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Impact of Saccharomyces cerevisiae metabolites produced during fermentation on bread

quality parameters: a review

Mareile Heitmanna, Emanuele Zanninia, Elke Arendta,*

Abstract

Although bread making with the use of Baker’s yeast has a long tradition in human history, little

attention has been paid to the connection between yeast addition and the final bread quality.

Nowadays, bakers mainly use different flour additives such as enzymes (amylases,

hemicellulases, and proteases) to change and improve dough properties and/or bread quality.

Another strategy is the use of modified industrial Baker’s yeast. To date, there is no yeast strain

used in the baking industry, which is genetically modified, despite some studies demonstrating

that the application of recombinant DNA technology is a possibility for improved strains suitable

for baking. However, due to the fact that the majority of consumers in Europe highly reject the

use of genetically modified microorganisms in the production of food, other strategies to

improve bread quality must be investigated. Such a strategy would be a reconsideration of the

selection of yeast strains used for the baking process. Next to the common criteria, the

requirement for adequate gas production, more attention should be paid on how yeast impacts

flavour, shelf life, colour and the nutritional value of baked products, in a similar way to which

yeast strains are selected in the wine and brewing industries.

Keywords

Yeast, Metabolism, Baked products, Yeast selection

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Abbreviations

(GDL) Glucono-delta-lactone

(ATP) Adenosine triphosphate

(I) Invertase

(G) Glucosidase

(GPD) Glycerol-3-phosphate dehydrogenase

(GP) Glycerol-3-phosphatase

(AD) Alcohol dehydrogenase

(AcD) Acetaldehyde dehydrogenase

(PD) Pyruvate dehydrogenase

(PDc) Pyruvate decarboxylase

(Ac-CoA-S) Acetyl-CoA synthase

(TCA) Tricaboxylic acid

(CS) Citrate synthase

(A) Aconitase

(ID) Isocitrat dehydrogenase

(α-KD) Alpha-Ketoglutarate dehydrogenase

(S-CoA-S) Succinyl-CoA synthase

(SD) Succinate dehydrogenase

(F) Fumarase

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(MD) Malat dehydrogenase

(IL) Isocitrate lyase

(MS) Malate synthase

(GS) Glutamate synthase

(QPS) Qualified Presumption of Safety

(GRAS) Generally Regarded as Safe

(GABA) γ-aminobutyric acid

(GAD) Glutamic acid decarboxylase

(S.) Saccharomyces

(P.) Penicillium

(M.) Meyerozyma

(vit.) Vitamin

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1. Introduction

Bread making is one of the oldest biochemistry processes in the whole world. Saccharomyces

cerevisiae also known as Baker’s yeast is one of the main ingredients for bread making. The

term “cerevisiae” (meaning beer) signifies closely linked relationship between beer and bread

making, originating centuries ago in Egypt and the Middle East. Historically, the same strain of

yeast was used for both processes. In the nineteenth century, yeasts’ left over from the brewing

industry were shared with the bakers for bread production. Today, thousands of different

genetically improved microbial cultures are used for different applications, like baking, brewing

and wine making. Although bread making has a long tradition throughout human history, little

attention has been focused on the connection between yeast addition and the final bread product

quality (Dequin, 2001; Mondal and Datta, 2008). To date, bakers mainly use different flour

additives, such as enzymes (amylases, hemicellulases, and proteases) to change and improve

dough properties and/or bread quality. Another strategy is the use of modified industrial Baker’s

yeast. During fermentation yeast produces mainly carbon dioxide, but the role of yeast goes

much deeper than just gas production (Randez-Gil et al., 1999), concerning the production of

ethanol and other secondary metabolites, which have an impact on the final product quality.

Yeast affects the volume, structure, flavour, colour and shelf life of each fermented product

(Fleet, 2007). The characteristic volume and aerated cell structure of bread are mainly influenced

by the addition of yeast its metabolism and carbon dioxide production during fermentation. Due

to the production of secondary metabolites through different metabolic pathways, yeast

influences the flavour (by producing precursors such as esters, aldehydes and ketones), colour

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(carbohydrates, amino acids) and shelf-life (acids, glycerol) of baked products. All these

metabolic products demonstrate the important role of yeast in bread making. Nevertheless, the

most important characteristic, which is usually considered during strain selection is the ability to

ferment sugars anaerobically with adequate gas production (Reed and Nagodawithana, 1991). In

our opinion, the production of other metabolites by the yeast is underestimated when considering

the selection of yeast strains. The purpose of this study is therefore to describe the impact of

yeast in view of final bread quality parameters. This article reviews critically published literature

on studies related to yeast and bread quality and identifies potential future investigations for

applied yeast research with particular reference to the production of wheat bread. The main

intention of this study is to better understand the complex dough fermentation reactions

performed by yeast and their impact on product quality. This may enable the adaption of new

yeast strains including those currently used in other applications, such as the brewing and wine

industries, which can specifically enhance bread properties and so the final bread quality. The

present review further examines the metabolites produced by yeast during dough fermentation

and their impact on bread quality parameters. This knowledge could help to create new

procedures and criteria for yeast strain selection for application in bread making.

2. Yeast in bread making

Yeast is an ubiquitous, unicellular, asexual eukaryote belonging to the kingdom Fungi, which is

able to ferment sugars (added or produced by enzymatic hydrolysis) into alcohol and carbon

dioxide (Cauvain and Young, 2007) and therefore is known as the leavening agent in baked

goods. Their shape is typically spherical, oval or cylindrical with an average diameter of around

8 µm. The cells contain a double layered cell wall through which the cell is able to absorb

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nutrients and release metabolites. The main yeast strains related to bread making are from the

species S. cerevisiae (Cauvain and Young, 2007; Fleet, 2007). Fresh Baker’s yeast comprises of

30-33 % of dry materials, 40.6-58.0 % of proteins, 35.0-45.0 % of carbohydrates, 5.0-7.5 % of

minerals, 4.0-6.0 % of lipids and several vitamins (vit.) (Bekatorou et al., 2006). The European

yeast industry produces 1 million tonnes of yeast annually of which around 30 % is exported

globally (http://www.cofalec.com/business-and-economy/). The annual growth rate of the global

market is expected to be 8.8 % from 2013 to 2018 (http://www.marketsandmarkets.com/Market-

Reports/yeast-industry-268.html). Typically, an aerobic fed-batch process with molasses as a

nutrient source is used for the commercial production of yeast (Attfield, 1997). The process

consist of growing, separating, washing and processing to remove extracellular and intracellular

water by filtration or pressing (Randez-Gil et al., 1999). To decrease damage to the yeast cells

additives like emulsifiers and/or antioxidants are added during production. Growth conditions

including temperature (25-30 °C), moisture, and nutrients (starch, sugar) must be optimised.

