Trends in Food Science & Technology 17 (2006) 557e566

Review

Flavour in sourdough
breads: a review 1995; Onno & Rouseel, 1994; Ottogalli, Galli, & Foschino,
1996). The key groups of fermenting organisms, in addition
to the yeasts, are strains of the lactic acid bacteria (LAB).
Salim-ur-Rehman*, Alistair In traditional sourdough systems initially part of the flour
is mixed with all the yeast and sufficient water to make
Paterson and John R. Piggott ‘‘sponge’’ that is allowed to ferment for some hours, typi-
cally overnight, exposed to the atmosphere. The sponge is
Centre for Food Quality, Department of Bioscience, then mixed with rest of the flour, water, salt and fat to a suit-
University of Strathclyde, 204 George Street, Glasgow able consistency, and then given a short period for fermen-
G1 1XW, Scotland, UK (Permanent address: Institute tation before final proving and baking. The outcome is
of Food Science and Technology, University of considered to be bread of a richer flavour due to bacterial
Agriculture, Faisalabad, Punjab 38000, Pakistan. souring of the sponge through airborne contamination or
Fax: D92 41 9201105; D44 141 553 4124; incorporation of 2e5% of sponge from the previous batch
e-mail: [email protected]) as a microbial inoculum (Bruemmer & Lorenz, 1991).
In good bakery practice, a sponge should contain meta-
Flavour compounds are key elements for consumer accep- bolically active lactic acid bacteria at 108e109 cfu g
1 and
tance and product identification in bread. One category of yeasts at 106e107 cfu g
1, primarily responsible for acidifi-
speciality breads, the sourdoughs have a fermentation process cation and leavening action of dough, respectively. How-
affected by a complex microflora of yeasts and lactic acid ever, the LAB may either originate from natural flour
bacteria which confer specific flavour characteristics. Al- contaminant, a fermented dairy product or from a commer-
though yeasts have the primary leavening role, lactic acid cial starter culture containing characterised strains of LAB
bacteria (LAB), with trophic and non-trophic relationships, produced in batch fermentation (Vuyst & Neysens, 2005).
produce important flavour components. Sourdoughs are be-
coming important as consumers move away from pan breads
towards speciality products. However, successful new product Microorganisms of sourdough sponges
development requires an understanding of variations in carbo- The microbial ecology of the sourdough fermentation is
hydrates’ metabolism, roles of endogenous enzymes and inter- complex and varied with identification of more than 50 spe-
actions of microorganisms for generation of non-volatile and cies of LAB, mostly of the genus Lactobacillus, and more
volatile flavour compounds. The potential of sourdough baking than 20 species of yeasts, are dominated by the genera
remains to be developed through specifications and optimisa- Saccharomyces and Candida. Sourdough microflora have
tions of process conditions and introduction of exogenous primarily stable associations with lactobacilli and yeasts,
enzymes and other ingredients. With effective new product having important metabolic interactions contributing to-
development sourdough characteristics could be matched to wards production of flavour compounds (Vuyst & Neysens,
relate with consumer tastes. 2005). Sourdough flavour components identified as pro-
duced by yeast (Table 1) and LAB (Table 2) fall into
many classes of compounds.
Several yeasts are found in sourdoughs but Saccharomy-
Introduction
ces cerevisiae is considered the dominant organism for
Sourdough is an important modern fermentation method
leavening of bread (Corsetti et al., 2001) (Table 1). It has
of cereal flours and water (Vogel et al., 1999) based upon
been suggested that numbers of S. cerevisiae are overesti-
the earlier spontaneous process. A slack flour dough is
mated due to the lack of robust systems for quantifying
inoculated with microbial starter, ‘‘mother culture’’, which
individual yeast in this habitat (Vogel, 1997). Important
is constantly renewed in a cyclical way, using specified
yeasts in sourdough starters include Saccharomyces exiguus
recipes and ripening conditions (Hammes & Ganzle,
(physiologically similar to Candida milleri), Candida kru-
sei, Pichia norvegensis and Hansenula anomala. Other
* Corresponding author. yeasts isolated from sourdough sponges include
0924-2244/$ - see front matter Ó 2006 Elsevier Ltd. All rights reserved.
doi:10.1016/j.tifs.2006.03.006
558 S.-ur-Rehman et al. / Trends in Food Science & Technology 17 (2006) 557e566

Table 1. Volatile and non-volatile compounds present in wheat flour sourdough fermented with various yeast strains (Damiani et al., 1996)

Compounds A B C D F
Lactic acid





Acetic acid





Ethanol þ

þ þ
1-Propanol þ þ
þ þ
2-Methyl-1-propanol þ

þ þ
Ethyl acetate þ þ þ þ þ
3-Methyl-1-butanol þ

þ þ
2-Methyl-1-butanol þ
þ þ þ
1-Pentanol





2-Methyl-1-pentanol





1-Hexanol þ
þ þ þ
3-Hexen-1-ol





1-Heptanol





1-Octanol





Acetaldehyde þ þ þ þ þ
3-Methyl-1-butanal þ þ þ þ þ
2-Methyl-1-butanal þ
þ þ þ
Hexanal þ þ þ þ þ
3-Methyl-hexanal





Heptanal





Trans-2-heptenal





Octanal þ þ þ þ þ
Nonanal þ þ þ þ þ
Benzaldehyde





Diacetyl þ




Hexane þ þ þ þ þ
Heptane
þ þ


Octane
þ þ


þ, Present and
, not present.
A e Saccharomyces cerevisiae; B e Candida krusei; C e Candida norvegensis; D e Saccharomyces exiguus; and E e Hansenula anomala.

Saccharomyces delbrueckii, Torulopsis holmii and Torulop- sourdoughs (Gül et al., 2005). In wheat sourdough, mixed
sis unisporus (Gül, Özçelik, Sadıç, & Certel, 2005). cultures were estimated as: Lb. Sanfranciscensis e 20%;
It is generally considered that in sourdoughs, the ratio of Lb. Alimentarius e 14%; Lb. Brevis e 12%; Leuconostoc
LAB to yeast should be 100:1 for optimal activities citreum e 7%; Lb. Plantarum e 6%; Lactococcus lactis
(Gobbetti, Corsetti, Rossi, La Rosa, & De Vincenzi, 1994). subsp. lactis e 4%; Lb. fermentum, Lb. acidophilus and
Although, Saccharomyces exiguus is a dominant species in Weissella confusa e each 2%; Lb. delbrueckii subsp. del-
starters, strains do not ferment maltose and grow by fermenting brueckii (Corsetti et al., 2001). Although, Lb. sanfrancisco
glucose released by heterofermentative lactic acid bacteria. In is considered a key sourdough lactic acid bacterium
return, the yeast supplies an electron source, fructose, to the (Gobbetti & Corsetti, 1997) in rice sourdough, Lb. fermen-
LAB that assist both growth and acetic acid production: a tum, Lb. gallinarum, Lb. kimchii, Lb. plantarum, Lb. pontis,
co-operation with important technological and sensory conse- Lb. paracasei, and Lb. paralimentarius have been reported
quences. Production of acetic acid improves quality in leavened to dominate (Merotha, Hammesa, & Hertela, 2004).
dough and other bakery products (Vernocchi et al., 2004). Mixed cultures of LAB and yeasts vary in composition
Typical sourdough prokaryotes are members of the LAB in sourdough sponges. The use of mixed cultures has a num-
genus Lactobacillus with obligately homofermentative, and ber of important advantages, such as improved flavour and
facultatively or obligately heterofermentative strains with texture and retained freshness for longer compared to
strains of Lb. sanfranciscensis (Catzeddu, Mura, Parente, baker’s yeast bread (Meignen et al., 2001). In such mixed
Sanna, & Farris, 2006) (synonym Lb. brevis subsp. lindneri), cultures, yeasts act mainly as leavening agents, while
Lb. plantarum and Lb. brevis most frequently isolated LAB contribute mainly to the flavouring compounds of
(Trüper & de Clari, 1997). Certain strains, initially classified bread. Kefir is a natural mixed culture which can produce
as Lb. brevis, are recently allotted to the new species bread of good quality. Many microorganisms have been iso-
Lb. Pontis (Vogel et al., 1994). Other LAB strains including lated from kefir microflora, sharing symbiotic relationships
Carnobacterium divergens (Lb. Divergens, Lb. amylophilus, including yeasts (Kluyveromyces, Candida, Torulopsis and
Lb. sake, Lb. acetotolerans, L. plantarum, Pediococcus Saccharomyces sp.), lactobacilli (Lb. brevis, Lb. acidophi-
pentosaceus and P. acidilactici, and Tetragenococcus halo- lus, Lb. casei, L. helveticus, and Lb. delbrueckii), strepto-
philus (Pediococcus halophilus) have been isolated from cocci (Streptococcus salivarius) lactococci (Lc. Lactis ssp.
 S.-ur-Rehman et al. / Trends in Food Science & Technology 17 (2006) 557e566 559

