AGRIEAST 2015 (10) P.

16 - 26
Review Article

Biochemistry of cheese ripening

M. Pagthinathan and M.S.M. Nafees

Department of Animal Science, Faculty of Agriculture, Eastern University,
Chenkalady, Sri Lanka
pagthymp@[Link]

ABSTRACT

Cheese ripening basically includes the breakdown of proteins, lipids and carbohydrates which
releases flavour compounds and modifies cheese texture. Principal ripening agents are milk
enzymes (plasmin and lipoprotein lipase), milk coagulant, starter lactic culture, secondary
culture and ripening agents. The ripening process of cheese is very complex and involves
microbiological and biochemical changes to the curd resulting in the flavour and texture
characteristics of the particular variety. Microbiological changes during ripening include the
death and lysis of starter cells, nonstarter lactic acid bacteria, and secondary microflora in
many varieties of cheese. Moulds in mould-ripened varieties and a complex Gram-positive
bacterial flora in smear cheeses are of great importance to the flavour and texture of cheese.
Cheese texture softens during ripening as a consequence of proteolysis of the casein micelle
and changes to the water-binding ability of the curd and in pH. The biochemical changes
occurring during ripening may be grouped into primary events that include the metabolism of
residual lactose, lactate and citrate (glycolysis), lipolysis and proteolysis. Following the primary
events, secondary biochemical events occur which are responsible for the development of
many volatile flavour compounds of ripened cheese varieties.

Keywords: Amino acid catabolism, cheese flavour, cheese ripening, metabolism of fatty acids,
metabolism of lactate, proteolysis
Introduction

The ripening of cheese is a very complex process make their choice of cheese primarily based
which involves microbiological and biochemical on flavour and texture characteristics (Awad,
changes leading to the development of flavor and 2006). Flavour development in cheese is of
texture characteristics of cheese. Microbiological major economic interest since final flavour of
ripening includes the death and auto-lysis of starter cheese determines consumer choice.
cells, the growth of an adventitious flora (non- Development of flavour in cheese is a complex
starter lactic acid bacteria) and secondary series of microbiological, biochemical and
microflora (McSweeney, 2004b). The biochemical chemical processes that occur during ripening
reactions occurring during ripening are classified in cheese (Yvon and Rijnen 2001; Walstra et
into primary and secondary biochemical events. al., 2006; Hannon et al., 2007). Flavour
Primary events are the metabolism of residual compounds are formed by various processes
lactose, lactate and citrate (glycolysis), lipolysis acting in a concerted and/or sequential manner
and proteolysis. Secondary biochemical reactions including conversion of lactose and citrate, fatty
are followed by primary events, which are very acids, and peptides and amino acids
important for the generation of many volatile flavor (McSweeney, 2004a; McSweeney, 2004b).
compounds in cheese (Fox et al., 2000; Lactic fermentation is responsible for the initial
McSweeney, 2004b). Biochemical processes of stage of flavour compounds formed by the
cheese ripening have previously been elucidated starter bacteria, and plays an important role in
by several researchers (McSweeney and Sousa, cheese ripening (Fox et al., 2000. The salt
2000; Sousa et al., 2001; Smit et al., 2002; content is also important to the flavour and
Katechaki et al., 2009). quality of cheese (Pastorino et al., 2003; Awad,
Flavour, texture and aroma are very important 2007). However, many studies have found that
features of cheese (Awad, 2006). Consumers most of the flavour components are developed

