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    Sood Si Technol (December 2015) $2(12}: 7588-7598 OL 10,10071813197.015-1977-y REVIEW. Flavor impacts of glycerol in the processing of yeast fermented beverages: a review Xiangdong Zhao! - Susanne Procopio! « ‘Thomas Becker! evied: $ July 2015 /Acepted: 21 July 2015 /Published online: | August 2015 (© Association of Food Ssintts& Technologists (India) 2015 Abstraet Glycerol conrbutes to the beverage body’ and fll- ness, Moreover, italsoinuences the Navor intensity. As a major byproduct, glycerol not only serves critical role in yeast osma- regulation and redox balancing, but also acts as the carbon come pettor against ethanol in alcoholic fermentation. Therefor, ine creasing glyeerol yield benefits both the flavor and ethanol re- duction forthe fermented beverages. Glycerol yield has been clevated either by fermentation optimization or by yeast genetic ‘modification, The fermentation optimizations reached maxi- ‘mum 14 g/L glycerol through screening yeast stains and opti- mizing fermentation parameters. Meanwhile the yeast overex- pressing GPDI (encoding glyeerol-t-phosphate dehydroge- nase) produced upto 6 folds more glycerol fr beer and wine. Except for glycerol improvement, the geneticelly modified yeasts accumulated dramatically undesirable compounds such as acetaldehyde, acetate and acetoin which are detrimental for beverage favor. Tn comparison, the natural high elyerolpro- ducers showed stain.speciic manner onthe yeast-derved aro- rma compounds Fike volatile aids, fase! aleohols, esters, and aldehydes. Temperature, sugar concestation, nitrogen compo- sition, oxygen and pH-vaue, which influence glycerol biosyne thesis, also obtained various effects onthe production of aromat- ic compounds. Inthe current review, we fist deliberate the Highlights: > Giycerol and major beverage flavor interactions “ Impats of glyceral optimizations on the genes transcription level + Changes of aroma compounds formation during glycerol yield ‘optimizations + Glycerol yield improvements in beverage fermentations Susanne Procopio ‘susan procopiogum de Lebel fir Brau- nd Gstrinksteshnologie, Technische Univesitt “Miinchen, Wethenstephan, Wethestephane Stig 20, 5354 Freising, Germany 2 Springer ‘organoleptic contributions of glycerol for fermented beverages, Furthermore, glycerol optimization stategies are discussed re- garding to the yield improvement, the genes expressions, the ‘overall flavor impacts and the fesibilities in beverage applica tions. Lastly, for improving beverage flavor by glycerol optimi- zation, a high-throughput platform is proposed to increase the sereening capacity of yeast strains and parameters in the pro- cessing of fermented beverages. Keywords Glycerol - Yeast - Fermentation - Flavor Beverage » High throughput screening (HTS) Introduction As a viscous polyalcohol with slight sweet taste, glycerol mainly contributes to the body for fermented beverage (Scanes et al. 1998), Additionally, it has been also reported to enhance the flavor intensity (Jones etal. 2008; Omori etal 1995), suppress the perceived roughness (Jones tal. 2008; Peleg et al. 1999), inerease the worty off-lavor retention and influence on the aroma volatility (Jones et al. 2008; Perpete and Collin 1999; Perpete and Collin 2000), Glyceral was ranging from 1 to 3 g/L against its claimed threshold of 10 giL in beers (Briggs et al. 2004; Nykanen and Suomaleinen 1983; Parker and Richardson 1970). In wine, the general concentration was across 4 to 15 g/L opposing to the thresholds at 5.2 and 9 giL (Nieuwoudt et al. 2002; Noble and Bursick 1984; Nykiinen and Suomaleinen 1983), For oth- cr fermented beverages, approximate 2 to 6 giL could be found in Asian rice wines, cider and barley shochu, a Japanese spirit (Chen and Xu 2010; del Campo et al. 2003; Liu et al. 2014; Omori et al. 1995), Glycerol serves two vital physiological roles during yeast fermentation. The accumulation of osmolyte glycerol allows 4 Food Si Technol (Desember 2015) $2(12): 7548-1598 1589 ‘yeast re-adjust osmotic gradient across he cell membrane to remain cellular functions under osmotic stress condition (Albertyn etal, 1994; Ansell etal, 1997), Glycerol is also involving in the maintenance of eytosolic redox balance for that the surplus NADH ean be reoxiized via glycerol synthe- sis by serving as the reaction cofactor (Albers eta 1996). Interestingly, more glyeerol formation also means that less carbon source will be diverted to ethanol synthesis in aleohol- ic fermentation [Namerous efforts have been devoted to redirect the glyco! ysis flow towards glycerol for beverage flavor improvements and ethanol reduction. The major strategy is by optimizing the yeast culture conditions. Up 10 14 g/L has been observed by Sscrecning yeast stains followed by optimizations on the fr= mentation parameters and substrates (Berovié t al. 2007; Erasmus etal, 2004; Nieuwoudt etal, 2002), Primarily, the prolonged or increased transcription level of GPDI, which encodes NAD-dependent glyeerol-3-phosphate dehydroge- DH), was responsible for the elevated glycerol level under the optimized conditions (Du etal. 2011; Kajiwara tal 2000). However, except fr glyeerol, the optimized conditions also impacted on the yield of fermentative favor compounds. ‘The higher temperature and higher sugar concentration, which were in favor of glycerol formation, led to over accumulation of acetic acid, actaléchyée and higher alcohols, Even