Table 2. Volatile and non-volatile compounds present in wheat flour sourdough fermented with hetero- (AeE) and homofermentative (FeJ) LAB
strains (Damiani et al., 1996)

Compounds A B C D E F G H I J
Lactic acid þ þ þ þ þ þ þ þ þ þ
Acetic acid þ þ þ þ þ þ þ



Ehanol þ þ þ þ þ





1-Propanol þ









2-Methyl-1-propanol










Ethyl acetate þ þ þ þ þ þ þ þ þ þ
3-Methyl-1-butanol










2-Methyl-1-butanol










1-Pentanol










2-Methyl-1-pentanol þ þ








1-Hexanol þ þ þ þ þ þ þ þ þ þ
3-Hexen-1-ol þ









1-Heptanol þ



þ þ



1-Octanol þ þ



þ



Acetaldehyde þ þ þ þ þ þ þ þ þ þ
3-Methyl-1-butanal þ



þ þ



2-Methyl-1-butanal










Hexanal þ þ þ þ þ þ þ þ þ þ
3-Methyl-hexanal




þ




Heptanal þ þ
þ
þ þ



Trans-2-heptenal þ þ


þ þ



Octanal þ þ þ þ þ þ þ þ þ þ
Nonanal þ þ þ þ þ þ þ þ þ þ
Benzaldehyde þ



þ þ



Diacetyl




þ þ þ þ þ
Hexane þ þ þ þ þ þ þ
þ þ
Heptane þ þ þ þ
þ þ þ þ þ
Octane þ þ þ þ
þ þ þ þ þ
þ, Present and
, not present.
A e Lactobacillus brevis lindneri; B e Lactobacillus brevis; C e Lactobacillus fructivorans; D e Lactobacillus fermentum; E e Lactobacillus
cellobiosus; F e Lactobacillus plantarum; G e Lactobacillus farciminis; H e Lactobacillus alimentarius; I e Lactobacillus acidophilus; and
J e Lactobacillus delbrueckii.

thermophilus, Leuconostoc mesenteroides and L. cremoris) lambica, three each as C. krusei and Candida valida and two
and occasionally acetic acid bacteria (Plessas, Pherson, as Candida glabrata were identified (Succi et al., 2003).
Bekatou, Nigam, & Koutinas, 2005; Simova et al., 2002).
The phenotypic characterisation of the isolates identifies
the presence of 80% as facultative heterofermentative Factors affecting sourdough volatiles
LAB (Lb. plantarum, Lb. paracasei, Lb. casei) and 12% and non-volatile compounds
as heterofermentative LAB (Lb. brevis and L. mesenter- Sourdough sponges vary in proportions of LAB in dif-
oides) in sourdough of Altamura bread (Ricciardi, Parente, ferent speciality breads. The primary metabolic products
Piraino, Paraggio, & Romano, 2005). of LAB are L-lactic and acetic acids with lesser amounts
The conditions and substrates of the sourdough sponge are of citric and malic acids. Ratio of lactic acid to acetic
fundamental with respect to microbiological stability of acid is important for final product flavour (Linko, Javanainen,
dough. Mixed commercial starters containing Lb. brevis & Linko, 1997). LAB multiply and produce lactic and acetic
and S. cerevisiae (baker’s yeast), fermented at 30  C for acids more slowly in the mixed culture with yeasts than in
20 h, showed reduced yeast growth and ethanol production monocultures (Merseburger, Ehret, Geiges, Baumann, &
in dough but more glycerol (80%) and acetic acid (55%) Schmidt-Lorenz, 1995).
whereas lactic acid production was unchanged (Meignen Heterofermentative LAB species can be discriminated
et al., 2001). In contrast to the production of durum wheat from homofermentative on the basis of volatile compounds
bran flour sourdough breads, more than 95% strains were (Table 2). Certain aldehydes are derived both from enzy-
of the yeasts Candida humilis and C. krusei and S. cerevisiae matic oxidation or autoxidation of the lipid fraction of
were dominant in rice sourdough (Merotha et al., 2004). the wheat (Frankel, 1982; Hann & Morrison, 1975) and
While in Triticum aestivum wheat flour sourdough, 58 strains bacterial metabolism. Homofermentative LAB, with the
as S. cerevisiae, five as Candida colliculosa, four as Candida exception of Lb. delbrueckii subsp. delbrueckii strains are
560 S.-ur-Rehman et al. / Trends in Food Science & Technology 17 (2006) 557e566

