Flavour formation by lactic acid bacteria and biochemical flavour profiling of cheese products.

Flavour development in dairy fermentations, most notably cheeses, results from a series of (bio)chemical processes in which the starter cultures provide the enzymes. Particularly the enzymatic degradation of proteins (caseins) leads to the formation of key-flavour components, which contribute to the sensory perception of dairy products. More specifically, caseins are degraded into peptides and amino acids and the latter are major precursors for volatile aroma compounds. In particular, the conversion of methionine, the aromatic and the branched-chain amino acids are crucial. A lot of research has focused on the degradation of caseins into peptides and free amino acids, and more recently, enzymes involved in the conversion of amino acids were identified. Most data are generated on Lactococcus lactis, which is the predominant organism in starter cultures used for cheese-making, but also Lactobacillus, Streptococcus, Propionibacterium and species used for surface ripening of cheeses are characterised in their flavour-forming capacity. In this paper, various enzymes and pathways involved in flavour formation will be highlighted and the impact of these findings for the development of industrial starter cultures will be discussed.

[1]  S. Skeie,et al.  Influence from raw milk flora on cheese ripening studied by different treatments of milk to model cheese. , 2000 .

[2]  J. Steele,et al.  Peptidases and amino acid catabolism in lactic acid bacteria , 2004, Antonie van Leeuwenhoek.

[3]  P. Schieberle,et al.  Formation of aroma-active strecker-aldehydes by a direct oxidative degradation of Amadori compounds. , 2000, Journal of agricultural and food chemistry.

[4]  L. Moio,et al.  Powerful odorants in water buffalo and bovine Mozzarella cheese by use of extract dilution sniffing analysis , 1993 .

[5]  M. Yvon,et al.  CodY-Regulated Aminotransferases AraT and BcaT Play a Major Role in the Growth of Lactococcus lactis in Milk by Regulating the Intracellular Pool of Amino Acids , 2003, Applied and Environmental Microbiology.

[6]  B. Smit Formation of Amino Acid Derived Cheese Flavour Compounds , 2004 .

[7]  G. de Roo,et al.  Purification and Characterization of Cystathionine (gamma)-Lyase from Lactococcus lactis subsp. cremoris SK11: Possible Role in Flavor Compound Formation during Cheese Maturation , 1997, Applied and environmental microbiology.

[8]  G. Urbach Relations between cheese flavour and chemical composition , 1993 .

[9]  I. Blank Gas Chromatography – Olfactometry in Food Aroma Analysis , 2001 .

[10]  M. Boekel,et al.  A review of Maillard reaction in food and implications to kinetic modelling , 2000 .

[11]  Werner Grosch,et al.  Evaluation of Key Odorants of the Neutral Volatiles of Emmentaler Cheese by the Calculation of Odour Activity Values , 1994 .

[12]  F. Archibald,et al.  Manganese: its acquisition by and function in the lactic acid bacteria. , 1986, Critical reviews in microbiology.

[13]  W. Engels Volatile and non-volatile compounds in ripened cheese: their formation and their contribution to flavour. , 1997 .

[14]  G. Reineccius,et al.  Identification and quantification of potent odorants in regular-fat and low-fat mild Cheddar cheese , 1997 .

[15]  J. Bosset,et al.  Comparison of the volatile flavour compounds of six european ‘AOC’ cheeses by using a new dynamic headspace GC-MS method , 1993 .

[16]  J. Accolas,et al.  Metabolism and biochemical characteristics of yogurt bacteria. A review [Streptococcus salivarius subsp. thermophilus, Lactobacillus delbrueckii subsp. bulgaricus] , 1992 .

[17]  V. Yaylayan,et al.  Carbohydrate and amino acid degradation pathways in L-methionine/D-[13C] glucose model systems. , 2001, Journal of agricultural and food chemistry.

[18]  R. Raya,et al.  Acetaldehyde metabolism in lactic acid bacteria , 1986 .

[19]  M. Yvon,et al.  Cheese flavour formation by amino acid catabolism , 2001 .