worse, inadequate optimizations aso impaired yeast grow and fure ther affected the fermentation kinetics and the overall flavor balance (Csutonis et al. 2014; Du etal. 20115 Liu ta. 2 Piddocke etal, 2009; Yu etal. 2012) Glycerol overproduction has been realized by the yeast ‘overexpressing genes involved in glycerol synthesis, disrup- tion oF deletion of the genes in glycolysis and fermentation pathways (Eglinton etal. 2002; Ehsani et al. 2009; Nevoigt et al, 2002; Nevoigt and Stahl 1996; Varela et al, 2012). Alternatively, fermentative glycerol has been also ‘overproduced by raising the surplus intracellular NADH and by “opening” the flux facilitator (Overkamp et al. 2000; ‘Varela tal, 2012). The genetically modified yeasts promoted lyeeto yield up t 5.6 and 6.1 folds and decrease ethanol concentration by 18 % and 21 % under beer and wine fermen- tations individually (Nevoigt etal. 2002; Varela et al. 2012) Nonetheless, the engineered strains accumulated dramatic amounts of detrimental flavors such acetaldehyde, acctate and acetoin in auton to the impaired growth ofthe yeast and poor social acceptance for their application (Cambon etal 2006; Eglinton etal. 2002; Lopes et al. 2000; Nevoigt et al. 2002; Remize et al, 1999; Varela et al, 2012) In order to realize beverage flavor improvement by glyoet= ol optimization, we present this review fry, to diseuss the ‘favor impacts of glycerol for fermented beverages based on the analytical and sensory studies. Secondly, strategies for slyeerol optimization are evaluated respecting othe effective- ness, side effects and feasibilites for beverage application. Lastly, we propose a high-throughput platform for optimizing _slycerol yield with flavor consideration in the processing of fermented beverages Sensory contribution of glycerol to fermented beverages lycerol has favorable impacts on the quality of beer, wine and spirit (Langstaf etal. 1991; Liu et al. 2014; Nieuwoudt et al, 2002; Omori et al. 1995; Omori et al. 1996), Large surveys showed that obvious higher level of glycerol was present in wines of superior grade than that of ordinary (Nicuwoudt et al. 2002; Nykinen and Suomaleinen 1983), Nevertheless, flavor profile is unique for cach wine. Individual wine, as well as other beverage, has debating ben- fits from glycerol, ‘Wide agreement on the body contribution Itis widely accepted for the contribution of glycerol to bever- ages body and fullness. Glycerol was reported to contribute to the “mouth-feel”, “viscosity”, “density”, “smoothness” and “roundness”, These sensory terms were closely correlated in subordinate or synonymic manner (Gawel etal. 2007; Jones tal, 2008; Langstaff eal, 1991; Nurgel and Pickering 2005), In beers, glycerol was found significant correlated to the den- sity and viscosity which were categorized as the mouth-feel attributes (Langstaff etal. 1991), Alcohol-free beer benefited from glycerol addition across a range fom 0.3 % to 2.0 % ‘with an improved body and fuliness (Sohrabvandi etal. 2010), For wines, although the majority research has reached an agreement on the mouth-fecl contribution (Gawel ct al 2007; Jones et al. 2008; Noble and Bursick 1984; Nurgel and Pickering 2005), the effective levels of glycerol varied even contradicted, Over 25 gil. glycerol was required for a noticeable effect in the perceived viscosity while the decrease ‘rend was observed in the range of 0-15 g/L in the wine (Noble and Bursick 1984; Nurgel and Pickering 2005). However, recent researches revealed that practical level of | _lycerol at around 10 g/L. was already enough to produce a noticeable improvement of the perceived viscosity (Gawel et al. 2007; Jones etal. 2008), Beverage body benefits from glycerol in certain concentra- tions or ranges. It might because of that the viscosity and density of glycerol are higher than most of the fermented beverages. In addition, by balancing the alcoholic strength and astringency, glycerol conferred a degree of “smoothness” and “roundness” on the palate (Jones et al. 2008). Glycerol ‘was also found in the total extract of beverages and might participate in the fullness contribution in associated with high extract content (Nieuwoudt et al. 2002). Nevertheless, the body contribution was highly dependent on the individual D Springer 7590 4 Food Sei Technol (Docensber 2015) $2(12): 7588-7598 ‘wine (Gawel et al. 2007; Jones et al, 2008) It was rather less consistent for sweet wines than that for the dry wines, For ‘example, glycerol has a doubtful contribution on the mouth: feel of ice wine since the high contents of aleohol and resid sugars tended to deliver greater impacts on the viscosity and body (Erasmus et al. 2004), This observation partly explains the controversial concentrations of glycerol for perceived viscosity Debate on the flavor impact Glycerol influences beverage flavor. Glycerol addition above ‘or below the threshold (10 g/L) was found to modify the beer flavor (Parker and Richardson 1970). The overall flavor inten sity was positively influenced by glycerol (10 g/L) especially ‘when the volatile concentration was low in wine (lones etal 2008). Glycerol together with other components were indicat- ced to responsible forthe delicate tastes and flavors of Chinese rice wine (Chen and Xu 2010; Liu et al. 2014). The flavor ‘compounds of shochu was enhanced by varying glycerol con- centration in the mash (Omori et al. 1994), Exeept for the impacts on overall flavor profile, glycerol also affected the