
acids transport (Thiele et al., 2003). Thus, primary proteolysis is
exerted by wheat or rye endogenous enzymes, which are activated
by the low pH. Further hydrolysis of peptides into amino acids is
then exerted by intracellular peptidases of LAB in a strain-specific
manner: the type and amount of released amino acids depend on
the fermenting strain (Di Cagno et al., 2002). Though LAB do not
influence the overall amount of hydrolysed proteins in respect to
aseptically acidified doughs, they affect the pattern of hydrolysed
679
products, increasing the amount of dipeptides and amino acids
(Di Cagno et al., 2002; Thiele et al., 2003). Indeed, the proteolysis
exerted by LAB induce softening of the dough in comparison to
chemically acidified doughs (Di Cagno et al., 2002).
Proteolysis occurs in the GF product towga, a traditional Tan-
zanian fermented food prepared by fermentation of either
sorghum, maize, cassava, millet or a combination thereof. Mugula
et al. (2003) showed that towga fermentation resulted in break-
down of sorghum proteins, but no distinction between the activity
of endogenous- and LAB-proteinases was outlined. Recently, sour-
dough fermentation was effectively applied for the production of
GF bread based on sorghum and potato starch (Schober et al.,
2007). In order to improve bread quality, hydroxypropyl-methyl-
cellulose (HPMC) was added in bread formulation, but only when
sourdough fermentation of the sorghum flour was performed,
superior quality bread was obtained. Water soluble proteins were
degraded into small peptides during fermentation and, as shown by
confocal laser scanning microscopy, no protein aggregation was
induced in bread crumb upon baking, whereas protein aggregates
were present in the chemically acidified bread. Sourdough
fermentation also resulted in the formation of a stronger starch gel
during cooking of the bread, as shown by rheological tests. There-
fore, it was concluded that the proteolytic events occurring during
sourdough fermentation on water soluble proteins are desirable for
producing high quality GF sorghum bread. Accordingly, Elkhalifa
et al. (2006) showed that only water soluble proteins were
hydrolysed by fermentation of sorghum flour for preparation of
kisra, a typical Sudanese food. As shown by light and confocal laser
scanning microscopy, the hydrolysis of water soluble proteins
resulted in the release of small starch granules, thus affecting the
properties of fermented sorghum.
Proteolysis can improve the quality of GF bread, as recently
shown for sorghum bread. However, considering the number and
variety of GF flours available, further studies are required to better
understand the role and impact of proteins and proteolysis during
GF bread-making. As recently shown by Renzetti and Arendt
(2009), proteolysis improves the quality of some GF bread, e.g. oats
and brown rice, but can also lead to no improvements or even to
a loss of quality of other GF breads.
7. Proteolysis for reducing toxicity of wheat flour
During endoluminal digestion, gliadins of wheat, rye and barley
release a family of peptides rich in Pro and Gln, which are
responsible for the autoimmune response in celiac patients
(Anderson et al., 2000; Kendall et al., 1972). Recently, LAB-induced
proteolysis has been suggested as an efficient method for
decreasing toxicity of wheat and rye flours. Gliadins in wheat are
among the most affected proteins by sourdough fermentation and
the extent of hydrolysis of monomeric gliadins (a-, b-, g-, u-glia-
dins) is strain-specific (Di Cagno et al., 2002; Wieser et al., 2008). Di
Cagno et al. (2002) showed that selected proteolytic LAB could
efficiently hydrolyse the 31–43 fragment of the toxic peptide A-
gliadin in wheat sourdough. On the basis of these results, the same
authors developed a novel bread technology (Di Cagno et al., 2004).
They showed that selected LAB could completely hydrolyse the
highly toxic 33-mer peptide over prolonged (12–24 h) and semi-
liquid fermentation of a mixture of wheat and non-toxic flours.
Breads produced with 12 h-sourdough fermentation still retained
acceptable quality and when consumed by celiac individuals, no
alterations on the baseline values could be observed. The selected