primary contributor of ethyl acetate and diacetyl (Hansen & due to the greater flour enzyme activity, which increases the
Hansen, 1993; Hansen, Lund, & Lewis, 1989a). availability of soluble carbohydrates (Röcken & Voysey,
The composition of sourdough flavour compounds is, 1993). In wheat sourdough fermentations, maltose remains
however, not only influenced by microbial composition, between 2 and 5 g/kg (Martinez-Anaya, Pitarch, & Bene-
but also interactions between bread making processes and dito de Barber, 1993) as yeast preferentially depletes avail-
ingredients (Collar, 1996). These are important for process able glucose and fructose (Barber, Baguena, Benedito de
stability and production of variety of regional specialities Barber, & Martinez-Anaya, 1991).
that meet consumer and commercial demands (Gobbetti, Co-fermentations are other metabolic strategies to
1998). In sourdoughs produced from Triticum aestivum enable sourdough LAB to use non-fermentable substrates,
(Succi et al., 2003) and Triticum durum wheat flours, a num- increasing adaptability. Co-fermentation of fructose and
ber of lactic acid bacteria and yeasts range from ca. log 7.5 maltose or glucose has been observed specifically in a
to log 9.3 colony forming units (cfu)/g and from log 5.5 to fructose-negative strain of Lb. Sanfranciscensis (Gobbetti,
log 8.4 cfu/g, respectively (Corsetti et al., 2001). In slack Corsetti, & Rossi, 1995; Stolz, Böcker, Hammes, & Vogel,
doughs, there are changes in yeast and lactic acid bacterial 1995), as has co-metabolism of citrate and maltose or glu-
growth with more abundant flavour compounds. Stimulat- cose (Gobbetti & Corsetti, 1996). Moreover, changes in the
ing effects on yeasts of NaCl can indirectly be attributed availability of citrate and fructose can influence the ratios
to a reduction in competition with lactic acid bacteria for of acetic acid and lactic acid and micro-structural rheolog-
available sugars (Neysens, Messens, & Vuyst, 2003), and ical features of the doughs (Gianotti et al., 1997), through
with content of other soluble carbohydrates, in the cereal change in acetic acid production (Calderon, Loiseau, &
flours at typical fermentation temperatures. Guyot, 2003). Certain strains of S. cerevisiae are sensitive
Ingredients in sourdough bread influence generation of to the acetic acid produced by LAB, especially at the
volatile compounds, volatile compounds of germinated, normal sourdough pH of 4.0e4.5 that favours the undisso-
sourdough fermented, and native rye substantially different, ciated lipophilic and membrane-diffusible form of the
variable even after the second treatment (Heiniö et al., organic acid. Thus, it is necessary to add baker’s yeast
2003). Type of cereal flours, endogenous and exogenous to compensate for poor survival of wild-type yeasts in
cereal components and processing steps such as the heat consecutive sourdough fermentations (Suihko & Makinen,
treatment in baking has significant effects on the generation 1984).
of flavouring compounds in breads (Hansen, 1995). One key Lb. sanfranciscensis hydrolyses maltose and accumu-
driver of flavour is the carbon source available to the micro- lates glucose in the medium in a molar ratio of about 1 malt-
organisms, notably low-molecular weight sugars that is, at ose to 1 glucose (Gobbetti, Corsetti et al., 1994; Gobbetti,
least in part, flavour precursors (Martinez-Anaya, 1996). Simonetti et al., 1994; Stolz, Böcker, Vogel, & Hammes,
Sucrose addition to wheat dough stimulates both yeast 1993). A study of maltose uptake and glucose excretion
and LAB growth and increases bacterial production of in Lb. sanfranciscensis (Neubauer, Glaasker, Hammes,
lactic and acetic acids (Corsetti, Gobbetti, & Rossi, 1994). Poolman, & Konings, 1994) showed that once the maltose
However, titratable acidity (TTA) of the dough increases is depleted, consumption of the excreted glucose is initi-
in proportion to the addition of sucrose over the range of ated. Glucose excreted during fermentation may be used
0e6%, ascribed to acetic acid accumulation which affects by maltose-negative yeasts such as S. exiguus or induced
the generation of flavouring compounds (Simonson, glucose repression of maltose catabolism in competitors
Salovaar, & Korhola, 2003). San Francisco sourdough from using maltose, thereby giving an ecological advantage
breads are produced with specific cultures of Lactobacillus to Lb. sanfranciscensis (Nout & Creemers-Molenaar, 1987).
(Linko et al., 1997; Seitz, Chung, & Rengarajan, 1998). However, among the Lactobacillaceae, only certain species e
A lack of competition between Lb. sanfranciscensis and Lb. sanfranciscensis, Lb. pontis, Lb. reuteri, and Lb.
S. exiguus for maltose consumption was noted which is fun- fermentum (Vogel et al., 1994) e phosphorylate maltose
damental for the stability of both the microorganisms for and the enzyme maltose phosphorylase may be considered
the production of good quality San Francisco French bread the key for LAB growth during sourdough fermentation.
(Sugihara, Kline, & McCready, 1970). However, LAB Conversely, with sucrose and Lb. plantarum and S. cer-
growth and lactic and acetic acid productions may be re- evisiae or S. exiguus in co-culture (Gobbetti, Corsetti, &
duced by more rapid consumption of maltose and, notably, Rossi, 1994a) cell yield and lactic acid production increase
of glucose by S. cerevisiae in an association with Lb. san- (Robert et al., 2006). Invertase hydrolysis of sucrose by
franciscensis in synthetic media (Collar, 1996). Even so, yeasts into glucose and fructose, may enhance LAB metab-
imbalances between yeast consumption and starch hydroly- olism (Aksu & Kutsal, 1986) because yeasts can hydrolyse
sis by endogenous flour enzymes lead to rapid depletion of sucrose to about 200 times faster than the released mono-
soluble carbohydrates during wheat sourdough fermenta- saccharides are fermented (Martinez-Anaya, 1996), with
tion which, in turn, decreases LAB acidification due to rapid sucrose depletion during sourdough fermentation
microbial competition (Rouzaud & Martinez-Anaya, 1993). (Seppi, 1984). S. exiguus preferentially utilises glucose or
This situation is less pronounced in rye dough fermentation sucrose and shows a high tolerance for the acetic acid
 S.-ur-Rehman et al. / Trends in Food Science & Technology 17 (2006) 557e566 561

from heterolactic LAB metabolism (Suihko & Makinen, (Spicher, Rabe, Sommer, & Stephan, 1981; Spicher, Rabe,
1984). Sommer, & Stephan, 1982).
Flavour component release may also be influenced by Temperature optima for co-culture of C. humilis and
the environmental conditions such as temperature. Yeast Lb. sanfranciscensis are 28 and 32  C, respectively, but
and LAB growth increase in a direct relationship with reduced production of acetate by Lb. sanfranciscensis has
temperature between 15  C and 27  C. Optimal growth been observed at 35  C, although, both lactate and ethanol
temperature of the two Candida milleri strains was between formations are not affected at this temperature (Brandt,
26  C and 28  C in monoculture. Nevertheless, decrease in Hammes, & Ganzle, 2004). Lactic acid production varied
the growth of yeast may be accompanied by a similar from 3.11 to 5.14 g/kg and traces of acetic acid may be pro-
reduction in the growth of LAB (Neysens et al., 2003; duced by some of the Lb. plantarum and Lb. farciminis
Simonson et al., 2003). strains during fermentation (Table 1).
Processing conditions such as proofing time, tempera-
ture and slackness of sourdough may also affect the aroma Volatile compounds
volatiles. Low temperature (25  C) and sourdough firmness Microbial metabolisms verify production of different
are considered appropriate for LAB souring activities but volatile compounds for hetero- and homo-lactic LAB
limited yeast metabolism. Raising the temperature to (Table 1) fermentations. Abundant products of yeast fer-
30  C and semi-fluid sourdough fermentation can generate mentation are 2-methyl-1-propanol, 2,3 methyl-1-butanol
more complete volatile profiles. While at 3 h fermentation, and other iso-alcohols. Heterofermentative LAB products
the sourdough may mainly be characterised by iso-alcohols are dominated by ethyl acetate and certain alcohols and
but an increase of leavening time of up to 9 h can produce aldehydes (Spicher et al., 1982) whereas major homofer-
volatiles about three times higher than that at 5 h as a result mentative LAB products are diacetyls and carbonyls
of LAB contribution (Lund, Hansen, & Lewis, 1989). (Spicher et al., 1981) (Table 2).
Lactobacillus brevis subsp. Lindneri and Lactobacillus
Types of flavouring compounds in sourdough breads plantarum have been considered to have the most appropri-
There are two categories of flavour compounds, produced ate profiles of flavour components. (Gobbetti, Corsetti, &
during sourdough fermentation. Non-volatile compounds De Vincenzi, 1995a). However, with the exception of
including organic acids produced by homo- (Gobbetti, Lb. plantarum DC400eSaccharomyces exiguus M14 asso-
Corsetti, & De Vincenzi, 1995a) and heterofermentative bac- ciation, both hetero- and homofermentative LAB enhance
teria (Gobbetti, Corsetti, & De Vincenzi, 1995b) which acidify, yeast formation volatile compounds (Damiani et al., 1996;
decrease pH and contribute aroma to the bread dough (Barber, Gobbetti, Corsetti, & De Vincenzi, 1995b).
Benedito de Barber, Martinez-Anaya, Martinez, & Alberola, The LAB strains of sourdough vary in metabolism and
1985; Galal, Johnson, & Varriano-Marston, 1978). aroma compounds. Monoculture fermentation of dough
The second category e volatile compounds of sour- for 15 h at 30  C, followed by mixing and a further 10 h
dough bread e includes alcohols, aldehydes, ketones, esters fermentation has shown increase in the production of typi-
and sulphur. All these compounds are produced by bio- cal sourdough volatile compounds (Merotha et al., 2004).
logical and biochemical actions during fermentation and The production of volatile aroma compounds during
contribute to flavour (Spicher, 1983). fermentation with combinations of single strains of S. cer-
evisiae and C. guilliermondii, Lb. plantarum has been
Non-volatile compounds investigated using both wheat doughs and simple substrates
Interactions between lactic acid bacteria and exogenous including maltose and glucose (Stolz et al., 1993). Using
enzymes influence microbial kinetics of acidification, only yeasts in wheat bread, seven volatiles were found
acetic acid production and bread textural characters. It has abundant: acetaldehyde, acetone, ethyl acetate, ethanol,
been shown to be desirable to augment endogenous hexanal, isobutyl alcohol, and propanol. In this fermenta-
enzyme activities in flour and has been obtained from tion, S. cerevisiae produced more volatiles than C. guillier-
LAB in association with microbial glucose oxidase, lipase, mondii, but quantity of volatile flavour compounds can be
endo-xylanase, -amylases and proteases. The growth of improved by the addition of glucose and sucrose less by
Leuconostoc citreum 23B, Lb. lactis subsp. lactis 11M and maltose. It is still not clear whether yeasts produce more
Lb. hilgardii 51B has been influenced by enzyme addition, flavouring components than LAB in simple substrate which
which increases lactic acid production (Cagno et al., appeared to produce a greater range of volatiles from wheat
2003). However, in a continuous sourdough fermentation, flour (Hansen, 1995; Torner, Martı́nez-Anaya, Antuña, &
an association between Lb. sanfranciscensis and S. cerevi- Benedito de Barber, 1992; Vollmar & Meuser, 1992).
siae has showed optimal production of acetic acid (Vollmar Addition of enzymes to sourdough sponges can also
& Meuser, 1992), whereas Torulopsis holmii improved enhance bread volatile compounds (Martinez-Anaya,
dough acidification in association with Lb. sanfranciscensis 1996), lipid oxidations through addition of enzymes in the
and S. cerevisiae and further enhanced acid production if form of active soya flour increase concentrations of hexanal,
associated with Lb. sanfranciscensis and Lb. plantarum 1-hexanol, 1-penten-3-ol, 1-pentanol and 2-heptanone, and
562 S.-ur-Rehman et al. / Trends in Food Science & Technology 17 (2006) 557e566