perception of specific flavors, The sweetness threshold of selycerol in wine was revealed at 5.2 g/L in earlier study (Noble and Bursick 1984). However, no obvious effect on sweetness perception was observed in the model white wine by 10 giL, in Riesling wines by 10.2 g/l and in ice wines by 8 giL in recent investigations (Erasmus etal. 2004; Gawel et al. 2007; Jones et al, 2008). Neither acidity was affected (ones et al. 2008). For the bittemess, glycerol was reported to suppress bitter and roughness perception of a model wine ones et al. 2008; Noble 1996). Notice that, the perception of individual flavor is highly depend on the overall beverage flavor matrix, In addition, the changed sensations may also ddue to the increased viscosity. For that reduced perception ‘on astringency and flavor intensity were reported in the high viscous solution although no measurable changes were pro- vvided by the analytical results (Hollowood etal. 2002; Peleg etal. 1999), Glycerol may have an obscure and indirect role on the aroma volatility, Results ffom both sensory and analytical in- vestigations denied the detectable effet of glycerol on the perception of overall aroma intensity of wines (Gawel ct al 2007; Jones et al. 2008; Lubbers et al. 2001), However, at lower level of volatiles, ethanol seemed have enhanced the ‘overall aroma in the presence of glycerol while suppressed it inits absence (Jones etal. 2008). Additionally, the volatility of two duty aroma (3-methyl butyl acetate and ethyl hexanoate) ‘were positively influenced by glycerol concentration ranging, from 01015 g/L although no overall aroma impacts have been claimed for the wine (Lubbers etal. 2001). The above obser- vations highlight the flavor-binding effect of glycerol (lones et al. 2008; Lubbers et al. 2001; Nawar 1971). 50 % glycerol ‘was reported to change headspace concentration of the vola- tiles such as 2-heptanone and 2-pentanone (Nawar 1971). The esters concentrations in distillate were also affected by the amounts of glycerol in the mash (Omori et al. 1995; Omori et al, 1994), The isoamnyl acetate concentration in the barley shochu was highest at 10 g/l of glycerol, while that of B- phenylethyl acetate increased gradually when increased glyc- cero] from 6 to 14 g/L in the mash (Omori et al. 1995). The concentration of octenol, 2-ethylhexanol, B-phenylethyl aleo- hhol and some fatty acid ethyl esters were influenced by the ~lycerol concentration inthe mash as well (Omori etal. 1995; ‘Omori et a. 1994; Omori et a. 1996) ‘Glycerol was found to increase the worty offflavor retention for the non-alcoholic beer (Perpete and Collin 1999; Perpete and Collin 2000), 3-methylbutanal and 2-methylbutanal were described a the predominant compounds responsible for the ‘worty taste in alcohol-free beer (Perpete and Collin 1999; Sohrabvandi et al, 2010), In regular beers, ethanol restored the thresholds of higher aldehyde flavors. However, both the absence of ethanol and the higher residual sugar could strength- ‘en worty off-flavor in the low-non-aleoholie beers, Glycerol seemed to have an cthanol-mimic effect which was able to increase the aldchydes retention as low as the level of 0.5 and 45 % of glycerol retained up to 40 % of the investigated aldehydes (Perpete and Collin 2000), So far, the flavorimpacts of glycerol, especially the optimal ranges of such impacts, still remain as a topic for debate. By varying glycerol and ethanol concentrations in 20 wines, the highest desirability were obtained at 6.19 gil. (Kaur et al 2013). The preference shows that glycerol in certain concen- tration may conduct an optimum sensory impacts for the fermented beverages. This range is highly depending on the beverage type, on the production methods and most ofall on the overall flavor matrix, Further sensory investigations should be conducted to clarify the optimal concentration range of glycerol in specific fermented beverages, especially in beers which are comparably lacking of attention, Moreover, the sensory research in the ow-/non- alcoholic beverages tends to be more attractive considering the body and flavor contributions of glycerol Increasing glycerol by fermentation optimization ‘Glycerol isthe thitd largest fermentation product after ethanol ‘and carbon dioxide, As illustrated in Fig, 1, the phosphorylat- ced glucose is converted into dihydroxyacetone phosphate (DHAP) and glyceroladchyde-3-phosphate (GAP) in equimo- Tar amounts. However, most of the DHAP is isomerized into GAP which is further converted towards ethanol. On the other side, the rest of DHAP is transformed into glycerol via two steps of enzymatic reactions. The first step transforms the DHAP into glycerol-3-phosphate (G3P) with NADH as a 4 Food Sl Technol (December 2015) $2(12) 7588-1598 1591 roy \ “ES ole. See. Le \ pee ese IE ae | Fig. 1 Glycol biosynthesis in respond t osmotie stress and rodox statue andthe interelationship with yeat-derived aroma compounds in 5 eereusige NADH: reduced form oF nicotinamide adenine dinucleotide, ‘TCA: tncaboxylic acid eyele: Acety-CoA: acetyloenzyme A: Acy! (CoA: aeylooenzyme A; Undetne gees: the expressions regulated by osmotic sess; Solid line: metabolic reaction(s; Dask line: schematic csmorogulaory signal uansducton caseade (HOG signaling pathway), “Many rections sogenes, ATP, Pi,CO; and CoA are omitted i the figure for clarity. The organoleptic impression ofthe aromi-active metabolites in associate with glyerol sensory contributions and yield optimizations