[20]  T. Morishita,et al.  Nutritional Requirements in Multiple Auxotrophic Lactic Acid Bacteria: Genetic Lesions Affecting Amino Acid Biosynthetic Pathways in Lactococcus lactis, Enterococcus faecium, and Pediococcus acidilactici. , 1992, Bioscience, biotechnology, and biochemistry.

[21]  M. Yvon,et al.  An aminotransferase from Lactococcus lactis initiates conversion of amino acids to cheese flavor compounds , 1997, Applied and environmental microbiology.

[22]  W. Hammes,et al.  Effect of enhanced proteolysis on formation of biogenic amines by lactobacilli during Gouda cheese ripening. , 1998, International journal of food microbiology.

[23]  A. Palva,et al.  Genetic characterization of an oligopeptide transport system from Lactobacillus delbrueckii subsp. bulgaricus , 2002, Archives of Microbiology.

[24]  W. Bockelmann,et al.  The surface flora of bacterial smear-ripened cheeses from cow's and goat's milk , 2001 .

[25]  B. E. Davidson,et al.  Identification and characterization of a cystathionine beta/gamma-lyase from Lactococcus lactis ssp. cremoris MG1363. , 2000, FEMS microbiology letters.

[26]  G. Smit,et al.  SCREENING FOR AND CONTROL OF DEBITTERING PROPERTIES OF CHEESE CULTURES , 1996 .

[27]  P. Bonnarme,et al.  Identification and Functional Analysis of the Gene Encoding Methionine- (cid:2) -Lyase in Brevibacterium linens , 2022 .

[28]  P. Renault,et al.  Pleiotropic transcriptional repressor CodY senses the intracellular pool of branched‐chain amino acids in Lactococcus lactis , 2001, Molecular microbiology.

[29]  A. Driessen,et al.  Bioenergetics and solute transport in lactococci. , 1989, Critical reviews in microbiology.

[30]  G. Smit,et al.  Diversity of l-leucine catabolism in various microorganisms involved in dairy fermentations, and identification of the rate-controlling step in the formation of the potent flavour component 3-methylbutanal , 2004, Applied Microbiology and Biotechnology.

[31]  J. Bosset,et al.  Key odorants in various cheese types as determined by gas chromatography-olfactometry , 2002 .

[32]  A. Chopin Organization and regulation of genes for amino acid biosynthesis in lactic acid bacteria. , 1993, FEMS microbiology reviews.

[33]  J. Gripon,et al.  Production of phenyl ethyl alcohol and its esters during ripening of traditional camembert , 1988 .

[34]  S. Skeie,et al.  Cooperation between Lactococcus lactis and Nonstarter Lactobacilli in the Formation of Cheese Aroma from Amino Acids , 2003, Applied and Environmental Microbiology.

[35]  J. Holbrook,et al.  NAD(+)-dependent D-2-hydroxyisocaproate dehydrogenase of Lactobacillus delbrueckii subsp. bulgaricus. Gene cloning and enzyme characterization. , 1994, European journal of biochemistry.

[36]  F. Ehrlich Über die Bedingungen der Fuselölbildung und über ihren Zusammenhang mit dem Eiweißaufbau der Hefe , 1907 .

[37]  A. Thierry,et al.  Conversion of l-Leucine to Isovaleric Acid by Propionibacterium freudenreichii TL 34 and ITGP23 , 2002, Applied and Environmental Microbiology.

[38]  Peter Ruhdal Jensen,et al.  Minimal Requirements for Exponential Growth of Lactococcus lactis , 1993, Applied and environmental microbiology.

[39]  T. Parliment,et al.  Volatile components of Limburger cheese , 1982 .

[40]  F. Kosikowski,et al.  Origins and principles , 1997 .

[41]  Patrick F. Fox,et al.  Cheese: Chemistry, Physics and Microbiology , 1993, Springer US.

[42]  S. Ehrlich,et al.  The complete genome sequence of the lactic acid bacterium Lactococcus lactis ssp. lactis IL1403. , 2001, Genome research.