When yeast is grown outside these optimal parameters, a complex stress response occurs

(Attfield, 1997). Stresses can cause direct and/or indirect cell damage influencing the membrane

permeability, inhibiting enzymes activity and result in the formation of reactive oxygen species.

Cell responses include a decrease in intracellular pH through the production of glycerol,

formation of several antioxidant defences such as glutathione and increased membrane

permeability for intracellular components. Stress response is an important factor for survival and

growth in industrial applications (Attfield, 1997). Intensive biochemical, microbiological and

technical knowledge has led to commercial Baker’s yeast preparations which contain one or

more strains from the species S. cerevisiae. Through the addition of sourdough, other species can

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be incorporated in the bread making process like Pichia and Candida (De Vuyst and Neysens,

2005). In general, a yeast used for the bakery industry should fulfil specific requirements with

respect to the application and processing characteristics, such as adequate gas production to

ensure a uniform dough leavening, tolerance to a wide range of pH, temperature and salt/sugar

concentrations, as well as formation of desirable aroma compounds (Linko et al., 1997).

Specialised brewers and/or distillers yeast could be incorporated in the bread making process, but

because they are not adapted for the bread making process, it is common knowledge that they are

unsuitable due to their different metabolism and tolerances (Cauvain and Young, 2007).

Nevertheless, from the safety point of view European Union regulations allow these yeasts for

use in dough fermentation and bread production. In fact, the safety of food is a major concern of

consumers. Therefore, regulations and safety assessment guidelines are available in the European

Union. The Qualified Presumption of Safety (QPS) list summarises a wide variety of biological

agents including bacteria, yeasts, fungi and viruses that may be used in the food and feed chain

(EFSA, 2012). In the United States, food and substances used in food are regulated by the U.S.

Food and Drug Administration and are summarised in the Generally Regarded as Safe (GRAS)

status.

Interesting yeast species that could be used as an alternative to Saccharomyces include

Debaromyces, Kluyveromyces and Schizosaccharomyces. Heitmann et al., (2015) recently

studied the impact of different beer yeasts in comparison to Baker’s yeast on wheat bread

quality. This study showed that various beer yeasts are suitable for bread making and the

resulting wheat bread showed both superior and inferior characteristics in comparison to the

Baker’s yeast control bread. Nowadays, yeast is produced in a huge variety of different forms

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throughout the world. However, these “domestic” yeasts are different from “wild strains” due to

genetic modification and adaption, which allows them to grow in inappropriate situations (Ali et

al., 2012). The main formats in which yeast is available include fresh, compressed, active dry

and instant active dry yeast. The difference between these formats is related to their physical

appearance due to differences in moisture content. Reduction in moisture content is used to

prolong the shelf life of the strain but such preservation methods have an impact on yeast

performance factors such as metabolic activity, acid- and osmo-tolerance as well as temperature

stability (Cauvain and Young, 2007). Product shelf life ranges from 3 weeks (fresh and

compressed yeast) to 1 (dry yeast) or 2 (instant dry yeast) years (Hui, 2006). Fresh and

compressed yeasts are most commonly used in industry, since they are considered to be the most

reliable. The format of the yeast also has an influence on the fermentation intensity. Fresh yeast

produces the most carbon dioxide during fermentation resulting in superior dough-rising

capacity. Considering the fermentation speed, the yeast acts in the following order: compressed

yeast > instant active dry yeast > active dry yeast (Hui, 2006). Although fresh yeast is slightly

dehydrated it doesn’t need hydration time like dry yeast. The biggest problem is the shorter shelf

life of fresh yeast in comparison to processed yeast. Instant yeast is available since the 1960s and

is characterised by its very low moisture content and its fine particle size. In comparison to dried

yeast, instant yeast can be directly added to the flour and its main use is for bread and pizza

premixes (Cauvain and Young, 2007). Some studies have already investigated the impact of

these different formats on product quality parameters. Codina and Voica, (2010) studied the

impact of different yeast (S. cerevisiae) forms on carbon dioxide retention using a

rheofermentometer. They found that active dry yeast had the highest fermentation rate followed

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by compressed yeast and active instant dry yeast (Codina and Voica, 2010). Rollini et al., (2007)

analysed four commercial compressed Baker’s yeast (S. cerevisiae) strains which were originally

used in different applications (pastries, bread, frozen doughs and Panettone) and tested them in

complex dough formulations. They found that the different strains were indeed suitable for

different applications in contrast to what was indicated by the producer. However, all their

Baker’s yeasts belonged to the species S. cerevisiae and they showed variations regarding their

growth efficiency and gassing power.

To the authors’ knowledge, there is no yeast strain used in the baking industry which is

genetically modified. However Randez-Gil et al., (1999) showed that recombinant DNA

technology is a possibility of constructing new strains with improved suitability for the baking

industry.

3. Yeast vs. chemical leavening agents

Besides yeast, it is a common practice to produce leavened products using chemical leavening

systems which produce carbon dioxide either through chemical decomposition using heat or

through an acid-base reaction. For bakery products, particularly pastries, the two major gas

producing chemicals used are sodium bicarbonate (baking soda) and ammonium bicarbonate

(Amendola and Rees, 2003). Baking soda is a powerful leavening agent which starts as soon as it

comes into contact with an acidic environment like batter or dough (Amendola and Rees, 2003).

The disadvantages of these chemical leavening agents are the creation of off-flavours as well as

an over browning. An advantage is the short production time; no fermentation step has to be

included in the production, since some gas is already released at room temperature with the

majority released during baking. In comparison to yeast the release of gas is much faster and the

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gas cells are therefore much bigger. A few products naturally containing acids can be used for

leavening like lemon juice or sour milk. Other chemical leavening agents include salts of

phosphoric acid such as aluminium phosphate, mono-calcium phosphate, sodium acid

pyrophosphate and di-calcium phosphate. Mono-potassium tartrate (cream of tartar) and

glucono-delta-lactone (GDL) can be also used. The common baking powder consists of a

mixture of different acids, mostly in combination with baking soda and starch as a carrier to

separate the acids and bases to prevent premature reactions (Hui, 2006). Due to the neutralising

effect of the different ingredients, no off-flavour is left and the pH is not influenced (Amendola

and Rees, 2003). Similar to yeast, chemical leavening agents affect the structure, colour, flavour

and pH of the final product. Each leavening agent creates a slightly different texture, so when

choosing the appropriate leavening agent, the reaction rate and desired effects in the finished

products must be known (Hui, 2006). Plessas et al., (2005) produced leavened bread by using

kefir grains instead of yeast or chemical leavening. The produced bread showed a smaller

specific volume but better ability to retain moisture after production, with a firmer texture.