LAB were also successfully used for the detoxification of rye sour-
dough (De Angelis et al., 2006a). Additionally, the same pool of LAB
was used in association with L. sanfranciscensis strains and fungal
proteases for the production of detoxified wheat sourdough, which
680 A.V. Moroni et al. / Food Microbiology 26 (2009) 676–684
also showed to be suitable for bread production (Rizzello et al.,
2007). The probiotic preparation VLS#3 also showed potential of
decreasing the toxicity of wheat flour (De Angelis et al., 2006b).
Giuliani et al. (2006) also showed that fermentation of GF ingre-
dients could degrade the gluten eventually present as contaminant
in such ingredients. Therefore, prolonged fermentation times with
selected LAB of wheat/rye flour and their combination with GF
flours might represent a novel technology for baking good quality
GF breads However, long-term in vivo tests are needed to confirm
the suitability of these products for celiac patients, and the
consumer acceptance towards a GF product containing wheat/rye
could still represent a major issue for marketing of these products.
8. Exopolysaccharides
Many food grade microorganisms produce exopolysaccharides
(EPS) (De Vuyst and Degeest, 1999). EPS act as biothickeners that
can be added to a variety of food products, where they serve as
viscosifying, stabilising, emulsifying or gelling agents (Tieking and
Gänzle, 2005). They are distinguished in two classes: homo-
polysaccharides (HoPS), mainly glucan or fructans polymers, and
heteropolysaccharides, with (ir)regular repeating units (De Vuyst
and Degeest, 1999. Heteropolysaccharides are mainly used in dairy
products, instead only HoPS are effective in bread-making. HoPS
are synthesised from sucrose by glucosyltransferares or fructosyl-
transferases, respectively, and various HoPS-producers are found
among cereal-associated LAB (Di Cagno et al., 2006; Tieking and
Gänzle, 2005). Indeed, HoPS producing LAB have been successfully
used in wheat sourdough fermentations (Tieking et al., 2003). The
structure and molecular weight of HoPS varies among the
producing LAB species, and the production yield also depends on
the carbohydrate concentration and composition of the source (Di
Cagno et al., 2006; Korakli et al., 2003). Glucan and fructans
produced by fermenting LAB can strongly influence the quality of
wheat bread in terms of bread volume and crumb firmness (Di
Cagno et al., 2006; Lacaze et al., 2007). In particular, the production
of EPS in situ is more effective than addition of the same poly-
saccharide in the bread formulation (Brandt et al., 2003).
Together with EPS, LAB can also produce gluco- or fructo-oligo-
saccharides (FOS) (Monsan et al., 2001), among which FOS, together
with the fructan inulin, have been well described for their prebiotic
effects (Biedrzycka and Bielecka, 2004; Cummings et al., 2001). In
addition, the levan produced by L. sanfranciscensis was proved to
stimulate bifidobacterial growth in vitro (Dal Bello et al., 2001). In
sourdough, L. reuteri, Lactobacillus acidophilus and L. sanfranciscensis
showed the ability to produce the prebiotic FOS 1-kestose (Korakli
et al., 2003; Tieking and Gänzle, 2005). In summary, the production
of HoPS by sourdough-LAB exerts two different functions, i.e., while
improving bread structure, it improves the nutritional value of
bread. Both features are extremely interesting for GF bread-making.
Even though EPS production in wheat and rye sourdoughs is
extensively described (Tieking and Gänzle, 2005), little information
is available on EPS formation and their role in GF sourdoughs.
Nonetheless, there is a widely accepted consensus on the possibility
of using EPS produced by sourdough-LAB as replacers for the
expensive hydrocolloids. One major concern is, however, the ability
of LAB to produce the EPS in considerable amounts while fer-
menting a GF substrate. Furthermore, studies investigating the
rheological performances of EPS in comparison to those of
commercially available hydrocolloids are required. Only recently,
the formation of EPS and FOS was investigated in GF sourdoughs
(Schwab et al., 2008). In sorghum sourdoughs, L. reuteri LTH5448
and Weissella cibaria 10M produced amounts of levan and dextran,