[43]  A. Hagting,et al.  The proteolytic systems of lactic acid bacteria. , 1996, Antonie van Leeuwenhoek.

[44]  R. Axel,et al.  A novel multigene family may encode odorant receptors: A molecular basis for odor recognition , 1991, Cell.

[45]  M. Yvon,et al.  Genetic Characterization of the Major Lactococcal Aromatic Aminotransferase and Its Involvement in Conversion of Amino Acids to Aroma Compounds , 1999, Applied and Environmental Microbiology.

[46]  Glutamate dehydrogenase activity: a major criterion for the selection of flavour-producing lactic acid bacteria strains , 2002 .

[47]  M. E. Aly,et al.  Effect of ripening temperatures on proteolysis and lipolysis in the outer and inner regions of Ras-type cheese made by various salting methods , 1988 .

[48]  R. Simard,et al.  Bitter flavour in dairy products. II. A review of bitter peptides from caseins: their formation, isolation and identification structure masking and inhibition , 1992 .

[49]  M. Yvon,et al.  Expression of a Heterologous Glutamate Dehydrogenase Gene inLactococcus lactis Highly Improves the Conversion of Amino Acids to Aroma Compounds , 2000, Applied and Environmental Microbiology.

[50]  G. Smit,et al.  Population dynamics of lactococci from industrial, artisanal and non-dairy origins in defined strain starters for Gouda-type cheese , 2001 .

[51]  Edmund R. S. Kunji,et al.  Kinetics and specificity of peptide uptake by the oligopeptide transport system of Lactococcus lactis. , 1998, Biochemistry.

[52]  Nierop Groot MN,et al.  Conversion of phenylalanine to benzaldehyde initiated by an aminotransferase in lactobacillus plantarum , 1998, Applied and environmental microbiology.

[53]  M. Yvon,et al.  Enhancement of amino acid catabolism in Cheddar cheese using α-ketoglutarate: amino acid degradation in relation to volatile compounds and aroma character , 2001 .

[54]  L. M. Libbey,et al.  Identity of additional aroma constituents in milk cultures of Streptococcus Lactis var. Maltigenes. , 1966, Journal of Dairy Science.

[55]  P. Fox,et al.  Effect of adding free amino acids to Cheddar cheese curd on proteolysis, flavour and texture development , 1997 .

[56]  W. Grosch,et al.  Evaluation of potent odorants of Camembert cheese by dilution and concentration techniques , 1997 .

[57]  ScienceDirect FEMS microbiology reviews , 1993 .

[58]  S. Collin,et al.  Evidence of Strecker aldehyde excretion by yeast in cold contact fermentations. , 2000, Journal of agricultural and food chemistry.

[59]  H. Gruppen,et al.  Partial purification and characterization of two aminotransferases from Lactococcus lactis subsp. cremoris B78 involved in the catabolism of methionine and branched-chain amino acids , 2000 .

[60]  J. Villadsen,et al.  Metabolic Behavior of Lactococcus lactis MG1363 in Microaerobic Continuous Cultivation at a Low Dilution Rate , 2001, Applied and Environmental Microbiology.

[61]  G. Smit,et al.  Identification, Cloning, and Characterization of a Lactococcus lactis Branched-Chain α-Keto Acid Decarboxylase Involved in Flavor Formation , 2005, Applied and Environmental Microbiology.

[62]  A. Bolotin,et al.  Characterization and Role of the Branched-Chain Aminotransferase (BcaT) Isolated from Lactococcus lactissubsp. cremoris NCDO 763 , 2000, Applied and Environmental Microbiology.

[63]  Pieter Walstra,et al.  Dairy chemistry and physics , 1984 .

[64]  D. Glaser,et al.  Taste responses and thresholds obtained with the primary amino acids in humans , 1990 .

[65]  B. Weimer,et al.  Conversion of Methionine to Thiols by Lactococci, Lactobacilli, and Brevibacteria , 1998, Applied and Environmental Microbiology.

[66]  E. A. Day,et al.  Identification of compounds responsible for fruity flavor defect of experimental cheddar cheeses. , 1965, Journal of dairy science.