Another advantage was the lower acidity when using kefir grains with a positive effect on the

mould-free shelf life. This study highlights that other substrates should be considered as

leavening agents for baked goods beside Baker’s yeast and chemical leavening agents.

4. Main Metabolic pathways

Yeasts are facultative anaerobes which means that they can grow with or without oxygen. In

general, yeasts convert sugars into carbon dioxide, energy and biomass in the presence of

oxygen. In the absence of oxygen they use alcoholic fermentation to convert sugar into ethanol,

carbon dioxide and glycerol. The dominant fermentation products, which have the greatest

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impact on bread quality are carbon dioxide and ethanol (Pronk et al., 1996; Trevelyan and

Harrison, 1952). They are formed as soon as the yeast has been added to the dough/batter. S.

cerevisiae also produces a range of other secondary metabolites as glycerol, organic acids,

flavour compounds and precursors. The production of these compounds is linked to several

different metabolic pathways, like glycolysis, alcoholic fermentation, the tricarboxylic acid cycle

(TCA) and the glyoxylate cycle, which are summarised in Figure 1. The primary carbon

metabolism is performed by glycolysis. During glycolysis the yeast produces energy by

consuming low molecular sugars available in the dough (sucrose, maltose, glucose and fructose).

Hexoses such as glucose and fructose, are the preferentially utilised sugars which enter the

glycolytic metabolic pathway. However, glucose is preferred over fructose, since they are

transported with the same carrier into the cell which has a greater binding specificity for glucose

(Verstrepen et al., 2004). When glucose and fructose are consumed, the yeast starts to deplete

maltose but without hydrolysing it to glucose, as Baker’s yeast lacks the necessary enzyme. In

beer production the most fermentable sugars are maltose, maltotriose and glucose. Again,

glucose is the preferred sugar, but to obtain appropriate substantial alcohol content, the complete

fermentation of maltose and maltotriose is also required. Consequently, the majority of brewing

yeasts are able to ferment maltose and maltotriose after glucose. However, some yeast cells are

not able to take up maltotriose for their metabolism which can lead to difficulties in beer brewing

leading to lower ethanol yields or atypical beer flavours (Alves-Jr et al., 2007). To utilise maltose

the yeast requires an active transport system across the plasma membrane. Subsequently, the

maltose is hydrolysed by glucosidase enzymes (G) into two glucose molecules (Alves-Jr et al.,

2007). The repression of the synthesis of glucosidase enzymes is a major concern in limiting the

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dough fermentation rate (Dequin, 2001) which is also the reason for a lag phase in the carbon

dioxide production. Osinga et al., (1988) suggested a means of avoiding this lag phase by

replacing the promoters of the maltose permease and maltase with constitutive promoters to

increase the metabolic conversion of maltose. Other higher sugars like sucrose need to be

degraded by invertase (I) before the yeast can use them for metabolism. Therefore, the yeast

harbours two different invertase enzymes. One invertase is located in the cytoplasm of the yeast

cell and therefore it requires sucrose uptake. The second invertase is located between the plasma

membrane and cell wall. The hexoses formed by this enzyme are taken up by hexose transport

systems, and made available for yeast metabolism (Pronk et al., 1996). Codina and Voica, (2010)

showed that after mixing no sucrose was left in the dough samples due to the presence the yeast

invertase, which degraded the sucrose to glucose and fructose for yeast fermentation. Maltose

concentration increases during dough fermentation due to activity of amylases found in wheat

flour. A common pathway which is involved in all sugar-metabolising microorganisms is the

lower part of the Embden-Meyerhof pathway and the formation of pyruvate (Koshland and

Westheimer, 1950; Pronk et al., 1996). Pyruvate has a central position in many metabolic

pathways as it can be seen in Figure 1. The production of pyruvate and, therefore glycolysis,

plays a key role in the fermentation metabolism of yeast. The definition of glycolysis is well

known as a sequence of ten enzyme-catalysed reactions, which converts sugars like glucose to

pyruvate coupled with the production of ATP as an energy source. Di-hydroxy acetone

phosphate as an intermediate in glycolysis and a precursor of glycerol, a compound which plays

an important role in the cytosolic redox balance during anaerobic growth (Ansell et al., 1997;

Bakker et al., 2001; Nevoigt et al., 2002; Nevoigt and Stahl, 1997; van Dijken and Scheffers,

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1986). Di-hydroxy acetone phosphate is reduced to glycerol-3-phosphate by glycerol-3-

phosphate dehydrogenase (GPD) and finally dephosphorylated to glycerol by glycerol-3-

phosphatase (GP) (Nevoigt et al., 2002; Sigler and Hofer, 1991). In addition, during growth of

yeast, pyruvate is transformed into many different compounds, such as carbon dioxide, ethanol

and other organic metabolites, which have an influence on bread quality (Pronk et al., 1996).

Since yeast favours an alcoholic fermentation metabolism over respiration (“Crabtree effect”)

(De Deken, 1966), in the presence of high sugar concentrations the main metabolic pathway

which must be considered is the alcoholic fermentation, starting from pyruvate (Fiaux et al.,

2003; Gancedo, 1998). This “Crabtree effect” can cause several problems, such as an incomplete

fermentation, production of off-flavours, undesirable by-products and loss of biomass yield

(Verstrepen et al., 2004). During alcoholic fermentation ethanol is via pyruvate decarboxylase

(PDc) conversion of pyruvate into acetaldehyde and carbon dioxide. Further, alcohol

dehydrogenase (AD) reduces acetaldehyde to ethanol, by oxidation of NADH.