reaction cofactor. This step was reported as the rate limiting reaction for glycerol synthesis. The second step is to phos- phate G3P into glycerol (Albertyn et al. 1994; Ansell etal 1997; Cordier etl. 2007). Two paits of isogenes are involved in glycerol synthesis: GPDJ/GPD2 encoding GPDH and GPPIIGPP2 encoding the paralogs of glycerol-5-phospha- tase. The isogenes work in a cross combination manner and serve distinct physiological roles. The GPDI follows by GPP2 are essential for the growth under osmotic stress con- ditions while their expressions are regulated by high osmolar- ity glycerol (HOG) response pathway, whereas the GPD2 and GPPI axe primatily functioned in maintaining the redox bal- ance under anaerobic conditions (Albertyn etal. 1994; Ansell etal. 1997; Pahiman etal. 2001; Remize etal. 1999), Glycerol biosynthesis is predominantly strain-dependent Variances of industry strains in glycerol and flavor compounds production ‘The predominant role of yeast strains is confirmed by the results that the strains which were initially selected as high cor low glycerol producers remained their ability in glycerol biosynthesis in different cultivation conditions (Du etal. 20115, Erasmus etal, 2004; Radler and Schitz 1982; Remize et al 2000). The trancriptomie study showed that, except for the difference in intracellular glycerol level, the six wine strains showed remarkably differences in th transcription level ofthe (Briggs otal, 2008) vicinal diketones: diacetyl (buster, actoin (msn), 2.-butanediol (slighty biter, relative neural); Organic acids: ats, male, lactic and cic acids (Sou), sucinic acid (iter sal), Aldehydes: acetaldehyde (grassy), 3-methylbutaal (mally, chocolare- ie), 2-metnyoutanal (mal, cheesy), Higher alobols: oetenal earthy, ‘mushroom-lie), 2-ethylhexanol (mushroomlike, sweet fruit), phenylethyl alcohol (rosy, honey-like), Acetate estes: ethyl acetate (solvent, isoaryl acetate ot Samehyl butyl sectate (banan-lk, Bephenyl ethyl acetate (ras); Faty acid ethyl ester ethyl hoxanoate (applet) genes GPD1, GPD2, STLI (encoding a glycerol-proton symportr), FPS1 (encoding the aquaglyceroporin, a plasma ‘membrane channel involved inthe influx and efflux of glve- exo) and GUPI (encoding the plasma membrane protein i volving in active glycerol uptake) (Fig. 1) (Jiménez-Mar tal, 2011). High activity of GPDH was found in the high alyeero! producers. In comparison, forthe strains formed very litle glycerol, GPDH was hardly detectable though excep- tions existed (Radler and Schit 1982), Furthermore, the high slycerol producing strains tended to acquire higher ability for the adaption of osmotic stress or lower ability to retain glye- rol as osmolyte in the cytoplasm (Brasmus et al. 2004). Meanwhile, higher ratio of glycerol to ethanol was observed in those strains although the alcohol dehydrogenase (ADH) activity seemed not correlated with yeast ability of glycerol production (Chen and Xu 2010; Raller and Sehiitz 1982), In audition, unlike the engineered strains, the natural high glye- «rol producers did not show collectively higher concentration of acetic acid, succinic acid, acetaldehyde and acetoin (Du tal 2011; Brasmus etal, 2004; Remize et al. 2000). The formation ofthe yeast derived flavor-ctive compounds were not related with glycerol, To some extent, the productions of higher alcohols, estrs and organic acids were also strane dependent (Chen and Xu 2010; Du et al. 2011; Erasmus etal. 2004; Bstéver etal, 2004; Radler and Schitz 1982; Remize et sl 2000), Therefor, screening yeas strains seems of special importance for flavor improvement of fermented beverages by glycerol optimization D Springer 1592 4 Food Sei Technol (Devers 2015) $2(12) 7588-7598 The main difference in glycerol productivity was observed in non-proliferating growth phase of yeast whereat over 70 % glycerol was produced (Du et al, 2011; Kajiwara et al. 2000; Kukee et al. 2003; Michnick et a. 1997; Remize et al. 2000), In the exponential growth phase, yeast possessed the highest GPDH activity and the highest level of GPDI expression but not glycerol yield (Du et al. 2011; Remize et al. 2000). In ‘comparison, the ethanol yield, ADH activity and expression level of ADHI all achieved peaks in the middle fermentation phase (Du et al. 2011). The significance of non-proliferating phase in glycetol production are also confirmed by the fact ‘that sugars are mainly consumed in this stage during beverage fermentations. Besides, glycerol was suspected to be mainly synthesized during the early phase of fermentation because of | the osmotic stress but released inthe later stage due to cellular lysis and higher membrane permeability (Orlic etal, 2010). Higher intial cell density is slightly in favor of glycerol formation for beer andl wine yeasts depending on the strains (GohthofTand Piendl 1973; Radler and Sehiitz 1982; Verbelen et al. 2009; Yalgin and Gzbas 2006). Ultra-high cell pitching ‘numbers led to faster glycerol formation and higher end eon centtation but no obvious influences was observed under the practical inoculum range (GehlhofT and Piendl 1973; Radler and Schiit, 1982), However, maximum glycerol concentration (8.6 pL) was also obtained at the low inoculum of the wine ‘yeast when varying the inoculum size between volume ratio of 2.5 to 12.5 % (Yalgin and Ozbas 2006). For glycerol optim zation, the improper inoculation strongly disturbed ferments- tion kinetics and further led to imbalanced flavor formation (Radler