[67]  J. Steele,et al.  Use of 13C Nuclear Magnetic Resonance and Gas Chromatography To Examine Methionine Catabolism by Lactococci , 1998, Applied and Environmental Microbiology.

[68]  G. Cilento,et al.  Oxidation of phenylpyruvic acid. , 1987, Biochimica et biophysica acta.

[69]  W. Bockelmann,et al.  The microflora of Tilsit cheese. Part 1. Variability of the smear flora , 1997 .

[70]  M. E. Morgan,et al.  Decarboxylation of α-Keto Acids by Streptococcus lactis var. maltigenes , 1967 .

[71]  G. Urbach Contribution of lactic acid bacteria to flavour compound formation in dairy products , 1995 .

[72]  M. Yvon,et al.  Autolysis of Lactococcus lactis AM2 stimulates the formation of certain aroma compounds from amino acids in a cheese model , 2004 .

[73]  P. R. Elliker,et al.  The Nutritional Requirements of Lactic Streptococci Isolated from Starter Cultures. I. Growth in a Synthetic Medium , 1953 .

[74]  Weimer,et al.  Purification and characterization of L-methionine gamma-lyase from brevibacterium linens BL2 , 1998, Applied and environmental microbiology.

[75]  H. Joosten,et al.  Conditions allowing the formation of biogenic amines in cheese, 2: decarboxylative properties of some non-starter bacteria , 1987 .

[76]  R. Siezen,et al.  Cloning, Characterization, Controlled Overexpression, and Inactivation of the Major Tributyrin Esterase Gene of Lactococcus lactis , 2000, Applied and Environmental Microbiology.

[77]  V. Monnet,et al.  The EstA esterase is responsible for the main capacity of Lactococcus lactis to synthesize short chain fatty acid esters in vitro , 2002, Journal of applied microbiology.

[78]  H. Taguchi,et al.  Conversion of Lactobacillus pentosusd-Lactate Dehydrogenase to a d-Hydroxyisocaproate Dehydrogenase through a Single Amino Acid Replacement , 2003, Journal of bacteriology.

[79]  P. Walstra,et al.  Dutch-type varieties. , 1993 .

[80]  E. Hammond,et al.  Generation of Swiss Cheese Flavor Components by the Reaction of Amino Acids with Carbonyl Compounds , 1989 .

[81]  B. Ganesan,et al.  Role of Aminotransferase IlvE in Production of Branched-Chain Fatty Acids by Lactococcus lactis subsp. lactis , 2004, Applied and Environmental Microbiology.

[82]  P. McSweeney,et al.  Biochemical pathways for the production of flavour compounds in cheeses during ripening: A review , 2000 .

[83]  M. Kula,et al.  d-2-hydroxyisocaproate dehydrogenase from Lactobacillus casei , 2004, Applied Microbiology and Biotechnology.

[84]  Z. Libudzisz,et al.  Heterogeneity of the physiological activity of Lactococcus and Leuconostoc sp. strains , 1993 .

[85]  Francesco Addeo,et al.  Grana Padano cheese aroma , 1998, Journal of Dairy Research.

[86]  B. E. Davidson,et al.  Identification and characterization of a cystathionine β/γ-lyase from Lactococcus lactis ssp. cremoris MG1363 , 2000 .

[87]  F Addeo,et al.  Odour-impact compounds of Gorgonzola cheese. , 2000, The Journal of dairy research.

[88]  R. Lindsay,et al.  Evaluation of the Role of Microbial Strecker-Derived Aroma Compounds in Unclean-Type Flavors of Cheddar Cheese , 1985 .

[89]  M. Kleerebezem,et al.  Regulation of the metC-cysK Operon, Involved in Sulfur Metabolism in Lactococcus lactis , 2002, Journal of bacteriology.

[90]  M. Kleerebezem,et al.  Metabolic engineering of lactic acid bacteria for the improvement of fermented dairy products , 2000 .

[91]  G. Smit,et al.  Enhanced flavour formation by combination of selected lactococci from industrial and artisanal origin with focus on completion of a metabolic pathway , 2001, Journal of applied microbiology.