Another important metabolite is acetyl-CoA, which can be formed in two different pathways;

either from pyruvate (glycolysis) or acetaldehyde (alcoholic fermentation). In the latter pathway,

acetaldehyde is oxidised to acetate by acetaldehyde dehydrogenase (AcD). Acetate is converted

to acetyl-CoA by acetyl-CoA synthase (Ac-CoA-S). With lowering sugar concentrations the

yeast switches their metabolism from alcoholic fermentation to respiration which utilises the

tricarboxylic acid cycle (TCA), known as “diauxic shift” (DeRisi et al., 1997; Foulkes, 1951;

Galdieri et al., 2010; Gasch and Werner-Washburne, 2002). The production of acetyl-CoA from

pyruvate is performed by pyruvate dehydrogenase (PD). Acetyl-CoA can be used for the

production of fatty acids and fat. Thurston et al., (1982) showed that fatty acids are mainly

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produced during the first four hours of fermentation in beer with a four-fold increase over this

time. Another fate of acetyl-CoA is its funnelling into the TCA cycle within the mitochondria,

with the ultimate production of secondary metabolites and additional carbon dioxide. By

definition, the TCA cycle is known as a series of chemical reactions used for carbon dioxide and

ATP generation through oxidation of acetate. In this cycle, pyruvate is oxidised to carbon

dioxide and water with the concomitant production of ATP. Most of the carbon dioxide involved

in dough fermentation comes from alcoholic fermentation due to the “Crabtree effect” of yeast.

The primary role of the TCA cycle is production of additional ATP. The expression of genes

involved in the TCA cycle is down regulated during the first 30 min of dough fermentation,

however, because of the presence of glucose and oxygen, these enzymes still have a low level

activity which explains the production and excretion of organic acids such as citrate, malate and

succinate. Several research groups showed that the such organic acids are produced in the TCA

cycle (Arikawa et al., 1999a, 1999b; Whiting, 1976). Another enzyme in the TCA cycle is

aconitase (A), which converts citrate into isocitrate (Gangloff et al., 1990). Aconitase is located

in the mitochondria but also in the cytosol as part of the glyoxylate cycle (Duntze et al., 1969;

Regev-Rudzki et al., 2005). The enzyme isocitrate dehydrogenase (ID) oxidases isocitrate to α-

ketoglutarate, with the production of carbon dioxide, which also represents the starting point of

glutamate metabolism. Via the reductive pathway of the TCA cycle, beginning from

oxaloacetate, malate and fumarate can be produced (Arikawa et al., 1999a). Further oxidation to

succinyl-coenzyme A is catalysed by α-ketoglutarate dehydrogenase (αKD) with the production

of carbon dioxide. Beside the TCA cycle, the formation of succinate and malate by the enzymes

isocitrate lyase (IL) and malate synthase (MS) can take place in the glyoxylate cycle which

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occurs in the cytosol (Arikawa et al., 1999b; Fernandez et al., 1992). In addition to the

production of glycerol, ethanol and organic acids, the yeast is able to produce free amino acids

using the Ehrlich Neubauer-Fromherz pathway, which is linked to the shikimate pathway

(Herrmann and Weaver, 1999; Maga and Pomeranz, 1974). Amino acid biosynthesis is

controlled by about 30 enzymes involving different pathways (Pronk et al., 1996). Coming from

the amino acids flavour formation takes place. It can start with a deamination of free amino acids

like valine, leucine, phenylalanine or tryptophan followed by a decarboxylation, which can

produce aldehydes. These aldehydes can be reduced to higher alcohols (isobutyl alcohol, isoamyl

alcohol, phenylethanol) or transformed into acids by oxidation (Maga and Pomeranz, 1974). In

general, the biosynthesis of higher alcohols commences with a transamination reaction of amino

acid such as valine, leucine and phenylalanine and is catalysed by aminotransferases. The

produced α-keto acid is further converted by decarboxylation into fusel alcohols, and finally

reduced to higher alcohols via the Ehrlich pathway (Hazelwood et al., 2008; Procopio et al.,

2011). In addition, the corresponding organic acids can be produced, like phenylacetate,

hydroxyphenylacetate, or isobutyrate (Hazelwood et al., 2008).

5. Loaf volume/Cell structure

Achieving a desired loaf volume by yeast fermentation is only possible by providing a

favourable environment for yeast growth and for formation of gluten matrix that enables

maximum gas retention (Sahlström et al., 2004). The gas bubbles, which are incorporated in the

dough after mixing, grow during fermentation until they are saturated with carbon dioxide. This

growth leads to expanding of the dough and thinning of the dough matrix between the gas cells.

If over-fermentation occurs the dough is not capable of retaining the additional gas produced by

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the yeast and the gas bubbles fracture, which leads to a lower bread volume. The gas holding

capacity is an important characteristic for determining the bread quality and suitability of yeast

for baking. The more gas that is entrapped in the dough, the smaller the gas cells and the higher

their distribution is after proofing. These gas cells can resist more strength before they rupture,

which leads to lower extensibility and a higher specific volume (Dobraszczyk, 2003; Sroan et al.,

2009; Verheyen et al., 2014). During the baking process, the ethanol produced evaporates with

some of the water, which helps to develop the aerated structure of the cell crumb. It is well

known that dough mixing time can be reduced by adding instant active dry yeast, due to an effect

on the gluten network development (Pyler and Gorton, 2008a). In dried yeast some non-viable

cells are present which release glutathione as a stress response (Penninckx, 2002; Reed and

Nagodawithana, 1991; Verheyen et al., 2015). Rheological dough properties are influenced by

oxidising and reducing agents, which have an effect on the glutenin subunits that are linked by

disulphide bonds and can affect their degree of polymerization (Delcour and Hoseney, 2010).

The release of glutathione has a strong reducing effect and therefore increases the rate of thiol-

disulphide interchange reactions which leads to a modification of the viscoelastic gluten network

(Verheyen et al., 2015). As a result, gluten proteins with reduced size and lower molecular

weight are present (Delcour and Hoseney, 2010). For the reason, that rheological dough

properties are strongly influenced by thiol-disulphide exchange reactions; by removing thiol

groups the dough gets stronger Frater & Hird, (1963). Strong and weak flours differ in their

amount of protein-bond glutathione. Li et al., (2004) measured 10 different flours varying in

their amount of protein-bond glutathione; Only 5 flours resulted in bread doughs showing a

strong performance. They reported that flours with a significantly higher amount of protein-bond

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glutathione result in a strengthening effect on the dough and therefore a stronger gluten-network

development and better bread characteristics. Moreover, yeast is able to produce glycerol and

pyruvic acid in the early stage of fermentation (Whiting, 1976). Glycerol has a positive effect on

the texture of bread, especially during freezing. (Corsetti et al., 2000) reported that the addition

of glycerol reduces the firming of baked products during storage.