and Schiitz 1982; Verbelen et al. 2009; Yalgin and Gzbas 2006). Consequently, the cell pitching rate should be ‘optimized together with other parameters such as sugar, nitro ‘gen and oxygen in the processing of fermented beverages. Temperature shocks stimulate excessive glycerol production ‘Cold and heat shocks triggered glycerol accumulation by ac- tivating the HOG pathway via different branches of ‘osmosensors as shown in Fig. 1. The down shift of temper ture into 0-12 °C for 4-24 h caused the accumulation of intracellular glycerol, trehalose and heat shock proteins to protect the cells from freeze injury (Panadero et al. 2006), ‘The cold shock has been investigated with great interest for baker's yeasts in coping with the frozen dough technology. However, rare attention has been put in the beverage fermen- tation process conceming the risk of stuck and sluggish fer- ‘mentation, The pre-adaplation of the active dry wine yeast strains at low temperature (13 °C) was proved to increase ‘lycerol formation together with improved fermentation per- formance in terms of shortening fermentation period, effective uptake of ammonium and clear reduction of acotaldchyde, acetic acid and fusel aleohols but again the benefits were de- pend on strains (Csutords et al. 2014; Llaurado et al. 2005), Moreover, some strains, which hybridized from S, cerevisiae ‘and the cryotolerant species S, eubayanus, S. wvarum as well 8S, kudriayzevii, adapt effectively to the cold temperature in beet, wine and cider fermentations. Those strains may have better chance to overproduce glycerol in respond to the cold shock ( Arroyo-L6pez et al. 2010; Gibson et al. 2013). ‘The heat shocks have been applied to beverage fermenta- tion and up to two folds more glycerol has been produced by the brewing, wine, sake, shochu and whisky yeasts (Berlot and Berovit 2011; Berovie et al. 2007; Kajiwara et al. 2000; ‘Omori et al. 1996; Omori et al. 1997). The heat shocks often perform during cell preculture provess at the temperature of 30-50 °C for 20-120 min one or multiple times. In addition to the hyperosmotic stress induced glycerol accumulation, the hieat shock also initiated the changes in redox status where slycerol oversynthesis was responded to resume the redox homeostasis (Berlat and Berovig 2011; Kukee et al. 2003), Both glycerol production and GPDH activity were strongly bbut transiently induced by heat shock treatment, The prolonged time of intensive transcription of GPD1 was re- sponsible for the excessive glycerol production of shock yeast (Kajiwara et al, 2000), Moreover, the heat exposure tended to create the short memory effect in the yeast and it ‘was used to enhance glycerol production by repeating the ‘moderate heat shocks (Berlat and Berovig 2011; Berovié etal. 2007; Kukee etal, 2003), Excopt forthe impact on glycerol biosynthesis, heat shock ‘reatment caused a transient delay in growth but followed by ‘more biomass produetion and even complete sugar consusny tion after cell recovery (Berlot and Berovié 2011; Berovié et al, 2007; Kajiwara et al. 2000; Petropoulos et al. 2010), 'No consistent changes oa ethanol and acceptable level of var- iations on organic acids were found after heat shock trea ‘ments (Berlot and Berovié 2011; Berovit etal. 2007; Omori ct al, 1996; Petropoulos et al. 2010). While the yields of isoamyl acetate and the fatty acid ethyl esters were significant increase in the heat shock mutants overproducing glycerol (Omori tal. 1997). Hence, heat shock was claimed as an effective treatment for increasing glycerol without negative impacts on beverage flavors (Berlot and Berovié 2011; Omori et al. 1996). Further improvement can be made by carefully choosing the proper strains since the heat shock re sponse is also strongly strain-dependent (Omori etal. 1996; Petropoulos et al. 2010). The heat shock temperature, duration and interval should also be optimized. Environmental parameters regulate glycerol yield Ligh temperature isin favor of glycerol synthesis High fermentation temperature is in favor of glycerol synthe sis of S. cerevisiae ( Arroyo-Lopez et al. 2010; Scanes eta 1998; Yalgin and Ozbag 2008). 1.2-1.7 folds increase of 4 Food Si Technol (Desember 2015) $2(12): 7548-1598 1593 lycerol were observed by optimizing fermentation tempera tures for various beverage yeasts (Chen and Xu 2010; Ductal. 2011; Yalgin and Ozbas 2008). The enhanced level of GPDI expression and GPDH activity at higher temperature was re- sponsible for such inereases ( Arroyo-Lépez et el. 2010; Du etal. 2011). Generally, glycerol presents at higher level in ale beer and red wine than that in lager beer and white wine. This is probably due to thatthe former beverages are fermented at higher temperature regardless other variables. Except for the inerease in glycerol yield, higher temperature also resulted in obvious inerease in fermentation rate and decrease of yeast viability. Meanwhile it imputed negative impacts for the overall flavor developments (Csutoris et al 2014; Liu et al. 2014). Acetic acid and succinic acid were found increase with temperature (Du et al. 20115, Liu et al, 2014; Llaurado et al, 2005), Controversially, ethanol was higher at low temperature but not ADH activity and ADIT expression (Du etal, 2011), Besides, glye- rol production can also benefit from complete sugar con sumption when fermented at relative low temperature. The fermentation temperature should be optimized with great care due to the impacts on either