2-heptenal and 1-octen-3-ol have only been reported in of bread (Bredie, Mottram, & Guy, 2002; Damiani et al.,
breads containing soya flour (Luning, Roozen, Moëst, & 1996). Maillard reaction products from high temperature
Posthumus, 1991). Addition of lipase, endo-xylanase and processing include pyrazines, pyrroles, furans and sulphur-
a-amylases enhanced acetic acid production by Lb. hilgardii containing compounds and lipid degradation products such
51B. Textural analyses suggest that such sourdoughs with as alkanals, 2-alkenals and 2,4-alkadienals (Parker, Hassell,
single enzyme have greater stability and crumb softening Mottram, & Guy, 2000). Moreover, at higher temperatures
than breads produced with only exogenous enzymes (Cagno pyrroles, thiophenes, thiophenones, thiapyrans and thiazolines
et al., 2003). Moreover, addition of fructose and citrate to increase and furans and aldehydes decrease (Bredie et al.,
sourdough enhanced the production of volatiles by LAB 2002). The pyrazines are nitrogen-containing ring compounds
including ethanol, which was eliminated in baking and that have extremely powerful odour and flavour properties
2-methyl-1-propanal with no effect on lactic and acetic (Ji and Berhard, 1992). Hence, the production of these com-
acids’ production (Gobbetti, Corsetti, & De Vincenzi, 1995a; pounds has a striking effect on the flavour of bread.
Gobbetti & Corsetti, 1996; Gobbetti, Smacchi, Fox,
Stepaniak, & Corsetti, 1996).
In a French yeast sourdough, more than 40 flavour com- Sensory characteristics of bread
ponents were identified: 20 alcohols, 7 esters, 6 lactones, 6 The sourdough fermentation is central to acceptability in
aldehydes, 3 alkanes and a single sulphur compound flavour, as chemically acidified bread and breads prepared
(Frasse, Lambert, Richard-Molard, & Chiron, 1993). Com- with pure commercial starter cultures are not well scored
ponents, except for aldehydes and alkanes, increased in in sensory preference assessments (Lund et al., 1989; Rothe
abundance due to greater activity of S. cerevisiae notably & Ruttloff, 1983). The synergistic metabolic activities of
those related to formation of fusel alcohols. However, the microorganisms produce an acidification or souring
in this fermentation 2,3 butanedione, 3-methyl-1-butanol, influencing the final character in the bread, notably the tex-
2-methyl 1-butanol, methional, 2-phenylethanol and 2 ture, increased shelf-life by reducing the mould growth
unidentified compounds with a pungent and mushroom during storage (Corsetti et al., 2000; Oura, Soumalainen,
odour were also apparent. & Wiskari, 1982; Röcken & Voysey, 1995) and generate
A number of other factors influence the production of typical flavour components yielding typical sourdough
volatile compound during production of sourdough bread sensory attributes (Gobbetti, 1998; Katina et al., 2005).
including free amino acids formed during fermentation The flavour of sourdough wheat bread is richer and more
through proteolysis which influence volatile compound pro- aromatic than in wheat bread, a factor that can be attributed
file (Collar & Martinez, 1993; Collar, Mascaros, Prieto, & to the long fermentation time of sourdough (Bruemmer &
Benedito de Barber, 1991; Gobbetti, Corsetti, & Rossi, Lorenz, 1991). The most desirable sensory characteristics
1994b; Gobbetti et al., 1996; Spicher & Nierle, 1984). are obtained at pH 4.0e5.5 and at 140  C (Przybylski &
Thiazolines, thiophenes, thiophenones, thiaziles, polythia- Kamiński, 1983). The loaves made with the addition of
cycloakanes, pyrroles and pyrazines have been shown to 5e10% sourdoughs fermented with Lb. plantarum and
relate to acidity in wheat extrusion cooking (Hansen & 5e15% sourdoughs fermented with Lb. sanfrancisco are
Hansen, 1994a) and sourdough fermentation (Hansen preferred in odour and taste (Hansen & Hansen, 1996).
et al., 1989a). Sensory evaluation of sourdough of wheat bread crumb
showed that bread made from sourdough fermented with
Effect of Maillard and Caramelization reactions the heterofermentative Lb. sanfranciscensis had a pleasant,
on the flavour of bread mild, sour odour and taste, whereas sourdough bread
The production of volatiles is also influenced by proof- fermented with the homofermentative Lb. plantarum had
ing and baking processes (Hansen, Lund, & Lewis, an unpleasant metallic sour taste. But, when sourdough
1989b; Kamiński, Przybylski, & Gruchala, 1981). The was supplemented with sourdough yeast S. cerevisiae, the
Maillard and Caramelization reactions are overwhelmingly wheat bread received a more aromatic flavour (Katina,
responsible for flavour formation in cereal products. Upon Heiniö, Autio, & Poutanen, in press). Chemical analyses
heating, the reaction involves between simple the precur- of flavour compounds can be combined with sensory anal-
sors such as amino acids and simple aldose or ketose of ysis of bread. Compounds that have been positively
sugars (Rothe & Ruttloff, 1983). As shown previously, correlated with flavour of wheat crumb are acetaldehyde,
the highest amount of free amino acids was produced 2-methylpropanoic acid, 2/3-methyl-1-butanol, 3-methyl-
during sour dough fermentation with the association butanoic acid, isopentanal, 2-nonenal, benzylethanol, 2-
between LAB and S. exiguus M14 (Gobbetti, Corsetti phenylethanol, 2,3-butandione and 3-hydoxy-2-butanone,
et al., 1994; Gobbetti, Simonetti et al., 1994; Kratochivil dimethyl sulphide and 2-furfural (Hansen & Hansen,
& Holas, 1983; Rothe, 1974). These amino acids are pre- 1996; Rothe, 1974). Sensory evaluation of sourdough rye
cursors of iso-alcohols (Gobbetti, Corsetti, & De Vincenzi, bread crumb suggested that a most intense and bread-like
1995b; Hansen & Schieberle, 2005) that contribute directly flavour was related to propanone, 3-methylbutanal, benzyl
to bread flavour during sourdough fermentation and baking alcohol and 2-phenylethanol (Hansen et al., 1989a).
 S.-ur-Rehman et al. / Trends in Food Science & Technology 17 (2006) 557e566 563