6. Flavour and aroma

Aroma and flavour are important quality parameters for bread. These are mainly affected by

ingredients and secondary fermentation products produced by yeast and generated under baking

conditions (Birch et al., 2013b; Frasse et al., 1993; Gassenmeier and Schieberle, 1995; Maeda et

al., 2009; Schieberle and Grosch, 1991). The most influential compounds are volatile compounds

like alcohols, aldehydes and ketones and non-volatile compounds like acids, esters, sugars,

phenolic compounds free fatty acids and lipids (Hui, 2006). Non-volatile compounds act mainly

as precursors for reactions that form new flavour compounds (Hui, 2006). Sugars remaining

from the fermentation have an effect on aroma due to their high reactivity in Maillard reactions

(Nilsson et al., 1987). The Maillard reaction is a complex mechanism between reducing sugars

like maltose, glucose and fructose and amino acids like leucine and phenylalanine, peptides

and/or proteins during baking, influencing the colour, flavour and nutritional properties of baked

products (O’Brien et al., 1989). Dough fermentation with yeast results in a decrease of the

concentration in free amino acid content. An increased amount could influence the aroma

through Maillard reactions and the Ehrlich pathway. Some volatile compounds are lost during

baking, while others form complexes with other dough constituents, thus affecting the flavour

profile of the final product. Not all components which contribute to the overall flavour and

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aroma of yeast leavened breads have been identified thus far. The total number of contributing

components is enormous and their specific interactions in flavour and aroma formation are still

not fully understood (Reed and Nagodawithana, 1991). A few authors have reviewed flavour

formation in bread (Cho and Peterson, 2010; Maga and Pomeranz, 1974; Pyler and Gorton,

2008b; Rothe, 1988). Most of the compounds responsible for aroma formation in bread crumb

made from yeast fermented dough result from yeast metabolism (Frasse et al., 1993; Schieberle

and Grosch, 1991), whereas the aroma compounds of the crust are products of Maillard reactions

(Purlis, 2010). The most significant compounds reported in the literature are alcohols and

aldehydes such as 2,3-butanedione and 3-hydroxy-2-butanone and esters which are produced by

yeast cells using the Ehrlich Pathway to degrade amino acids (Hazelwood et al., 2008).

Nowadays in the baking industry the trend is to use a short bread making process in terms of

fermentation, whereby the development of aroma and flavour is very limited (Cauvain and

Young, 2007; Maeda et al., 2009). The application of different bacterial starter cultures, such as

wine or beer yeast could compensate for these short fermentation process and produce flavour

and aroma during such short fermentations (McKinnon et al., 1996; Suomalainen and Lehtonen,

1978). Research on alcoholic beverage fermentation and production reveals that the choice of

starter cultures is an important factor, related to the formation of aroma and flavour in the final

product (Procopio et al., 2011; Suárez-Lepe and Morata, 2012). Several studies have dealt with

the effects of yeast on aroma development during the production of wine and beer (Molina et al.,

2007; Saerens et al., 2008). In these industries, yeast identification and strain characterisation is

essential, due to the wide variety of different flavour and aroma profiles yeast can impart

(Dashko et al., 2015; Huangl et al., 2010; Katarína et al., 2014; Pires et al., 2014; Vararu et al.,

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2016). In the recent years the aroma of bread gain more focus and recognition as an important

bread quality parameter (Birch et al., 2014, 2013a, 2013b).

Birch et al., (2013a) studied the influence of seven commercial compressed Baker’s yeasts on the

formation of bread aroma using dynamic headspace extraction. They found significant

differences in the aroma profile of the bread crumb with fermentation time, between the breads.

Furthermore, they stated that the choice of Baker’s yeast is a very important decision for the

bakers with respect to fermentation activity and aroma formation potential. Another study by the

same group showed that with increasing yeast concentration, the main flavour components like

2-methyl-1-propanol, 2-phenylethanol, phenylacetaldehyde, 2,3-butanedione, ethyl acetate, ethyl

3- methylbutanoate, ethyl hexanoate, ethyl octanoate and phenyl-ethyl acetate increase

concomitantly (Birch et al., 2013b). On the other hand, an increase in fermentation temperature

caused an increase in lipid oxidation products, which are often described as off-flavours.

However, their formation is independent of yeast concentration. It was suggested that short

fermentation time at low temperatures and high yeast concentrations could be used to develop a

bread with a high concentration of aroma compounds and less off-flavour. Thurston et al., (1982)

suggested a relationship between yeast’s fatty acids and the aromatic profile of fermented foods.

Such fatty acids have been shown to contribute to the production of fatty acid ethyl esters

especially in beer. They are connected to the yeast cell wall and can be released when yeast cells

dies. Fatty acid esters are secondary metabolites produced by yeast and many bacteria, which

play a key role in the flavour of alcoholic beverages. Ethyl esters of short and medium fatty acids

are important flavour compounds characterised by their strong fruity flavour. Beside their

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application in food and beverage production, they are also used by the cosmetic and

pharmaceutical industries.

7. Colour

Another important attribute for consumer’s acceptability is colour. Surface colour is of

considerable importance in the baking industry (Pathare et al., 2013; Purlis, 2010; Zanoni et al.,

1995) as it is the first parameter assessed by consumers. Colour formation depends on physico-

chemical characteristics such as moisture, pH, sugar concentration, amino acid content and the

process conditions used during production, like baking temperature, fermentation time and

temperature and starter culture (Zanoni et al., 1995). Colour formation results chemical,

biochemical, microbial and physical changes, which arise during production (Pathare et al.,

2013). Colour formation on the crust develops mainly throughout bread baking due to chemical

changes via the Maillard reactions (Purlis, 2010). Maillard reaction occurs between proteins and

carbohydrates at temperatures higher than 50 °C at a pH range of 4-7. Another important reaction

is caramelisation (Kroh, 1994; Zanoni et al., 1995), which is the direct degradation of sugars and

starch occurring in high-sugar foods at higher temperatures, >120 °C or 9<pH<3 (Kroh, 1994;

Zanoni et al., 1995). Both reactions appear concurrent and depend on the type of sugar and

amino acids present as well as the pH and water activity of the product (Zanoni et al., 1995). The

residual reducing sugars remaining after fermentation strongly influence the crust and crumb

colour (Finot, 1990; O’Brien et al., 1989). Due to increased mobility of reactants, the reaction

rate increases exponentially with higher moisture content, up to a maximum at 30% moisture

(Wolfrom et al., 1948; Wolfrom and Rooney, 1953). Both the initial pH of the product and the

buffering capacity of the system influence the rate and direction of the reaction. The rate of

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browning is low at acidic pH values and intensifies with increasing pH to a maximum at a pH of

~10 (Ashoor and Zent, 1984; Wolfrom et al., 1946). In general, the rate of Maillard reaction is if

excess reducing sugars are present rather than excess amino compounds (O’Brien et al., 1989).