respectively, which were comparable to those measured in wheat
and rye sourdoughs. FOS produced by W. cibaria were not digested
by baker’s yeast in the batter, and therefore they were still available
in the final bread. Remarkably, improvements in the GF bread
quality were achieved only when W. cibaria-fermented sourdough
was added in the formulation, indicating that dextrans, rather than
levans, are suitable for replacing hydrocolloids in GF bread-making.
However, since the type (molecular weight, degree of branching
and type of linkages) and origin of the EPS as well as the interac-
tions with the matrix strongly influence its rheological behaviour
(Lacaze et al., 2007), more research is needed for determining the
EPS performance in GF bread and for identifying the most suitable
EPS for hydrocolloids replacement.
9. Aroma compounds
The flavour of leavened baked goods is influenced by raw
materials, by sourdough fermentation, by the type of starters, and
by proofing and baking conditions (Gobbetti, 1998). Upon the
whole process of bread production, from flour to bread, flavouring
compounds can be either removed or generated. Microbial and
enzymatic conversions of carbohydrates, amino acids and lipids in
the dough create the flavour compounds relevant for the crumb
odour, i.e., alcohols, esters and carbonyls, whereas crust odour is
influenced by the thermal reactions occurring during baking. An
equilibrated aroma can be reached by using the right combinations
of LAB and yeasts (Hansen and Schieberle, 2005). The ratio between
lactic acid and acetic acid is an important factor affecting the aroma
of the final bread (Corsetti and Settanni, 2007), and it is influenced
by the fermenting microorganism, the fermentation temperature
and the type of flour (Hansen and Schieberle, 2005).
A limited extent of proteolysis during sourdough fermentation
beneficially improves bread flavour without adverse effects on
texture and volume (Thiele et al., 2002). LAB can catalyse reactions
such as deamidation, transamination and decarboxylation, and their
amino acids metabolism also contribute to the flavour. As an example,
glutamine is converted to glutamate and then accumulated during
sourdough fermentation, imparting taste to the bread (Gänzle et al.,
2007). The expression of the arginine deiminase pathway in Lacto-
bacillus spp. promotes higher production of ornithine, and thus
enhances the formation of 2-acetyl pyrroline, which is responsible for
the roasty note of wheat bread crumb (Gänzle et al., 2007). Along with
proteins, also lipids can greatly affect bread quality. Lipids oxidation,
which occurs during flour storage and mixing of the dough, induces
the formation of (E)-2-nonenal and other aldehydes which are key
aroma compounds in wheat and rye breads (Hansen and Schieberle,
2005; Vermeulen et al., 2007). During sourdough fermentation, the
concentration of these aldehydes is reduced by alcohol dehydroge-
nase activity of LAB, and especially by heterofermentative LAB (Ver-
meulen et al., 2007). Yeasts also exert alcohol dehydrogenase activity
in sourdough, and Candida spp. can accelerate oxidation of fatty acids
by formation of hydrogen peroxide (Hansen and Schieberle, 2005).
Recently, fermentation was shown to induce production of fla-
vouring compounds also in GF products. For example, fermentation
of sorghum for production of towga generated different flavouring
compounds (Mugula et al., 2003). Alcohols were produced in high
concentration when fermentation was carried out with I. orientalis
in combination with Lactobacillus brevis or L. plantarum, and
diacetyl was produced in significant amounts by L. plantarum and
Pediococcus pentosaceus. The production of aldehydes was increased
by co-fermentation with L. plantarum and yeasts. Fermentation with
mixed cultures containing L. plantarum enhanced the diacetyl
content in maize fermented meals (Edema and Sanni, 2008). Thus,
even if further studies are required, it is not speculative to assume
that sourdough fermented by LAB and yeasts can be used to produce
flavour enhancing compounds which will improve the poor
sensorial quality of GF breads.
 A.V. Moroni et al. / Food Microbiology 26 (2009) 676–684 681