[92]  W. Grosch Evaluation of the key odorants of foods by dilution experiments, aroma models and omission. , 2001, Chemical senses.

[93]  G. Smit,et al.  Chemical conversion of alpha-keto acids in relation to flavor formation in fermented foods. , 2004, Journal of agricultural and food chemistry.

[94]  Catrienus de Jong,et al.  Flavour forming abilities and amino acid requirements of Lactococcus lactis strains isolated from artisanal and non-dairy origin , 1999 .

[95]  J. Broadbent,et al.  Tryptophan catabolism by Lactobacillus casei and Lactobacillus helveticus cheese flavor adjuncts , 1999 .

[96]  R. Raya,et al.  Threonine aldolase in Lactobacillus bulgaricus ATCC 11842 and YOP 12 , 1986 .

[97]  J. K. Rippe,et al.  Enzymic Generation of Methanethiol To Assist in the Flavor Development of Cheddar Cheese and Other Foods , 1986 .

[98]  Michael Rychlik,et al.  Flavour and off-flavour compounds of swiss Gruyère cheese. Evaluation of potent odorants , 2001 .

[99]  T. Kaneda,et al.  Biosynthesis of branched-chain fatty acids in Bacillus subtilis. A decarboxylase is essential for branched-chain fatty acid synthetase. , 1988, The Journal of biological chemistry.

[100]  V. Juillard,et al.  Diversity of Oligopeptide Transport Specificity in Lactococcus lactis Species , 2003, The Journal of Biological Chemistry.

[101]  M. Kleerebezem,et al.  Complete genome sequence of Lactobacillus plantarum WCFS1 , 2003, Proceedings of the National Academy of Sciences of the United States of America.

[102]  Werner Grosch,et al.  Detection of potent odorants in foods by aroma extract dilution analysis , 1993 .

[103]  L. M. Libbey,et al.  Lactobacillus maltaromicus, a New Species Producing a Malty Aroma1 , 1974 .

[104]  J. Steele,et al.  PURIFICATION AND CHARACTERIZATION OF OLIGOMERIC SPECIES OF AN AROMATIC AMINO ACID AMINOTRANSFERASE FROM LACTOCOCCUS LACTIS SUBSP. LACTIS S3 , 1998 .

[105]  G. Lees,et al.  Formation of acetaldehyde from threonine by lactic acid bacteria , 1976, Journal of Dairy Research.

[106]  J. Hugenholtz,et al.  Lysis of Lactococcus lactis subsp.cremoris SK110 and Its Nisin-Immune Transconjugant in Relation to Flavor Development in Cheese , 1998, Applied and Environmental Microbiology.

[107]  C. Peláez,et al.  Enhancement of 2-methylbutanal formation in cheese by using a fluorescently tagged Lacticin 3147 producing Lactococcus lactis strain. , 2004, International journal of food microbiology.

[108]  G. Smit,et al.  Application of wild starter cultures for flavour development in pilot plant cheese making , 2000 .

[109]  S. Skeie,et al.  The nature of aroma compounds produced in a cheese model by glutamate dehydrogenase positive Lactobacillus INF15D depends on its relative aminotransferase activities towards the different amino acids , 2004 .

[110]  Terry E. Acree,et al.  A Procedure for the Sensory Analysis of Gas Chromatographic Effluents , 1984 .

[111]  B. Law,et al.  Isolation and characterization of the L-methionine-γ-demethiolase from Brevibacterium linens NCDO 739 , 1989 .

[112]  M. Kula,et al.  d-(-)-Mandelic acid dehydrogenase from Lactobacillus curvatus , 1988, Applied Microbiology and Biotechnology.

[113]  M. Cocaign-Bousquet,et al.  Rational development of a simple synthetic medium for the sustained growth of Lactococcus lactis , 1995 .

[114]  A. Lepeuple,et al.  Involvement of a Prophage in the Lysis of Lactococcus lactis subsp. cremoris AM2 during Cheese Ripening , 1998 .