16
AGRIEAST 2015 (10) Pagthinathan and Nafees : Cheese ripening

during cheese ripening (Fox et al., 2000; drained through the whey. According to
McSweeney, 2004a; McSweeney, 2004b; Walstra McSweeney and Fox (2004), 0.2–0.5% of
et al., 2006). citrate is present in Cheddar cheese. In the
presence of mesophilic lactobacilli starter
Metabolism of residual lactose and citrate cultures, citrate can be metabolised to produce
certain flavour compounds, including diacetyl,
The primary glycolytic process is the metabolism acetoin, acetate and 2,3- butanediol, which
of lactose to lactate by the starter culture during contribute to the flavour of cheese (Cogan and
early stages of ripening. If glycolysis is incomplete Hill, 1993; de Figueroa et al., 2000,2001;
by the starter culture, the non-starter lactobacillis McSweeney, 2004b).
may take part in the process of glycolysis (Fox et
al., 2000). It has also been reported that the rate Degradation of lipids
and extent of lactose metabolism influence the
initial texture of the curd (McSweeney, 2004a). A Cheese lipids may involve hydrolytic or
large amount (96%) of lactose in milk is lost in the oxidative degradation by the action of
whey as lactose or lactate (Fox et al., 2000). indigenous, endogenous and/or exogenous
However, considerable amount of lactose remains lipases, which result in the liberation of free fatty
in fresh curd, which can significantly affect cheese acids in cheese during ripening, as shown in
quality (McSweeney, 2004b). In the early part of Figure 1. (Bosset et al., 1993; Fox et al., 1995;
ripening, residual lactose is quickly metabolised to McSweeney, 2004a; Walstra et al., 2006). The
form L-Lactate. The rate and extent of reaction milk fat is rich in short-chain fatty acids, which
are dependent on the temperature and salt-in- are important precursors for the production of
moisture (S/M) levels of the curd. When S/M levels volatile flavour compounds that considerably
increase, starter activity is slowly stopped at the contribute to the development of flavour in
end of manufacture (Fox et al., 2000; McSweeney, many cheese varieties (Collins et al., 2003;
2004a; Parente and Cogan, 2004). Alewijn et al., 2005). Lipolysis occurs in all
varieties of cheese, but excessive lipolysis can
The rate of lactic acid formation during cheese result in rancidity (Collins et al., 2003;
ripening has a direct effect on the taste of cheese, McSweeney, 2004b). Further, it has also been
especially in young cheese (Fox et al., 2000). D- reported that free fatty acids were counter-
Lactate could be produced directly from residual balanced with other flavour compounds to
lactose by starter lactobacilli or by racemisation develop an appreciated aroma (Bosset and
of L–lactate by non-starter lactic acid bacteria Gauch, 1993; Fox et al., 1995). Lipases
(NSLAB - eg: pediococci), during ripening of generally originate from the milk, rennet
cheese (Fox et al., 2000; McSweeney, 2004a). preparation, starter, adjunct starter, nonstarter,
The lactate racemisation of lactate generally and secondary microorganisms and exogenous
involves oxidation of L-lactate to form pyruvate lipase (Deeth, and Fitz-Gerald, 1995; Fox and
by L-lactate dehydrogenase, which is then Wallace, 1997; McSweeney and Sousa, 2000).
converted to D-lactate by D–lactate
dehydrogenase. DL-lactate is less soluble than L– Milk contains an indigenous lipase, which is
lactate, resulting in the formation of Ca-DL -lactate called as lipoprotein lipase (LPL) (Fox and
crystals which appear as white specks on the Stepaniak, 1993; Fox et al., 1993). The fat does
surface of the mature cheese (Dybing et al., 1998; not hydrolyse under normal circumstance
McSweeney, 2004b). While the crystals are because substrate (fat) and enzyme (LPL) are
harmless, they may reduce consumer’s interest compartmentalised by the milk fat globule
(McSweeney, 2004b). Lactate can also be membrane (MFGM) and most of LPL is
metabolised by lactic acid bacteria (LAB) to associated with the casein micelles (Collins et
produce flavour compounds including acetate, al., 2003; McSweeney, 2004b). The rupture of
ethanol, formate and CO2. However, the extent of MFGM due to homogenisation, agitation,
this process depends on the population of NSLAB foaming and freezing, could cause significant
and the presence of O2 (McSweeney, 2004b). lipolysis, resulting in off-flavors in cheese (Fox
Citrates have a relatively lower concentration in et al., 2000; Collins et al., 2003; McSweeney,
milk, most of which is in the soluble phase and is 2004b).

17
AGRIEAST 2015 (10) P. 16 - 26

LPL is preferable for hydrolysis of triglycerides production of volatile flavour compounds