10. Shelf life shown to produce in active concentrations reutericyclin, a low

Contamination of bread occurs after baking and airborne
distribution of dust and mould spores is the main cause for bread
spoilage (Legan, 1993). In addition to economical losses, bread
spoilage also represents a health hazard for the consumers, espe-
cially when bread is contaminated with mycotoxigen moulds.
Modified atmosphere packaging, irradiation and addition of
preservatives are among the most commonly used tools for
prevention of bread spoilage (Corsetti and Settanni, 2007).
However, there is a definite need for other safe and efficient ways
for preventing bread spoilage in order to respond to the increasing
demand for natural, high quality products. Sourdough represents
a natural tool to improve the shelf life of bread, since it can prevent
microbial spoilage and also retard bread staling. Over the last
decade, several studies have described the antifungal compounds
produced by (sourdough)-LAB and their activities against common
bread spoilage organisms. Sourdough-associated LAB produce
many antimicrobial substances, such as organic acids, CO2, ethanol,
hydrogen peroxide, diacetyl, fatty acids, phenyllactic acid, reuterin
and fungicins (Messens and De Vuyst, 2002; Schnürer and
Magnusson, 2005). Among the organic acids, acetic and propionic
acid produced by heterofermentative LAB are more effective than
lactic acid (Schnürer and Magnusson, 2005). Caproic acid produced
by L. sanfranciscensis CB1, together with a mixture of acetic, formic,
propionic, butyric, and n-valeric acids, play a key role in inhibiting
Fusarium, Penicillium, Aspergillus and Monilia growth in bread
(Corsetti et al., 1998). Also L. plantarum shows a very broad
antimicrobial activity, and the antifungal compounds 4-hydrox-
yphenlyllactic and especially phenyllactic acids were identified
as responsible for fungal inhibition (Dal Bello et al., 2007;
Lavermicocca et al., 2003; Ryan et al., 2009). A synergistic effect was
found when sourdough fermented with antifungal L. plantarum
strains was used in combination with calcium propionate (CAP) for
production of wheat bread (Ryan et al., 2008). In particular, the
presence of sourdough allowed reducing CAP levels by around 30%,
without affecting the shelf life of bread. In sourdough, L. reuteri was
molecular weight antibiotic acting against Gram-positive LAB and
yeasts (Gänzle et al., 2000). L. reuteri strains have also been shown
to produce reuterin, an antimicrobial substance active against
bacteria, yeasts and fungi (Gänzle, 2004). Sourdough-associated
LAB are also effective against rope spoilage of bread induced by
Bacillus spp., probably due to production of organic acids and other
still unknown antibacterial substances (Katina et al., 2002; Valerio
et al., 2008). However, since the metabolic activity of micro-
organisms is affected by the environment, in particular by the
fermentation substrate, studies are required to verify if production
of these antimicrobial compounds occurs also during GF sourdough
fermentation.
To date, only one work has investigated the use of sourdough to
extend the shelf life of GF bread. In this study, Moore et al. (2008)
used the antifungal strain L. plantarum FST 1.7 (Dal Bello et al.,
2007) to produce GF sourdough from a mixture of brown rice, corn
starch, buckwheat, and soya flours. Fermenting 20% of the GF flours
with L. plantarum FST 1.7 resulted in delaying the onset of staling in
respect to chemically acidified GF control breads. Furthermore, the
antifungal activity of L. plantarum was retained during fermenta-
tion of GF flours, and when 20% sourdough was added in the bread
formulation, the growth of Fusarium culmorum was retarded up to 3
days in comparison to the control breads. This study, thus, indicate
that production of antifungal GF sourdough can be regarded as
a promising and valuable alternative to the use of additives for
prolonging shelf life and retarding staling of GF breads. However,
further research is needed to identify the optimal starter/s and
fermentation conditions for achieving GF bread of extended shelf
life and high quality.
11. Conclusions
Sourdough fermentation positively influences all aspects of
bread quality: texture, aroma, nutritional properties and shelf life.
The data collected so far suggest that these positive effects can be
exploited also in GF bread production, and thus, the application of

Cornbread
Buttermilk (cultured fat-free milk, salt, vitamin
A palmitate, vitamin D3), cornmeal, eggs,
canola oil, evaporated cane juice, potato
starch, cornstarch, salt, baking powder (sodium
acid pyrophosphate, sodium bicarbonate,
cornstarch, monocalcium phosphate), xanthan

Prairie bread
Nonfat milk, rice flour, tapioca Sandwich bread
starch, eggs, evaporated cane Non fat milk, rice flour,
juice, canola oil, yeast, millet, tapioca starch, eggs,
buckwheat, pumpkin seeds, evaporated cane juice, canola
xanthan gum, salt, sunflower oil, yeast, salt, xanthan gum,
seeds, sesame seeds, flax, poppy lemon juice
seeds, lemon juice

GF bread

Sourdough bread
Sourdough (GF flours, malted
flour), (yeast), salt, palm oil

Fig. 1. Composition of commercially available gluten-free (GF) breads, compared to the composition of hypothetical GF sourdough bread.
682 A.V. Moroni et al. / Food Microbiology 26 (2009) 676–684
the sourdough technology represents the new frontier for the
production of high quality GF bread. Sourdough GF bread could also
be the answer to the demand for clean labels, natural products and
for a reduced use of additives (Fig. 1).
Acknowledgments
This study was financially supported by the European
Commission in the Communities 6th Framework Programme,
Project HEALTHGRAIN (FP6-514008). This publication reflects only
author’s views and the Community is not liable for any use that may
be made of the information contained in this publication. The
authors acknowledge that this research was partly funded also by
FIRM Ireland.
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