Effect of type and flour composition heterofermentative cultures, with the highest amounts de-
on flavour compounds tected in sourdough made from wholemeal and low-grade
The interactions between starter cultures and flour com- flours (Hansen & Hansen, 1994a). In contrast to white
ponents play an important role in the production of flavour wheat bread, the starch granules are very much swollen
compounds in the sourdough breads. A particular favoured in the bran sourdough bread with enzymatic mixture which
cereal for sourdough is rye flour that contains flavour pre- improves the texture, volume, flavour and shelf-life of
cursor amino acids, fatty acids and phenolic compounds bread (Katina, Salmenkallio-Marttila, Partanen, Forsell, &
that are converted into flavour-active components in differ- Autio, 2006).
ent production stages of baking (Schieberle & Grosch, With the high extraction rate (80e100%), the content of
1983; Schieberle & Grosch, 1985; Schieberle & Grosch, nutrients such as B vitamins and minerals increases com-
1987; Hansen, 1995). In rye, germination imparts cereal pared to low extraction rate flour (65e75%), as does the
and fresh flavour in extruded products. Key flavour compo- buffering capacity of flour chiefly due to the phytic acid
nents are dimethyl sulphide and 2-methylbutanal which (Batifoulier, Verny, Chanliaud, Rémésy, & Demigné,
could be related to the sensory attribute intensities. Ex- 2005). These factors stimulate the growth and biochemical
truded rye sourdough has been reported to have an intense, activity of the LAB followed by a higher production of
sour flavour and porous texture with high contents of furfu- acids and flavour compounds (Martinez-Anaya, Benedito
ral, ethyl acetate, 3-methylbutanol and 2-methylbutanol and de Barber, & Esteve, 1994; Salovaara & Valjakka, 1987).
such extrudates of rye flour are rich in 2-ethylfuran, 2- Soya flour, containing a range of enzyme activities, also
methylfuran, hexanal and pentanal (Heiniö et al., 2003). could influence the bread volatiles, isolated by dynamic
Other contributors to the overall crumb flavour of rye bread headspace techniques and quantified by gas chromatogra-
include 3-methylbutanal, 2-nonenal, 2,4-dcadienal, hexa- phy and gas chromatography/mass spectrometry (Luning
nal, phenylacetaldehyde, methional, vanillin, 2,3-butan- et al., 1991).
dione, 3-hydroxy-4,5-dimethyl-2(5H)-furanone and 2- and Wheat and rye flours are mostly used for sourdough
3-methylbutanoic acid (Kirchhoff & Schieberle, 2001). making but maize and rice flours can also be used (Buttery
Sourdough breads may vary in flavouring compounds as & Ling, 1999; Buttery, Orts, Takeoka, & Nam, 1999;
a result of flour composition and blending of the cereal Clarke & Arendt, 2005; Rocha & Malcata, 1999; Sanni,
flours (Schieberle & Grosch, 1985; Schieberle & Grosch, Onilude, & Fatungase, 1998). Major volatiles of rice flour
1992; Schieberle & Grosch, 1994). Wheat flour composition include 1-hydroxy-2-propanone, furfuryl alcohol, 2,5-dime-
influences the concentration of ethyl acetate and ethanol thylpyrazine, 2-methylpyrazine, pyrazine, hexanal, furfural,
production in heterofermentative LAB sourdoughs which pentanol, 3-hydroxy-2-butanone and ethyl-3,6-dimethyl-
are more abundant with high-grade flours (Hansen & pyrazine (Buttery et al., 1999) whereas major volatiles
Hansen, 1994a). Ash contents from 0.55 to1.00% had identified in corn flour include 2-methylpyrazine, 1-
positive influences on total volatiles. Wheat bread from hydroxy-2-propanone, 4-vinylguaicol, 2-acetyltetrahydro-
sourdough sponges including Lb. plantarum or Lb. sanfran- pyridine (E,E )- and (E,Z )-2,4-decadienal, acetic acid,
ciscensis had a higher content of 2,3-methyl-1-butanol; and 3-methylbutanal, g-butyrolactone, furfuryl alcohol and
through association of LAB and yeasts, bread was consid- 2,5-dimethylpyrazine (Buttery & Ling, 1999).
ered of enhanced flavour quality with higher contents of
2,3-methyl-1-butanol, 2-methyl-propanoic acid, 3-methyl-
butanoic acid and 2-phenyl-ethanol (Hansen et al., 1989b; Conclusion
Hansen & Hansen, 1996). It is considered that this is the In sourdoughs, flavour-active compounds are produced
result of the combination of greater bacterial acidification by LAB and yeasts individually and through their interac-
and proteolysis (Gobbetti, Corsetti et al., 1994; Gobbetti, tions. Heterofermentative LAB mainly produce ethyl
Simonetti et al., 1994; Levesque, 1991) which may be acetate and certain alcohols and aldehydes, whereas homo-
attributed to Lb. sanfranciscensis in the sourdough. Hansen fermentative LAB synthesise diacetyl and other carbonyls.
& Hansen (1994b) have suggested an association of In contrast, iso-alcohols are products of yeast fermentation
Lb. sanfranciscensis, Lb. plantarum and S. cerevisiae as but may contribute little towards final bread flavour. Addi-
optimal for aroma balance in wheat sourdough breads. tion of fructose/glucose/maltose or citrate to the dough
The amount of fermentable carbohydrate in the flour increases LAB contributions to volatile formation in baking
varies with the type of cereal, but in particular with the but Maillard and Caramelization reactions are also respon-
activities of amylases, xylanases and peptidases enzymes sible for flavour formation in baked cereal products. Inter-
of flour. In wheat flours, totals of maltose, sucrose, glucose actions between microbial strains and ingredients,
and fructose vary from 1.55 to 1.85% depending on extent fermentation temperature, pH and leavening time have all
of starch hydrolysis, and activities of microbial enzymes been evaluated. Further research should be conducted on
and microbial contaminants (Martinez-Anaya, 1996). optimization of fermentation process in wheat bread
The flour type has a momentous effect on the production production by varying biotechnical parameters, levels of
of ethyl acetate and ethanol in sourdough emulsifiers/improvers including lactose, milk solids and
564 S.-ur-Rehman et al. / Trends in Food Science & Technology 17 (2006) 557e566