Crust colour can be also controlled by using different starter cultures. Alpha amylase activity is

the main reaction to be considered in relation to crust colour formation, due to the production of

increasing amounts of maltose and dextrins, which participate both in the Maillard and

caramelisation reactions. Use of different starter cultures can have an influence on colour

formation due to differences in sugar metabolism (Goesaert et al., 2005; Heitmann et al., 2015;

Ormrod et al., 1991).

8. Shelf life

Shelf life is a parameter relating to the loss of perceived freshness. This can be correlated to

several different factors which are summarised in two different categories, staling and microbial

spoilage. These parameters will be discussed further in the next two paragraphs.

a. Staling/Hardness/Firmness

Modifications in crumb structure due to changes other than spoilage organisms, such as chemical

and physical changes of the crust (soft, leathery) and crumb (hard, dry, and crumbly) is referred

to as staling (Kulp and Ponte, 1981). Bread staling is mainly associated with the firming of the

crumb, which is an important factor in terms of consumer acceptability (Pateras, 2007). Although

bread staling is not yet completely understood, the baking industry uses different anti-staling

agents to inhibit staling. These include enzymes, alcohol, lipids, emulsifiers and sweeteners (Hui,

2006; Pateras, 2007). In particular, alpha-amylase is well known to retard crumb firming

(Giménez et al., 2007; Hui, 2006; Pateras, 2007). Lipases, lipoxygenases, endoxylanase,

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arabinosidase and protease are also known to prevent bread staling due to a crumb softening

effect. Heitmann et al., (2015) examined the effect of different yeast strains on bread hardness

during storage. By using different starter cultures they were able to produce a significant change

in crumb hardness, explained by the negative correlation of r = -0.90 (p < 0.05) between crumb

hardness and specific volume. It is known that breads produced with a bulk fermentation step

have a longer shelf life, due to larger amounts of alcohol produced during fermentation. Some

studies examining the effect of ethanol on bread staling, showed that the crumb modulus of

bread, treated with ethanol, increases during storage at a slower rate than the control bread using

a differential scanning calorimeter and crumb compressibility measurement (Fearn and Russell,

1982; Russell and Chorleywood, 1983). Russell and Chorleywood, (1983) showed, that bread

treated with ethanol firms at a slower rate than control bread. Increasing sugar and salt levels are

also known to slow the staling of baked products (Cairns et al., 1991; Taylor et al., 2009).

(I’Anson et al., 1990) reported a decreasing effect of ribose> sucrose> glucose on the

retrogradation of wheat starch, but the full mechanism of action is not fully understand. It is

suggested that sugars are able to increase the glass transition temperature and concurrently

decrease the diffusion of polymers to a crystal nucleus (I’Anson et al., 1990). On the other hand

(Levine and Slade, 1990) stated that sugars increase the glass transition temperature of the

amylose matrix and therefore the re-crystallization of amylopectin is repressed. Glycerol has

been reported to influence moisture distribution and staling of bread during storage. Yeast

leavened breads show a higher water content than unleavened breads which results in more

carbon dioxide and therefore a coarser bread crumb (Mondal and Datta, 2008).

b. Microbial spoilage

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Microbial spoilage is another important factor when considering bread shelf life due to post-

processing contamination (Pateras, 2007). Microbial spoilage is commonly caused by

microorganisms belonging to the species Aspergillus, Penicillium, Fusarium, Rhizopus, Mucor,

Endomyces and Cladosporium (Legan, 1993). Aside from the economic losses caused by these

microorganisms, consumers are concerned about the potential mycotoxins produced by these

microorganisms. Mycotoxins can cause severe health problems in humans (Legan, 1993; Pateras,

2007). The parameters determining the microbial shelf life are water activity, pH and storage

conditions. The most common method for preventing mould growth is the application of

chemical preservatives such as propionic acid and its salts or potassium sorbate. However, the

current trend is towards production without the use of additives. One solution is the

incorporation of sourdough as a natural bio-preservative to increase the mould-free shelf life of

baked products. Lactobacilli produce weak organic acids, other low molecular weight

compounds, peptides, cyclic dipeptides and proteins, which are known for their antifungal

activity (Axel et al., 2015, 2014; Magnusson and Schnürer, 2001; Niku-Paavola et al., 1999;

Okkers et al., 1999; Röcken and Vorsey, 1995; Stiles, 1996). Physical methods of prolonging the

shelf life are modified atmosphere packaging, pasteurisation or irradiation of packed bread

(Legan, 1993; Pateras, 2007). Some spoilage organisms such as spoilage yeast cause off-odours.

Post-processing contamination is likely due to physical contact with contaminated equipment.

The two main types of yeast associated in spoilage are filamentous yeast (“chalky moulds”) and

fermentative yeast (Berni and Scaramuzza, 2013; Pateras, 2007). S. cerevisiae is the most

common fermentative yeast spoilage organism, characterised by an alcoholic or estery off-odour.

Filamentous yeast like Pichia burtonii form white colonies on the surface of bread. This growth

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can be easily referred to as mould (Legan, 1993; Pateras, 2007). Berthels et al., (2004) identified

yeast strains with a discrepancy in their consumption preference for glucose and fructose, and

found the ability of such strains to reduce residual fructose levels and increase ethanol yield to be

helpful in partially solving the spoilage problem. They suggested use of such strains as a criteria

for selection of new yeast strains for wine production. Heitmann et al., (2015) demonstrated the

effect of different S. cerevisiae strains, originating from the brewing industry, on the shelf life of

white wheat bread. They showed both inferior and superior behaviour in terms of mold-free shelf

life of the breads, which ranged between 3 and 5 days. They also demonstrated differences

ability to propagate mould on breads baked with the different S. cerevisiae strains.