flavor development or the risk of | sluggish fermentation. Sugar concentration is positively related to glycerol yield Within the tolerance range of osmotic stress, more glycerol ‘were produced with the inereasing of sugar concentration ( Amroyo-Lopez et al. 2010; Erasmus et al, 2004; Jiménez Marti ot al. 2011; Orlic et al. 2010; Piddocke et al. 2009; Radler and Schiitz 1982). As illustrated in Fig, 1, the HOG signaling cascade is activated when the cells sense the hyperosmotic stress environment. On one hand, the Hog! ki- nase appear to stimulate the glycolytic flux and increase the transcription of the genes involved in glycerol synthesis (GPD1 and GPP2) and glycerol import (STL.1); On the other hhand, it may also participate in the closure the channel aquaglyceroporin (FPS1) to prevent glycerol efflux (Hohmann et al. 2007). Notice that, under wine fermentation conditions, the HOG pathway was reported to only parlly control glycerol formation, Glycerol yield was mainly medi tated by the GPDH activity and the expression level of GPDI (Pigcau and Inglis 2005; Remize ct al, 2003). Moreover, the channel protein Fps1p seemed play a critical role forthe over- all glycerol yield (Remize etal. 2001) ‘The sugar composition also impact glycerol yield (Myers et al. 1997; Ortic etal. 2010; Piddocke et al. 2009). Higher proportion of fermentable sugar enhanced glycerol formation (Myers eta, 1997), Glycerol was higher in the fermentation of alucose rich medium than in medium with maltose addition (Orlic otal, 2010; Piddocke etal. 2009). The reason could be that more DAP and NADH were generated in high mono saccharide condition. The drawbacks of the high sugar fermentation lies at that it often leads to lower growth rate, longer lag phase, incomplete sugar utilization and strongly accumulations of acetaldehyde and acetate esters ( Jiméne Mant et al. 2011; Piddocke et al. 2009; Yu et al. 2012). More than two folds increase in ethyl acetate and isoamyl acetate was accumulated and the disproportional changes of | flavor-active metabolites have resulted in the over-fruity char- acter of beer (Piddocke et al, 2009), Interestingly, yeast via- bility in the high sugar fermentation process was improved by the pre-adaptation in osmotic stress media ( Jiménez-Marti etal. 2011), The pre-adaptations of yeast have great potential to enhance glycerol production in the main process of bever- age fermentations. Effecs of other fermentation parameters ‘The quantity and variety of the nitrogen source have a pro- nounced effect on yeast fermentation performance and flavor development in beverage fermentations. However, the nitto- ‘gen concentration seemed only have little influences on glye~ cero] formation (Orie etal. 2010; Remize etal. 2000). Some amino acids may be of special importance for glycerol sya- thesis considering that the omission of thiamine inthe medium lowered one third of the overall glycerol yield (Radler and Schitz 1982). The ammonium grown culture led to double ‘amounts of glycerol than the cultures on amino acids or the ‘mixtures as nitrogen source. This was probably due to the swplus NADH generated in de novo synthesis of amino acids (Albers etal. 1996; Michnick etal, 1997; Remize etal. 2000; Scanes et al. 1998). Ethanol yield, on the other hand, was found higher in amino acids media but was strain-dependent (Albers et al. 1996), For the impacts of agitation, two controversy results have been obtained on glycerol yield (Bjorkqvist et al, 1997; Gardner et al. 1993; Petropoulos et al. 2010; Radler and Schitz 1982; Remize et al, 2000). Glycerol is dominantly formed under anaerobie conditions to consume the excessive NADH during yeast alcoholic fermentation, Nevertheless, considerable amount of glycerol ean be also accumulated un- der the osmotic stress condition. Under such condition, GPDJ and GPP? tend to be overexpressed out ofthe need of osmo- regulation (Albertyn et al. 1994). The difference in the tran- scription level of GPD/ might be in charge of the controver sal impacts of agitation on glycerol yield in previous studies (Ansell and Adler 1999; Bjorkqvist etal. 1997), ‘The high pH-value has slightly positive effect on glycerol formation in beverage fermentation process ( Arroyo-Lépez ct al. 2010; Nykiinen and Suomaleinen 1983; Scanes et al 1998; Yalgin and Ozbas 2008). The positive effect was prob- ably because of the increased activity of aldehyde dehydroge- nase at higher pH-value. Aldehyde dehydrogenase catalyze the oxidation of acetaldehyde into acetic acid. This step gen- crates extra NADH which requite reoxidization via glycerol D Springer 1594 4 Food Sei Technol (Devers 2015) $2(12) 7588-7598 synthesis in the absence of oxygen (Fig. 1) (Wang etal. 2001), However, for beverage fermentation, low pH-value is often adopted for its inhibition on the growth of spoilage mieroor- anisms and enhancement on flavor development (Briggs cet al. 2004; Jackson 2008). ‘Small addition of sulphur dioxide (SO,) in the medium resulted in elevated glycerol yield (Gardner et al. 1993; Nykinen and Suomaleinen 1983; Radler and Schitz 1982; Scanes et al. 1998). SO, is a natural fermentation byproduct presented in beer and wine. It can be also added to beer (up to 20 mg/L) to protect oxidative flavor deterioration and to must (50-100 mg/L) to inhibit wild yeast and spoilage bacteria Briggs et al, 2004; Jackson 2008). The positive effect of SO; is based on that the bisulfites ion binds the acctaldchyde which results