[115]  Bruce P. Bryant,et al.  Receptor mechanisms for flavour stimuli , 1994 .

[116]  Mark E. Johnson,et al.  Overexpression of Lactobacillus caseid-Hydroxyisocaproic Acid Dehydrogenase in Cheddar Cheese , 2004, Applied and Environmental Microbiology.

[117]  M. Gobbetti,et al.  Purification and characterization of cystathionine γ‐lyase from Lactobacillus fermentum DT41 , 1998 .

[118]  Kumiko Ninomiya Umami: a universal taste , 2002 .

[119]  S. Parayre,et al.  Enzymatic versus spontaneous S-methyl thioester synthesis in Geotrichum candidum. , 2000, FEMS microbiology letters.

[120]  B. Weimer,et al.  Purification and Characterization ofl-Methionine γ-Lyase from Brevibacterium linens BL2 , 1998, Applied and Environmental Microbiology.

[121]  Andrew M. Taylor,et al.  Flavour science : recent developments , 1996 .

[122]  G. Smit,et al.  Fermentative formation of flavour compounds by lactic acid bacteria , 2002 .

[123]  H. Spinnler,et al.  Review: Compounds Involved in the Flavor of Surface Mold-Ripened Cheeses: Origins and Properties , 1996 .

[124]  J. L. Quéré,et al.  Inactivation of lactococcal aromatic aminotransferase prevents the formation of floral aroma compounds from aromatic amino acids in semi-hard cheese , 1999 .

[125]  G. Smit,et al.  Development of a high throughput screening method to test flavour‐forming capabilities of anaerobic micro‐organisms , 2004, Journal of applied microbiology.

[126]  O. Kuipers,et al.  Molecular and Functional Analyses of themetC Gene of Lactococcus lactis, Encoding Cystathionine β-Lyase , 2000, Applied and Environmental Microbiology.

[127]  A Ott,et al.  Origin of acetaldehyde during milk fermentation using (13)C-labeled precursors. , 2000, Journal of agricultural and food chemistry.

[128]  C. Dacremont,et al.  Concept matching technique for assessing importance of volatile compounds for Cheddar cheese aroma , 1994 .

[129]  A. Alting,et al.  Purification and Characterization of Cystathionine (beta)-Lyase from Lactococcus lactis subsp. cremoris B78 and Its Possible Role in Flavor Development in Cheese , 1995, Applied and environmental microbiology.

[130]  V. Yaylayan Recent Advances in the Chemistry of Strecker Degradation and Amadori Rearrangement : Implications to Aroma and Color Formation , 2003 .

[131]  M. Yvon,et al.  Adding α-Ketoglutarate to Semi-hard Cheese Curd Highly Enhances the Conversion of Amino acids to Aroma Compounds , 1998 .

[132]  F. Kosikowski,et al.  Cheese and fermented milk foods. Volume 1: origins and principles. , 1997 .

[133]  G. Smit,et al.  Lactococcal aminotransferases AraT and BcaT are key enzymes for the formation of aroma compounds from amino acids in cheese , 2003 .

[134]  F. Exterkate,et al.  Bitter flavour in cheese. I: Mechanism of the formation of the bitter flavour defect in cheese , 1983 .

[135]  S. Visser,et al.  Isolation and comparative characterization of components that contribute to the flavour of different types of cheese , 1994 .

[136]  M. Gobbetti,et al.  Amino acid catabolism in cheese‐related bacteria: selection and study of the effects of pH, temperature and NaCl by quadratic response surface methodology , 2001, Journal of applied microbiology.

[137]  P. Bonnarme,et al.  Catabolism of volatile sulfur compounds precursors by Brevibacterium linens and Geotrichum candidum, two microorganisms of the cheese ecosystem. , 2003, Journal of biotechnology.

[138]  J. Gripon,et al.  The chemistry of flavour and texture generation in cheese , 1982 .

[139]  G. Smit,et al.  Antimicrobial-producing wild lactococci isolated from artisanal and non-dairy origins , 2002 .

[140]  M. Yokoyama,et al.  Production of Skatole and para-Cresol by a Rumen Lactobacillus sp , 1981, Applied and environmental microbiology.