It is well known that some fungi and bacteria are able to produce secondary metabolites with

antifungal properties. Nowadays most of the attention has been given to antifungal lactic acid

bacteria present in sourdough bread and little attention has been given to yeast as possible

producers of antifungal compounds. Yeasts, however, are promising candidates as fungicides.

Coda et al., (2013) screened 146 different yeast strains (genera including Candida,

Metschnikovia, Debaromyces, Pichia and Kazachstania) focusing on antifungal activity against

Penicillium roqueforti. They found six Meyerozyma guilliermondii with noticeable in vitro

activity. Their work showed the possibility of extending shelf-life of baked goods using M.

guilliermondii LCF1353 as a mixed starter while maintaining optimal taste and structure at the

same time. In another study the same group demonstrated, similarly, the potential of

Wickerhamomyces anomalus as a mixed starter to extend the shelf-life of baked goods (Coda et

al., 2011). Mo & Sung, (2014) investigated Pichia anomala SKM-T, which is known for its

antagonistic properties against some spoilage moulds like Penicilium paneum KACC44834, and

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found it to be suitable as a leavening agent for the production of white pan bread. The bread

containing this strain exhibited less P. paneum spoilage colonies on the surface than bread baked

with S. cerevisiae, due to a production of the flavour compounds 2-phenylethyl acetate, 2-

phenylethyl alcohol, 2-decenal and nonanal, which enabled a shelf life extension. The production

of antifungal compounds is considered in the selection of yeast strains for wine production.

However, it is not a characteristic considered when choosing Baker’s yeast.

9. Nutrition

Since ancient times, cereals have been a staple in the human diet. They are considered as an

important source of energy and supply macronutrients including complex carbohydrates, fibre,

protein as well as micronutrients such as calcium, phosphorus, iron, sodium, magnesium and

potassium. Cereal grains can be considered a source of vitamins, especially B vitamins like

thiamine (vit. B1), riboflavin (vit. B2) and niacin (vit. B3) (Cauvain and Young, 2007). Yeast

represent a nutritional source of carbohydrates, fats, vitamins, especially B vitamins, minerals

and amino acids, in particular, lysine (Rincón and Benítez, 2001). Studies on the effects of cereal

fermentation on nutritional quality are scarce. Due to bread being a staple food it represents an

important means to supplementing human nutrition. During fermentation yeast can have an

effect on the levels vitamins, phenolic compounds, phytates and folates, which is discussed more

detail below.

During the production of yeasted bread, a 48% loss of thiamine (vit. B1) and pyridoxine (vit. B6)

were observed (Batifoulier et al., 2005). However, a longer fermentation increased the levels

again. Compared to thiamine (vit. B1) and pyridoxine (vit. B6), folate (vit. B9) showed good

stability during bread making and an increased amount could be found in comparison to the flour

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(Osseyi et al., 2001). The content of thiamine (vit. B1) has also been reported to decrease in the

wheat and rye baking process, (Martinez-Villaluenga et al., 2009) but to increase with a longer

fermentation time (Batifoulier et al., 2005). The fermentation step can therefore have an effect on

the overall formation or retention of vitamins during baking. A short baking process was also

presented to reduce the content of thiamine (vit. B1) in whole-wheat, but a prolonged yeast or

sourdough fermentation maintained its levels (Batifoulier et al., 2005). Batifoulier et al., (2005)

also found that the thiamine content was increased when fermentation time was prolonged and

that the increase was significantly higher in white bread with yeast compared to sourdough,

despite comparable vitamin production by the microorganisms (0.25 mg/g dry matter). Long

fermentations could support a net synthesis of thiamine by yeast, while fermentation with lactic

acid production in sourdough bread could origin in a decrease of thiamine (Khetarpaul and

Chauhan, 1989). In contrast to these findings, (Rucker et al., 2006) reported a 35% loss of

thiamine during bread making. A small amount of the riboflavin (B2) in bread derives from

yeast. As a result, bread often contains more riboflavin than the original flour. Sourdough

fermentation does not lead to any enrichment of riboflavin (Batifoulier et al., 2005). Whole-

wheat bread making with yeast (from kneading to final bread) undergoing a long fermentation

process, resulted in a 30 % enrichment in riboflavin. The use of yeast and sourdough during

fermentation did not show a synergistic effect on B vitamin levels (Batifoulier et al., 2005), but a

longer fermentation time could increase the level of pyridoxine (Batifoulier et al., 2005). Yeast

fermentation has been shown to result in an increase of folate in the baking process of wheat and

rye breads (Kariluoto et al., 2006). Kariluoto et al., (2006) investigated the ability of yeasts and

lactic acid bacteria to have an influence on the folate content in a rye sourdough and showed that

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the effects of sourdough bacteria are negligible. Proofing does not influence the total folate

content but changes in vitamin distribution were observed. Folate losses during baking were

about 25 % (Kariluoto et al., 2004). However, the synthesis of folate by yeast results in an

increase of the content over three-fold in bread. Another important advantage of yeast

fermentation is the reduction of phytates (phytic acid) by phytase activity which results in an

increase of the bioavailability of magnesium and phosphorus. However, phytase activity depends

on the substrate flour, proofing temperature and time as well as dough pH and the amount of

yeast (Pozrl et al., 2009). Commercial Baker’s yeast has been shown to express phytase activity

(Tu et al., 2000). A wide variation in phytase activity was identified in sourdough starters

containing both yeast and lactic acid bacteria (Chaoui et al., 2003; Reale et al., 2004). Another

potential suggestion was the use of high-phytase yeast strains to act as phytase carriers in the

gastrointestinal tract. The reduction of phytic acid has repeatedly been reported in yeast and

sourdough processes. Although yeast fermentation reduces the unfavourable effects of phytic

acid, sourdough bread seems to be a better source of available minerals, especially magnesium,

iron and zinc (De Angelis et al., 2003; Turk et al., 1999). Therefore, it should be possible to

control the phytase activity by modifying the process conditions or by selecting specific

microbial starters. Losses have been observed for tocopherol (vit. E) during sourdough

preparation and dough making (Wennermark and Jägerstad, 1992). Katina et al., (2007) observed

reduction in tocopherol (vit. E) and tocotrienol (vit. E) content. This may have been due to

oxygen sensitivity. Fermentation has been shown to increase the antioxidant activity in the

methanol extracted fraction of rye sourdough, concurrent with increased levels of easily

extractable phenolic compounds (Katina et al., 2007). A reduction in kneading time combined

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with a longer fermentation time could be able to retain carotenoids and vitamin E contents.