oxidants. Effects of dried mixed cultures of yeasts-LAB wheat doughs started with pure culture of lactic acid bacteria.
also need further attention. Cereal Chemistry, 68, 66e72.
Corsetti, A., Gobbetti, M., & Rossi, J. (1994). Sourdough fermentation
in presence of added soluble carbohydrates. Microbiology
References Aliments Nutrition, 12, 377e385.
Corsetti, A., Gobbetti, M., De Marco, B., Balestrieri, F., Paletti, F.,
Aksu, Z., & Kutsal, T. (1986). Lactic acid production from molasses Russi, L., et al. (2000). Combined effect of sourdough lactic acid
utilizing Lactobacillus delbrueckii and invertase together. bacteria and additives on bread firmness and staling. Journal of
Biotechnology Letters, 8, 157e160. Agricultural and Food Chemistry, 48, 3044e3051.
Batifoulier, F., Verny, M. A., Chanliaud, E., Rémésy, C., & Demigné Corsetti, A., Lavermicocca, P., Morea, M., Baruzzi, F., Tosti, N., &
(2005). Effect of different breadmaking methods on thiamine, Gobbetti, M. (2001). Phenotypic and molecular identification
riboflavin and pyridoxine contents of wheat bread. Journal of and clustering of lactic acid bacteria and yeasts from wheat
Cereal Science, 42, 101e108. (species Triticum durum and Triticum aestivum) sourdoughs of
Barber, S., Baguena, R., Benedito de Barber, C., & Martinez- Southern Italy. International Journal of Food Microbiology, 64,
Anaya, M. A. (1991). Evolution of biochemical and rheological 95e104.
characteristics and breadmaking quality during a multistage wheat Damiani, P., Gobbetti, M., Cossignani, L., Corsetti, A.,
sour dough process. Zeitschrift fur Lebensmittel-Untersuchung Simonetti, M. S., & Rossi, J. (1996). The sourdough microflora.
und-Forschung, 192, 46e52. Characterization of hetero- and homofermentative lactic acid
Barber, S., Benedito de Barber, C., Martinez-Anaya, M. A., Martinez, J., bacteria, yeasts and their interactions on the basis of the volatile
& Alberola, J. (1985). Cambios en los acidos organicos volatiles compounds produced. LWT-Food Science and Technology, 29,
C2eC5 durante la fermentacation de mases panarias preparados 63e70.
con masas madres comerciales y con cultivos puros de micro- Frankel, E. N. (1982). Volatile lipid oxidation products. Progress in
organismos. Revista de Agroquimica y Technologia de Alimentos, Lipids Research, 22, 4e33.
25, 223e232. Frasse, P., Lambert, S., Richard-Molard, D., & Chiron, H. (1993).
Brandt, M. J., Hammes, W. P., & Ganzle, M. G. (2004). Effects of The influence of fermentation on volatile compounds in french
process parameters on growth and metabolism of Lactobacillus bread dough. LWT-Food Science and Technology, 26,
sanfranciscensis and Candia humillis during rye. European Food 126e132.
Research and Technology, 218, 333e338. Galal, A. M., Johnson, J. A., & Varriano-Marston, E. (1978). Lactic acid
Bredie, W. L. P., Mottram, D. S., & Guy, R. C. E. (2002). Effect of and volatile (C2eC5) organic acids of San Francisco sourdough
temperature and pH on the generation of flavor volatiles in French bread. Cereal Chemistry, 55, 461e468.
extrusion cooking of wheat flour. Journal of Agricultural and Gianotti, A., Vannini, L., Gobbetti, M., Corsetti, A., Gardini, F., &
Food Chemistry, 50, 1118e1125. Guerzoni, M. E. (1997). Modelling of the activity of selected
Bruemmer, J. M., & Lorenz, K. (1991). Europeon development in starters during sourdough fermentation. Food Microbiology, 14,
wheat sourdoughs. Cereal Food World, 36, 310e312. 327e337.
Buttery, R. G., & Ling, L. C. (1999). Additional studies on flavour Gobbetti, M. (1998). The sourdough microflora: interactions of lactic
components of corn tortilla chips. Journal of Agricultural and acid bacteria and yeasts. Trends in Food Science & Technology, 9,
Food Chemistry, 46, 2764e2769. 267e274.
Buttery, R. G., Orts, W. J., Takeoka, G. R., & Nam, Y. (1999). Volatile Gobbetti, M., & Corsetti, A. (1996). Co-metabolism of citrate and
flavour components of rice cakes. Journal of Agricultural and maltose by Lactobacillus brevis subsp. lindneri cb1 citrate-negative
Food Chemistry, 47, 4353e4356. strain: effect on growth, end-products and sourdough fermentation.
Cagno, R. D., Angelis, M. D., Corsetti, A., Lavermicocca, P., Zeitschrift fur Lebensmittel-Untersuchung und-Forschung, 203,
Arnault, P., Tossut, P., et al. (2003). Interactions between sourdough 82e87.
lactic acid bacteria and exogenous enzymes: effects on the Gobbetti, M., & Corsetti, A. (1997). Lactobacillus sanfranciscoa key
microbial kinetics of acidification and dough textural properties. sourdough lactic acid bacterium: a review. Food Microbiology, 14,
Food Microbiology, 20, 67e75. 175e188.
Calderon, M., Loiseau, G., & Guyot, J. P. (2003). Fermentation by Gobbetti, M., Corsetti, A., & De Vincenzi, S. (1995a). The sourdough
Lactobacillus fermentum Ogi E1 of different combinations of microflora. Characterization of homofermentative lactic acid
carbohydrates occurring naturally in cereals: consequences on bacteria based on acidification kinetics and impedance tests.
growth energetic and -amylase production. International Journal Italian Journal of Food Science, 2, 91e102.
of Food Microbiology, 80, 161e169. Gobbetti, M., Corsetti, A., & De Vincenzi, S. (1995b). The sourdough
Catzeddu, P., Mura, E., Parente, E., Sanna, M., & Farris, G. A. (2006). microflora. Characterization of heterofermentative lactic acid
Molecular characterization of lactic acid bacteria from sourdough bacteria based on acidification kinetics and impedance tests.
breads produced in Sardinia (Italy) and multivariate statistical Italian Journal of Food Science, 2, 103e112.
analyses of results. Systematic and Applied Microbiology, 29, Gobbetti, M., Corsetti, A., & Rossi, J. (1994a). The sourdough
138e144. microflora. Interactions between lactic acid bacteria and yeasts:
Clarke, C. I., & Arendt, E. K. (2005). A review of the application of metabolism of carbohydrates. Applied Microbiology and
sourdough technology to wheat breads. Advances in Food and Biotechnology, 41, 456e460.
Nutrition Research. Gobbetti, M., Corsetti, A., & Rossi, J. (1994b). The sourdough
Collar, C. (1996). Biochemical and technological assessment of the microflora. Interactions between lactic acid bacteria and yeasts:
metabolism of pure and mixed cultures of yeast and lactic acid metabolism of amino acids. World Journal of Microbiology and
bacteria in breadmaking applications. Food Science and Biotechnology, 10, 275e279.
Technology International, 2, 349e367. Gobbetti, M., Corsetti, A., & Rossi, J. (1995). Maltoseefructose
Collar, C., & Martinez, C. S. (1993). Amino acid profiles of fermen- co-fermentation by Lactobacillus brevis subsp. lindneri CB1
ting wheat sour doughs. Journal of Food Science, 58, 1324e fructose-negative strain. Applied Microbiology and Biotechnology,
1328. 42, 939e944.
Collar, C., Mascaros, A. F., Prieto, J. A., & Benedito de Barber, C. Gobbetti, M., Corsetti, A., Rossi, J., La Rosa, F., & De Vincenzi, S.
(1991). Changes in free amino acids during fermentation of (1994). Identification and clustering of lactic acid bacteria and
 S.-ur-Rehman et al. / Trends in Food Science & Technology 17 (2006) 557e566 565