[141]  D. Hemme,et al.  Production of methanethiol from methionine by Brevibacterium linens CNRZ 918. , 1985, Journal of general microbiology.

[142]  Frank Kosikowski,et al.  Cheese and Fermented Milk Foods , 1982 .

[143]  T. Kaneda,et al.  Coenzyme A- and nicotinamide adenine dinucleotide-dependent branched chain alpha-keto acid dehydrogenase. I. Purification and properties of the enzyme from Bacillus subtilis. , 1969, The Journal of biological chemistry.

[144]  O. Kuipers,et al.  Probing Direct Interactions between CodY and the oppD Promoter of Lactococcus lactis , 2005, Journal of bacteriology.

[145]  M. Salaspuro,et al.  Acetaldehyde production and metabolism by human indigenous and probiotic Lactobacillus and Bifidobacterium strains. , 2000, Alcohol and alcoholism.

[146]  A. P. DE Ruiz Holgado,et al.  Acetaldehyde Production by Strains Used as Probiotics in Fermented Milk. , 1994, Journal of food protection.

[147]  C. Foucaud-Scheunemann,et al.  Identification and characterization of an oligopeptide transport system in Leuconostoc mesenteroides subsp. mesenteroides CNRZ 1463 , 2002, Letters in applied microbiology.

[148]  G. Reineccius,et al.  Identification of aroma compounds in Parmigiano-Reggiano cheese by gas chromatography/olfactometry. , 2002, Journal of dairy science.

[149]  Shaoquan Liu,et al.  Esters and their biosynthesis in fermented dairy products: a review , 2004 .

[150]  M. Kleerebezem,et al.  Metabolic Engineering of Acetaldehyde Production by Streptococcus thermophilus , 2002, Applied and Environmental Microbiology.

[151]  Mark E. Johnson,et al.  Aromatic amino acid catabolism by lactococci , 1997 .

[152]  G. Smit,et al.  Properties of mesophilic lactic acid bacteria from raw milk and naturally fermented raw milk products , 1996 .

[153]  G. Smit,et al.  Starter culture development for improving the flavour of Proosdij-type cheese , 2003 .

[154]  M. Kula,et al.  l-2-hydroxyisocaproate dehydrogenase—A new enzyme from Lactobacillus confusus for the stereospecific reduction of 2-ketocarboxylic acids , 1984, Applied Microbiology and Biotechnology.

[155]  B. Poolman,et al.  On the binding mechanism of the peptide receptor of the oligopeptide transport system of Lactococcus lactis , 2000, The EMBO journal.

[156]  J. Bosset,et al.  Flavour and off-flavour compounds of Swiss Gruyère cheese. Identification of key odorants by quantitative instrumental and sensory studies , 2001 .

[157]  Gary A Reineccius,et al.  Aroma extract dilution analysis of aged cheddar cheese , 1995 .

[158]  R. Jensen,et al.  Intracellular roles of microbial aminotransferases: overlap enzymes across different biochemical pathways. , 1981, Critical reviews in microbiology.

[159]  C. Peláez,et al.  Biochemical and molecular characterization of alpha-ketoisovalerate decarboxylase, an enzyme involved in the formation of aldehydes from amino acids by Lactococcus lactis. , 2004, FEMS microbiology letters.

[160]  M. Kleerebezem,et al.  Flavour formation from amino acids by lactic acid bacteria: predictions from genome sequence analysis , 2002 .

[161]  D. G. Laing,et al.  Flavour perception mechanisms , 1996 .

[162]  V. Marshall,et al.  Threonine aldolase and alcohol dehydrogenase activities in Lactobacillus bulgaricus and Lactobacillus acidophilus and their contribution to flavour production in fermented milks , 1983, Journal of Dairy Research.

[163]  H. Maeda,et al.  Enzymatic Synthesis of Optically Pure (R)-( — )-Mandelic Acid and Other 2-Hydroxycarboxylic Acids: Screening for the Enzyme, and Its Purification, Characterization and Use , 1986 .