in ceumulation of cytosolic NADH and further shifts the carbon flow towards glycerol (Wang et al. 2001), Disputed glycerol yield was reported when varying SOs addition. Additionally, this method seems not applicable anymore for the trend to limit $02 addition in beverage industry, Sulfate at alkaline pH, sodium chloride (NaC), potassium chloride (KCD) and sodium carbonate (NaCOs) have been exploited as the selective agent (stress) for adaptive evolution tocreate high glycerol mutants (Kutyna et al. 2010). However, those mutants are often non-stable in addition to the acetic acid overproduction and prolonged fermentation time. On ‘grape must, S. cerevisiae yeast produced more glycerol com- pare with that on the synthetic medium (Du et al. 2011; Erasmus t al 2004; Radler and Schitz 1982). This indicates thatthe nutrients complexity may also influence glycerol bio- synthesis in beverage fermentation (Ansell and Adler 1999), In Fig, 2, we depict the yield improvement of glycerol by various optimizations in the process of beverage fermentation. To summarize, yeast glycerol production is influenced by ‘many growth, process and environmental factors but primarily it is suain-dependent. By she carefully choosing the yeast strains and further optimizing the fermentation conditions, beverage flavor could be improved by elevating glycerol yield. ‘Overproducing glycerol by genetically modified yeast Promote glycerol biosynthesis by overexpressing GPD and GPP genes ‘The two homologous genes GPDI and GPD2 are eracial for alycerol synthesis. The single deletions of GPDI or GPD2 only slightly reduced glycerol formation. While the god/A. and gpd2A double mutant did not produce detectable glycerol and was highly osmosensitive meanwhile failed to grow under anoxic conditions (Ansell et al. 1997; Bjorkgvist et al. 1997), In comparison, mutants lacking of GPP/ and GPP? genes ‘were devoid of glycerol-3-phosphatase activity but stil pro- duced a small amount of glycerol. Moreover, the two GPP ‘genes substituted well with each other for thatthe single mu- tant remained unaffected by the high osmolality stress (Pahlman et a. 2001). Glycerol overproduction has been realized by the adoption of yeasts overexpressing GPD and GPP genes for beverage fermentations (Cambon et al. 2006; Eglinton et al, 2002; Lopes et al. 2000; Nevoigt et al. 2002; Remize et al, 2003; Remize et al. 1999; Varela et al. 2012). Depending on the strains and fermentation environments, the GPDT overexpres- sion has yielded up to 6 folds mote glycerol compared with 2.6 times increase of the GPD2 overexpression (Eglinton et al, 2002; Varela et al. 2012). The single overexpression of GPP ‘genes did not significant changed glycerol yield whereat only 1.2 times inerease was observed (Pahiman et al, 2001). Mimic the beer fermentation conditions, the GPD/ overexpressing lager yeast yielded 8.73 g/L glycerol compared with 1.57 g/ L of the reference strain (Nevoigt etal. 2002). For wine fer- ‘mentation, the GPDJ mutant produced 34.3 g/L glycerol that ‘was 6 folds more compared with that of the wild type (Varela et al, 2012), To a less extent, 1.5 t0 3.8 folds increase in slycerol yield has boen lifted up by the overexpression of, GPDI in various commercial strains (Table 1) (Cambon etal, 2006; Remize etal. 1999), In comparison, the lab strains (uch as V5, W303-1A and OL1) overexpressing GPDI seemed to respond stronger in glycerol synthesis even some of them were unable to complete the sugar consumption (Cambon et al. 2006; Cordier et al. 2007; Lopes et al. 2000; Nevoigt and Stahl 1996; Remize et al, 1999). The overexpres- sion of GPD2, GPP/ and GPP2 are less predominant for ‘lycerol overproduction but have similar side effeets under beverage fermentation conditions (Pahlman et al. 2001; Remize et al. 2001). The engineered strains produced lower biomass and decreased ethanol formation (Nevoigt et al 2002). Minor changes in higher alcohols, estes and fatty acids were reported. However, strongly accumulation of aceta succinate, acetaldehyde, acetoin and 2,3-butanediol were ob- served but also strain-dependent (Table |) (Cambon et al 2006; Cordier etal. 2007; Nevoigt et al. 2002; Nevoigt and Stahl 1996; Remize et al. 1999), The effects for other organic acids such as male, citric, tartaric and lactic acids were not ‘obvious (Lopes et al. 2000; Remize et al. 2001). The “engineered beer” was described as shemy-like due to the imbalance flavor development. Majority of the sen- sory panel members were able to distinguish the “engineered wine” but did not fond of it (Lopes et al. 2000; Nevoigt et al. 2002) ‘To combat the over accumulation of acetic acid and 2,3- butanediol, ALDS encoding aldehyde dehydrogenase and BDHI encoding the 2,3-butanediol dehydrogenase (which 4 Food Si Technol (Desember 2015) $2(12): 7548-1598 oo Fig. 2 Glycerol yield improvement by the optimizations of various parameters during beverage fermentations The results were reported under beet (Piddocke et al. 2009; Vetelen et al, 2008), wine (Amoyo- Liner etal. 2010 Belt and Berovi 201: Berove etal, 2007; Castors tal 2014; Du etal, 2011; Erasmus etal. 2004; Kukee et al 2003; Orie catalyze the conversion of rancid acetoin to the innocuous 2,3- butanediol) were modified in the GPDI/2 overexpression mu- tants (Cambon etal, 2006; Condier etal, 2007; Eglinton etal 2002; Ehsani et al. 2009; Varela et al, 2012). The multiple ‘enginccred strains performed similarly in glycerol and ethanol formation compared