containing short and medium-chain fatty acids through a series of metabolic processes during
(Deckelbaum et al., 1990; Fox et al., 2000), and ripening of cheese (McSweeney and Sousa,
acts preferentially C4:0 and C6:0 located at the sn-1 2000; Walstra et al., 2006). The enzymatic
and sn-3 positions of triglycerides (Olivecrona et system permits the β–oxidation of fatty acid to
al., 1992; Collins et al., 2003). LPL is probably a β-keto acid, which is rapidly de-carboxilated
heat liable and is largely inactivated by HTST to give methyl ketone (Figure 2) (McSweeney,
pasteurization, although 78 oC for 10 s is required 2004a; Dimitrellou et al., 2009). Collins et al.
for complete inactivation of this enzyme (Fox et (2003) reported that the methyl ketones are the
al., 2000; McSweeney, 2004b). Collins et al. most important flavour compound, depending
(2003) reported that the LPL activity is most on types of cheese and concentration of methyl
significant in raw milk cheeses than in cheese made ketone. The formation of methyl ketones is
from pasteurised milk. Moreover, rennet paste, influenced by several factors including
used as a coagulant in certain Italian cheese temperature, pH of cheese (pH 5-7),
varieties, contains a potent lipase and pregastric concentration of precursor FFA and
esterase, which are responsible for lipolysis in physiological state of the mould. Methyl ketones
those type cheeses (McSweeney, 2004b). The may also be reduced to form the corresponding
Lactic acid bacteria are also capable to lipolysis secondary alcohol by the action of P. roqueforti
of fat, and their enzymes have been shown to enzymes (Collins et al., 2003; Walstra et al.,
contribute to significant level of lipolysis 2006).
characteristics in Cheddar cheese (Collins et al.,
2003; McSweeney, 2004a, 2004b). The lactones in cheese are γ-decalactone and
δ-decalactone, which contribute to the flavour
Catabolism of free fatty acid of a number of cheese varieties (McSweeney,
2004a). These compounds are formed from
Fatty acids have a direct impact on the flavour of hydroxyacids following intramolecular
many cheese varieties, e.g. C4-C 10 acids are esterification (Collins et al., 2003).
particularly strongly flavoured (Collins et al., Hydroxyacid are produced from oxidation of
2003). Free fatty acids, especially short-medium fatty acids by the action of lipoxygenases and
chain fatty acids, are important precursors for the other enzymes present in cowmilk (Collins et al.,2003;

18
AGRIEAST 2015 (10) Pagthinathan and Nafees : Cheese ripening

Figure 2 : Fatty acid metabolism during ripening

(Source : McSweeney, 2004a)

McSweeney, 2004a). Dodecalactones may be which may cause an off-flavour to cheese

formed by P. roqueforti from long-chain of (McSweeney, 2004a). However, McSweeney
unsaturated fatty acids (Chalier and Crouzet, and Sousa (2000) and Yvon and Rijnen (2001)
1992). Collins et al. (2003) reported that indicated that the role of proteolysis in cheese
disproportionate amounts of high molecular mass flavour is a production of peptides and amino
lactones caused rancidity in Cheddar cheese. In acids which act as precursors for catabolic
addition, esters are formed by reaction of fatty reactions, resulting in production of many
acids with an alcohol, and the ethyl esters are most important volatile flavour compounds. Sousa et
common in cheese (McSweeney, 2004a). al. (2001) reported that in most cheeses, the
Thioester compounds are also produced when initial proteolysis of caseins occurred mainly by
fatty acid reacts with catabolic products of the coagulant, and to some extent, by plasmin
sulphur-containing amino acids (Molimard and and somatic cell proteinases (cathepsin D), and
Spinnler, 1996; Collins et al.,2003). release the large and intermediate-sized
peptides which are subsequently hydrolysed by
Degradation of protein the coagulant and microbial enzymes. Microbial
proteinases generally produce the small peptides
Proteolysis is the most complex, and important and amino acids in cheese probably, by
biochemical event in cheese ripening (Yvon and hydrolyzing the large peptides produced from
Rijnen, 2001; Dimitrellou et al., 2010). Proteolysis αs1- casein and β-casein by action of chymosin
plays a vital role in the development of texture as and plasmin respectively, whereas free amino
well as flavour (Sousa et al., 2001). Proteolysis is acids are produced from short peptides by
contributing to the softening of cheese texture microbial peptidase (Sousa et al., 2001;
during ripening due to the hydrolyzing para-casein McSweeney, 2004a,b).
matrix, decreasing the water activity (aw) through
water binding by liberated carboxylic acids, During cheese ripening, proteinases and
forming the amino groups by hydrolysis (Sousa et peptidases originate from the coagulant, milk,
al., 2001; McSweeney, 2004b). Proteolysis may starter LAB, NSLAB, secondary starters and
have a direct impact on cheese flavour through exogenous proteinases or peptidases added in
the formation of short peptides and amino acids, curd to accelerate the ripening (Fox et al., 2000).