Fermentation of rye bran with yeast was also shown to increase the level of free ferulic acid

(Katina et al., 2007). The antioxidant capacity of rye breads baked with sourdough showed an

increase than that of white wheat bread. The highest values were reported for breads using

wholemeal flour (Martinez-Villaluenga et al., 2009; Michalska et al., 2007). Recently, it was

shown that a yeast fermentation using wheat bran together with cell wall hydrolytic enzymes

increased the bioaccessibility of phenolic compounds in breads as well as the metabolite 3-

phenylpropionic (Anson et al., 2009). (Poutanen et al., 2009) An increase in free ferulic acid was

observed as a result of dough mixing and proofing. However, the amount of released ferulic acid

was about 1 % of the total amount of ferulic acid originate from wholemeal rye. An increase of

the levels of total phenolic compounds and free phenolic acids could be found by sourdough and

yeast fermentation of wholemeal rye (Katina et al., 2007). In contrast, Boskov Hansen et al.,

(2002) did not observe a significant change in the content of phenolic acids during dough

proofing. Baking showed a slightly increase of the concentration of phenolic compounds in the

crust, probably through Maillard reaction (Gélinas and McKinnon, 2006). However, this effect

was not detected in wholemeal bread (Boskov Hansen et al., 2002; Dewettinck et al., 2008;

Gélinas and McKinnon, 2006). One other study used different yeast strains for the production of

selenium enriched baked products. Stabnikova et al., (2008) used a yeast, which biomass was

enriched with organic forms of selenium, to increase the amount of selenium in bread. The non-

protein monocarboxylic acid, γ–aminobutyric acid (GABA), plays an important role in the

animal and human nervous system as a neurotransmitter. An increased intake of GABA can be

related to different health benefits, such as lowering of blood pressure, prevention of diabetes,

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inhibition of leukaemia cell proliferation and cancer cell apoptosis. Collar et al., (1992) and

Benedito De Barber et al., (1989) suggested, however, that GABA is rapidly consumed by yeast

at the beginning of a fermentation, due to the high demand of nitrogen for cell growth, or takes

part in the Maillard reaction. More recently Lamberts et al., (2012) showed the important role of

yeast in the GABA dynamics during bread making. During dough mixing the level of GABA is

increasing, but during fermentation yeast consumes it as a nitrogen source. However, the authors

were able to produce GABA enriched bread through the addition of exogenous glutamic acid

decarboxylase (GAD) in the recipe. Hudec et al., (2015) screened different microorganisms from

10 different food applications as well as seven pure bacterial strains for GABA. They showed a

small production of GABA from S. cerevisiae Baker’s yeast and wine yeast of 0.8 and 1.3%,

respectively. The highest selectivity of GABA production could be detected using Lactobacillus

delbrueckii subsp. bulgaricus of 90.0 %. Using strains from the genera Lactobacillus, via

sourdough production, could be a good alternative to increase the GABA content in bread.

Rizzello et al., (2008) previously reported a GABA concentration of 258.7 mg/kg in a wholemeal

wheat sourdough by the addition of an adjunct culture using lactic acid bacteria.

10. Conclusion

The baking industry is currently selecting their yeast strains based on their ability to ferment

sugars anaerobically with adequate gas production. However, other important quality parameters

for consumer acceptance of bread including colour, texture and flavour, are not considered when

selecting yeast strains. At the moment the production of additional metabolites by yeast plays an

underestimated role in the selection of strains. Wine yeasts have been traditionally selected by

their fermentative power, suitable fermentation kinetics, in additional to their low acetic acid

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production and resistance to sulphur dioxide (Suárez-Lepe and Morata, 2012). Recently, new

selection criteria have been sought to improve the technological properties and sensorial features

of wines, since the metabolic uniqueness and physiological properties of yeast could, through the

production of metabolites, improve the sensorial properties of wine. Included in these criteria are

the ability to enhance wine colour, the absence of β-glucosidase activity, the facilitation of

colloidal stabilisation in red wines, the appropriate enhancement of aroma via the production of

volatile compounds and the provision of structure and body (Suárez-Lepe and Morata, 2012).

Similarly detailed selection criteria are commonplace in the production of beer. The brewing

industry, in general, separates yeast strains into ale and lager yeasts. In addition, they also use

more specific selection criteria, such as the fermentation behaviour (top or bottom fermentation),

fermentation performance (fermentation rate and degree of attenuation), the ability to ferment

meliobiose, temperature tolerance, ability to flocculate (powdery or flocculant yeast), oxygen

requirements and the ability to form or remove fermentation metabolites (aroma compound

formation) (Bokulich and Bamforth, 2013; Kunze, 2014). Therefore, specific selection of

Baker’s yeast should be as carefully considered as it is done for wine and beer yeasts,

particularly in terms of flavour, colour and shelf life. The wine industry has also recognised the

potential of non-Saccharomyces yeast strains, which haven’t yet been studied in the process of

bread making (Suárez-Lepe and Morata, 2012). Randez-Gil et al., (1999) previously suggested to

use recombinant DNA technology for the creation of new yeast strains expressing enzymes to

allow elimination of the extensive use of baking additives. Choosing the perfect starter culture

for bread/baked product manufacturing should not solely be determined by the gas production

capacity during fermentation. Other characteristics like enzyme activity are an important

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parameters to predict the final bread quality, due to their impact on shelf life (microbial and

staling) as well as colour and flavour formation. Consumer acceptance will not allow use of

genetically engineered yeasts. More targeted yeast selection, based on broader criteria, offers a

good way to obtain yeast strains from the species S. cerevisiae (even other genera and species)

with novel technological properties, within the limitation of current Food Legislation. Such

strains should enable improvements in the technological and/or sensorial qualities of baked

products.

11. Acknowledgment

The authors want to thank Claudia Axel and Kieran Lynch for correcting the manuscript. This

study was financed by the Seventh framework Program of the European Community for

research, technological development and demonstration activities (2013-2017); specific program

“FLOURplus” Intelligent and easy tool to categorise and characterise flour quality for consumer-

driven wheat baked goods in European SME-bakery and cereal sector (606198).

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Figure 1. Schematic representation of the most important metabolic pathways, following the

carbohydrate dissimilation, their enzymes (Abbreviation) and references in Saccharomyces

cerevisiae influencing bread quality parameters.

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