with GPDI mutants (Table 1). The GPDI1 ald6A mutants showed lower level of acetate, but the increase of acetaldehyde, diacetyl and acetoin yield were even stronger. The accumulations of acetoin and diacetyl were fur- ther decreased in the GPD/ ald6A BDH! matant (Cambon et al. 2006; Eglinton et al. 2002; Ehsani etal, 2009; Varela etal. 2012), Direct carbon flux towards glycerol by other genetic ‘modifications ‘The indirect genetic modifications have been also performed to increase glycerol production by deletion of TPII, which en- codes triose phosphate isomerase, deletion of PDC2, regulatory gene involved in the up-regulation of transcription factor of Wet gravy Super compotion: Muogen cancetaton ‘San; tivogen camcentaton Heat sock Anerbie (Detsinsrverant cera sd ld tal. 2010; Petropouls etal, 2010; Remize etal, 2000; Yagan and aba 2008) and rice wine (Chea and Xu 2010) femeaation condone. Some ‘ofthe resuks are estimated fom the published figures if not provided Sirctly PDCIIS encoding pyruvate decarboxylase and deletion of ADHI in S. cerevisiae yeasts (Cordier et al. 2007; Nevoigt and Stahl 1996; Varela et al, 2012), Other genetic modification strategies such as increasing the cytosolic NADH proportion by the deletion of GUT? and NDEUi2 (involving in mitochondrial ‘oxidation of eytosolie NADH) and “opening” the glycerol ef: flux facilitator by truncation of Epsip were also applied to promote glycerol overproduction (Cordier et al. 2007; ‘Overkamp etal. 2000; Remize etal. 2001; Varela et al. 2012), Although these modifications resulted in dramatic nerease in tlycerol yield even up to 18 times mote in some synthetic ‘medium. ‘The mutants showed significantly impaired growth ability and accumulated large amounts of pyruvate, acetate and acetaldehyde because of the diverted carbon flow (Cordier et al. 2007; Nevoigt and Stabl 1996; Varela et al 2012). The yields of esters and higher aleohols were slighlly decreased of remained unchanged. Even no negative sensory cffects was claimed for wine quality, the prolonged fenments- tion time and poor social acceptance still hinder the application of those genetically modified yeasts in beverage industry D springer 7596 4 Food Sei Technol (Devers 2015) $2(12) 7588-7598 999) (Cambon etal. 2006; Kglinton et al. 2002; Ehsani etal. 20095, Varela etal. 2012), round brackets the brackets are the yields ofthe mutants: GPDI, BDHT overexpression and ald deletion; Ferment Conclusion 5 Glycerol yield has been optimized intensively for improving beverage flavor and ethanol reduction by the strategies such as screening of yeast strains, optimization of fermentation parame- ters and yeast genetic modifications. However, in both the sen- sory rescarch and fermentation optimization of glycerol, the fol- Towing aspects deserve more attention, Firstly, obvious gap exists between the actual glycerol level and the sensory beneficial con- ‘centration, Whereas, the beneficial concentration (range) remains unclear for beer as well as other fermented beverage. Additionally, glycerol was suspected to contribute better for the rnon- and low-aleoholic beverage considering the reported organ- ‘oleptic impacts and it needs further confirmation. Secondly, yeast slycerol production is primarily strain-dependent and yeast strains have been screened for their glycerol productivity. Nevertheless, the sereening was restricted within dozens of strains which seemed inadequate to get favorable yeast for Savor ‘improvement, Besides, fermentation parameters have been opti= ‘ized to improve glycerol yield. However, the variables and effects have been only partly considered, In the process of fermented beverages, the change of single parameter would in- cevilably affect the whole fermentation kinetics and further impact ‘on glycerol yield and the overall flavor developments. Therefore, ‘comprehensive optimization with the multiple inputs (stains, ‘environmental and process parameters) and multiple outputs (elycerol, important flavor compounds and beverage sensory Jmpression) should be investigated in the upcoming rescarch, “To facilitate the screening of large numberof strains and optimni= zation of multiple variables, we propose a platform constructed by the high throughput screening (HTTS) technology in combina- tion with the design of experiments (DoE) methods. The cell- based HTS often adopt the microtiter plates to miniaturize the fermentation and has been widely applied in the sereening of al 2001; Remize et al (Camino eta, 2006; Ehsan al 2009; Varela etal. 2012) Ref (Newoigt (Mick etal. 1997; Remi etal 2012) ia] 604150 076 +034) 23-Butanedol 2274248, (32 +059) (46L051) ust (Ind) o boratory (Lab) stains; the da 3340.26 0014001) Acstoin 5314103 (6.03 +006) 465.4 3.08, (om +001) iat} 06 (002 +001) 022 ro s (Ref) on ete ‘Acctaldehyde 023.007 (a3 +090) 0.14 + 0.05, (002 £001) eal te] 023 roy 7228 36, 2 ares Seecinate fet) 027 (0.06) 5 =o) 04601 (0302) (04402) 05402 r 94 1243.6) (07 + 14) s992 124 (65 29 c 2 tion of DoE methods could farther enhance the effectiveness for _lycctol optimization inthe processing of fermented beverages 4466 as 73420 S41 os+19 6 a7 Acknowledgments The frst author e funded by China Scholarship Council (Lab sting) a3 as Boer (Lager sti) Wine Wine g 2 References Albers E, Larsson C, Lidén G, Niklasson C, Gustafsson L (1996) Influence ofthe nitrogen source on Saccharomyces cerevisiae an scrobi growth and product formation. 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