yeasts from wheat sourdoughs of central Italy. Italian Journal of Katina, K., Salmenkallio-Marttila, M., Partanen, R., Forsell, P., &
Food Science, 1, 85e94. Autio, K. (2006). Effects of sourdough and enzymes on staling of
Gobbetti, M., Simonetti, M. S., Rossi, J., Cossignani, L., Corsetti, A., high-fibre wheat bread. LWT-Food Science and Technology, 39,
Damiani, P. (1994). Free D- and L-amino acid evolution during 479e491.
sourdough fermentation and baking. Journal of Food Science, 59, Kirchhoff, E., & Schieberle, P. (2001). Determination of key aroma
881e884. compounds in the crumb of a three-stage dilution assays and
Gobbetti, M., Smacchi, E., Fox, P. F., Stepaniak, L., & Corsetti, A. sensory studies. Journal of Agricultural and Food Chemistry, 49,
(1996). The sourdough microflora. Cellular localization and 4304e4311.
characterization of proteolytic enzymes in lactic acid bacteria. Kratochivil, J., & Holas, J. (1983) Study of proteolytic processes Rye
LWT-Food Science and Technology, 29, 561e569. sour. Developments in food science, Vol. 5B. Oxford: Elsevier
Gül, H., Özçelik, S., Sadıç, O., & Certel, M. (2005). Sourdough bread (pp. 791e798).
production with lactobacilli and S. cerevisiae isolated from Levesque, C. (1991) ‘Etude des Potentialitès de la Souche S47 de
sourdoughs. Process Biochemistry, 40, 691e697. Saccharomyces cerevisiae à produire des alcools supérieurs à
Hammes, W. P., & Ganzle, M. (1995). The genus Lactobacillus. In partir de composés organiques identifiés et des précurseurs de la
B. J. B. Wood, & W. H. Holzapfel (Eds.), The genera of Lactic farine’ in thèse d’université, Nantes.
acid bacteria, Vol. II (pp. 19e54). London: Blackie Academic & Linko, Y., Javanainen, P., & Linko, S. (1997). Biotechnology of
Professional. bread baking. Trends in Food Science & Technology, 8,
Hann, D., & Morrison, W. R. (1975). Effects of ingredients on the 339e344.
oxidation of linoleic acid by lipoxygenase in bread doughs. Journal Lund, B., Hansen, Å., & Lewis, M. J. (1989). The influence of dough
of Science of Food and Agriculture, 26, 493. yield on acidification and production of volatiles in sourdoughs.
Hansen, A. (1995). Flavour compounds in rye sourdough and rye LWT-Food Science and Technology, 22, 150e153.
bread. In K. Pountanen, & K. Autio (Eds.), International rye Luning, P. A., Roozen, J. P., Moëst, R. A. F. J., & Posthumus, M. A.
symposium: technology and products (p. 160). Helsinki, Finland: (1991). Volatile composition of white bread using enzyme active
VTT, VTT Symposium 161. soya flour as improver. Food Chemistry, 41, 81e91.
Hansen, A., & Hansen, B. (1993). Flavour compounds in wheat Martinez-Anaya, M. A. (1996). Enzymes and bread flavour. Journal
sourdoughs. In C. Benedito De Barber, C. Collar, M. A. Martinez- of Agriculture and Food Chemistry, 44, 2469e2480.
Anaya, & J. Morel (Eds.), Progress in food fermentation, Proceed- Martinez-Anaya, M. A., Benedito de Barber, C., & Esteve, C. C. (1994).
ings of Euro Food Chem. VII. Valencia: IATA CSIC. Effect of processing conditions on acidification properties of wheat
Hansen, A., & Hansen, B. (1994a). Influence of wheat flour type on the sourdough. International Journal of Food Microbiology, 22,
production of flavour compounds in wheat sourdoughs. Journal 249e255.
of Cereal Science, 19, 185e190. Martinez-Anaya, M. A., Pitarch, B., & Benedito de Barber, C. (1993).
Hansen, A., & Hansen, B. (1994b). Volatile compounds in wheat Biochemical characteristics and breadmaking performance of
sourdoughs produced by lactic acid bacteria and sourdough yeasts. freeze- dried wheat sour dough starters. Zeitschrift fur Lebensmittel-
Zeitschrift fur Lebensmittel-Untersuchung und-Forschung, 198, Untersuchung und-Forschung, 196, 360e365.
202e209. Meignen, B., Onno, B., Gélinas, P., Infantes, M., Guilois, S., &
Hansen, A., & Hansen, B. (1996). Flavour of sourdough wheat bread Cahagnier, B. (2001). Optimization of sourdough fermentation
crumb. Zeitschrift fur Lebensmittel-Untersuchung und-Forschung, with Lactobacillus brevis and baker’s yeast. Food Microbiology,
202, 244e249. 18, 239e245.
Hansen, A., Lund, B., & Lewis, M. J. (1989a). Flavour of sourdough Merotha, C. B., Hammesa, W. P., & Hertela, C. (2004). Characterisa-
rye bread crumb. LWT-Food Science and Technology, 22, 141e tion of the microbiota of rice sourdoughs and description of
144. Lactobacillus spicheri sp. nov. Systematic and Applied
Hansen, A., Lund, B., & Lewis, M. J. (1989b). Flavour production Microbiology, 27, 151e159.
and acidification of sourdoughs in relation to starter culture and Merseburger, T., Ehret, A., Geiges, O., Baumann, B., & Schmidt-
fermentation temperature. LWT-Food Science and Technology, Lorenz, W. (1995). Microbiology of dough preparation vii.
22, 145e149. production and use of preferments to produce wheat-bread.
Hansen, A., & Schieberle, P. (2005). Generation of aroma compounds Mitteilungen aus dem Gebiete der Lebensmitteluntersuchung
during sourdough fermentation: applied and fundamental aspects. und Hygiene, 86, 304e324.
Trends in Food Science & Technology, 16, 85e94. Neubauer, H., Glaasker, E., Hammes, W. P., Poolman, B., &
Heiniö, R., Katina, K., Wilhelmson, A., Myllymäki, O., Rajamäki, T., Konings, W. N. (1994). Mechanism of maltose uptake and glucose
Latva-Kala, K., et al. (2003). Relationship between sensory excretion in Lactobacillus sanfrancisco. Journal of Bacteriology,
perception and flavour-active volatile compounds of germinated, 176, 3007e3012.
sourdough fermented and native rye following the extrusion Neysens, P., Messens, W., & Vuyst, L. D. (2003). Effect of sodium
process. LWT-Food Science and Technology, 36, 533e545. chloride on growth and bacteriocin production by Lactobacillus
Ji, H., & Berhard, R. A. (1992). Effect of microwave heating on amylovorus DCE 471. International Journal of Food Microbiology,
pyrazine formation in a model system. Journal of the Science of 88, 29e39.
Food and Agriculture, 59, 283e289. Nout, M. J. R., & Creemers-Molenaar, T. (1987). Microbiological
Kamiński, E., Przybylski, R., & Gruchala, L. (1981). Thermal degra- properties of some wheatmeal sourdough starters. Chemistry
dation of precursors and formation of flavour compounds during Mikrobiology and Technology Lebensmittel, 10, 162e167.
heating of cereal products. Part I. Changes of amino acids and Onno, B., & Rouseel, P. (1994). Technologie et microbiologie de le
sugars. Die Nahrung, 25, 507e518. panification au levain. In H. De Roissart, & F. M. Luquet (Eds.),
Katina, K., Arendt, E., Liukkon, K.-H., Autio, K., Flander, L., & Bacteries lactiques, Vol. 2 (pp. 293e321). France: Lorica Uriage.
Poutanen, K. (2005). Potential of sourdough for healthier cereal Ottogalli, G., Galli, A., & Foschino, R. (1996). Italian bakery products
products. Trends in Food Science and Technology, 16, 104e112. obtained with sour dough: characterization of the typical micro-
Katina, K., Heiniö, R. L., Autio, K., & Poutanen, K. (2006). flora. Advances in Food Science, 18, 131e144.
Optimization of sourdough process for improved sensory Oura, E., Soumalainen, H., & Wiskari, R. (1982). Sourdough. In
profiles and texture of wheat bread. LWT-Food Science and A. H. Rose (Ed.), Fermented foods (pp. 123e146). London:
Technology, 39, 1189e1202. Academic Press.
566 S.-ur-Rehman et al. / Trends in Food Science & Technology 17 (2006) 557e566

Plessas, S., Pherson, L., Bekatou, A., Nigam, P., & Koutinas, A. A. Seppi, A. (1984). Studio sulla biochimica degli zuccheri nella
(2005). Bread making using kefir grains as baker yeast. Food fermentazione del pane. Revista Soc. Ital. Science Alimentos, 25,
Chemistry, 93, 585e589. 223e232.
Parker, J. K., Hassell, G. M., Mottram, D. S., & Guy, R. C. (2000). Simonson, L., Salovaar, H., & Korhola, M. (2003). Response of wheat
Sensory and instrumental analyses of volatile generated during sourdough parameters to temperature, NaCl and sucrose varia-
the extrusion cooking of oat flour. Journal of Agricultural and tions. Food Microbiology, 20, 193e199.
Food Chemistry, 48, 3497e3506. Spicher, G. (1983). Baked goods. In J. H. Rehm, & G. Reed (Eds.),
Przybylski, R., & Kamiński, E. (1983). Thermal degradation of precur- Biotechnology (pp. 1e80). Weinheim: Verlag Chemie.
sors and formation of flavour compounds during heating of cereal Spicher, G., & Nierle, W. (1984). The microflora of sourdough. xviii.
products. Part II. The formation and changes of volatile flavour communication: the protein degrading capabilities of the lactic
compounds in thermally treated malt extracts at different acid bacteria of sourdough. Zeitschrift fur Lebensmittel-
temperature and pH. Die Nahrung, 27, 487e496. Untersuchung und-Forschung, 178, 389e392.
Ricciardi, A., Parente, E., Piraino, P., Paraggio, M., & Romano, P. Spicher, G., Rabe, E., Sommer, R., & Stephan, H. (1981). The micro-
(2005). Phenotypic characterization of lactic acid bacteria from flora of sourdough. xiv. communication: about the behaviour of
sourdoughs for Altamura bread produced in Apulia (South Italy). homofermentative sourdough bacteria and yeasts in mixed culture.
International Journal of Food Microbiology, 98, 63e72. Zeitschrift fur Lebensmittel-Untersuchung und-Forschung, 173,
Robert, H., Gabriel, V., Lefebvre, D., Rabier, P., Vayssier, Y., & 291e296.
Fontagné-Faucher, C. (2006). Study of the behaviour of Lacto- Spicher, G., Rabe, E., Sommer, R., & Stephan, H. (1982). Communi-
bacillus plantarum and Leuconostoc starters during a complete cation: on the behaviour of heterofermentative sourdough bacteria
wheat sourdough breadmaking process. LWT-Food Science and and yeasts in mixed culture. Zeitschrift fur Lebensmittel-
Technology, 39, 256e265. Untersuchung und-Forschung, 174, 222e227.
Rocha, J. M., & Malcata, F. X. (1999). On the microbiological profile of Stolz, P., Böcker, G., Hammes, W. P., & Vogel, R. F. (1995). Utilization
traditional Portuguese sourdough. Journal of Food Protection, 62, of electron acceptors by lactobacilli isolated from sourdough.
1416e1429. Zeitschrift fur Lebensmittel-Untersuchung und-Forschung, 201,
Röcken, W., & Voysey, P. A. (1993). Sourdough fermentation in 91e96.
bread making. Journal of Applied Bacteriology (Supplement), 79, Stolz, P., Böcker, G., Vogel, R. F., & Hammes, W. P. (1993). Utilization
38Se39S. of maltose and glucose by lactobacilli isolated from sourdough.
Röcken, W., & Voysey, P. A. (1995). Sourdough fermentation in FEMS Microbiology Letters, 109, 237e242.
breadmaking. Journal of Applied Bacteriology (Symposium Succi, M., Reale, A., Andrighetto, C., Lombardi, A., Sorrentino, E., &
Supplement), 79, 38Se48S. Coppol, R. (2003). Presence of yeasts in southern Italian sour-
Rothe, M. (1974). Aroma in bread (in German). Berlin: Akademie. doughs from Triticum aestivum flour. FEMS Microbiology
Rothe, M., & Ruttloff, H. (1983). Aroma retention in modern bread Letters, 225, 143e148.
production. Die Nahrung, 27, 505e512. Suihko, M. L., & Makinen, V. (1984). Tolerance of acetate, propionate
Rouzaud, O., & Martinez-Anaya, M. A. (1993). Effect of processing con- and sorbate by Saccharomyces cerevisiae and Torulopsis holmii.
ditions on oligosaccharide profile of wheat sourdoughs. Zeitschrift fur Food Microbiology, 1, 105e110.
Lebensmittel-Untersuchung und-Forschung, 197, 434e439. Sugihara, T. F., Kline, L., & McCready, L. B. (1970). Nature of San
Salovaara, H., & Valjakka, T. (1987). The effect of fermentation tem- Francisco sour dough French bread process. II. Microbiological
perature, flour type and starter on the properties of sour wheat aspects. Bakers Digest, 44, 51e56.
bread. International Journal of Food Science and Technology, 22, Torner, M. J., Martı́nez-Anaya, M. A., Antuña, B., & Benedito de
591e597. Barber, C. (1992). Headspace flavour compounds produced by
Simova, E., Beshkova, D., Angelov, A., Hristova, T., Frengova, G., & yeasts and lactobacilli during fermentation of preferments and
Spasov, Z. (2002). Lactic acid bacteria and yeasts in kefir grains and bread doughs. International Journal of Food Microbiology, 15,
Kefir made from them. Journal of Industrial and Microbiological 145e152.
Biotechnology, 28, 1e6. Trüper, H. G., & de Clari, L. (1997). Taxonomic note: necessary
Sanni, A. I., Onilude, A. A., & Fatungase, M. O. (1998). Production correction of specific epithets formed as substantives (nouns) ‘in
of sour maize bread using starter culture. World Journal of apposition’. International Journal of Systematic Bacteriology, 47,
Microbiology and Biotechnology, 14, 101e106. 908e909.
Schieberle, P., & Grosch, W. (1994). Potent odorants of rye bread Vernocchi, P., Valmorri, S., Gatto, V., Torriani, S., Gianotti, A.,
crust-differences from the crumb and from wheat bread crust. Suzzi, G., et al. (2004). A survey on yeast microbiota associated
Zeitschrift fur Lebensmittel-Untersuchung und-Forschung, 198, with an Italian traditional sweet-leavened baked good fermenta-
292e296. tion. Food Research International, 37, 469e476.
Schieberle, P., & Grosch, W. (1992). Changes in the concentration Vogel, R. F. (1997). Microbial ecology of cereal fermentations. Food
of potent crust odorants during storage of white bread. Flavour Technology and Biotechnology, 35, 51e54.
and Fragrance Journal, 7, 213e218. Vogel, R. F., Bocker, G., Stolz, P., Ehrmann, M. D., Ludwig, W., Pot, B.,
Schieberle, P., & Grosch, W. (1987). Quantitative analysis of aroma et al. (1994). Identification of lactobacilli from sourdough and
compounds in wheat and rye bread crusts using a stable isotope description of Lactobacillus pontis sp. nov. International Journal of
dilution assay. Journal of Agricultural and Food Chemistry, 35, Systematic Bacteriology, 44, 223e229.
252e257. Vogel, R. F., Knorr, R., Müller, M. R. A., Steudel, U., Gänzle, M. G., &
Schieberle, P., & Grosch, W. (1985). Identification of volatile flavour Ehrmann, M. A. (1999). Non-dairy lactic acid fermentations: the
compounds of wheat bread crust-comparison with rye bread crust. cereal world. Antonie van Leeuwenhoek, 76, 403e411.
Zeitschrift fur Lebensmittel-Untersuchung und-Forschung, 180, Vollmar, A., & Meuser, F. (1992). Influence of starter cultures
474e478. consisting of lactic acid bacteria and yeast on the performance
Schieberle, P., & Grosch, W. (1983). Identification of flavour com- of a continuous sourdough fermenter. Cereal Chemistry, 69,
pounds from the crust of rye bread. Zeitschrift für Lebensmittel- 20e27.
Untersuchung und-Forschung, 177, 173e180. Vuyst, L. D., & Neysens, P. (2005). The sourdough microflora:
Seitz, L. M., Chung, O. K., & Rengarajan, R. (1998). Volatiles in biodiversity and metabolic interactions. Trends in Food Science
selected commercial breads. Cereal Chemistry, 75, 847e853. & Technology, 16, 43e56.