19
AGRIEAST 2015 (10) P. 16 - 26

Plant proteases have also been investigated as tripeptidases (Christensen et al., 1999;
source of milk coagulants for cheese production. McSweeney, 2004b). It has been reported that
So far, only a few plant aspartic proteinases have proteinase (PrtP) contributes to the formation
been isolated and partially characterised (Tavaria of small peptides in cheese by hydrolysing
et al., 1997; Bruno et al., 2010). Most plant larger peptides produced from αs1 – casein and
coagulants were reported to have low ratio of β – casein by chymosin and plasmin,
milk-clotting to proteolytic activity, which results respectively, whereas intracellular peptidases
in bitter peptides in ripened cheese, and low yield such aminopeptidases, dipeptidases and
of cheese production (Sousa et al., 2001). tripeptidases liberate the free amino acids from
small peptides ( Fox et al., 2000; McSweeney,
2004b). In addition, non – starter LAB
The major source of milk coagulant is calf rennet, (NSLAB) is involved in the ripening of nearly
which contains two types of proteolytic enzymes all types of cheese, especially Cheddar cheese
naturally, chymosin (EC [Link]) (88-94%) and (Fox et al., 2000). The activity of the NSLAB
bovine pepsin (EC [Link]) (6–12%) (Sousa, et appears to supplemets the proteolytic action of
al., 2001). The principal and essential role of the the starter (McSweeney, 2004a). However,
coagulant in cheese making is to coagulate milk. NSLAB seems to be not only contributing
Some coagulant activity is retained in the curd to proteolysis in Cheddar cheese, but also to the
contribute to proteolytic activity during ripening in release of free amino acids (Fox and
many varieties of cheeses (Sousa et al., 2001). McSweeney, 1998).
Other enzymes found to be a source of coagulants
for cheese production include pepsin Catabolism of free amino acid
(Rhizomucor) and proteinases (Cryphonectria
parasitcia, Cynara cardunculus) (Fox et al., The catabolism of the free amino acids during
2000; McSweeney, 2004b; Nourani et al., 2009). ripening has produced many flavor compounds
(McSweeney and Sousa, 2000; Yvon and
Milk contains several indigenous proteinases. Rijnen, 2001). Products of amino acid
Plasmin is a principal indigenous proteinase, and catabolism contribute as precursors to the
is a trypsin-like serine proteinase. It acts optimally development of volatile flavour compounds in
at pH 7.5 and 37°C and heat stable up to 55oC. cheese (Katechaki et al., 2009).
The plasmin activity is more significant in cheese
types which are cooked to high temperature Amino acids are grouped into branched-chain
because high temperatures probably inactivate the amino acids, aromatic amino acids and
plasmin inhibitors (McSweeney, 2004a). The methionine. They act as a substrate for
plasmin is active on caseins in the order β-casein aminotransferase (lyase), decarboxylation,
H” αs2- casein > αs1-casein. κ-Casein seems to deamination, dehydrogenation, oxidation and
be quite resistant to the action of plasmin (Fox et reduction reactions to produce a variety of
al., 2000). In addition, milk contains many other flavor compounds during cheese ripening (Yvon
proteolytic enzymes such as cathepsin D and and Rijnen, 2001; Marilley and Casey, 2004;
cathepsin B, which have been confirmed for the McSweeney, 2004a,b; Dimitrellou et al., 2010).
milk coagulation activity (Hurley et al., 2000;
Magboul et al., 2001; McSweeney, 2004b).
Catabolism of branched-chain amino
Lactic acid bacteria (LAB) are proteolytic, and acids
possess a proteinase and a wide range of
peptidase, which are important for the final stages Catabolism of branched-chain amino acids such
of proteolysis in cheese ripening, ultimately as leucine, isoleucine and valine are initiated by
producing small peptides and amino acids in cheese the action of aminotransferase producing
(Fox et al., 2000). The proteolytic system of corresponding α–keto acids (α-ketoisocaproate,
Lactococcus is basically composed of a cell α-keto-β-methyl valerate and α-ketoisovalerate,
envelope-associated proteinase (CEP) or PrtP, and respectively) (Yvon and Rijnen 2001; Marilley
intracellular proteinases especially peptidases and and Casey, 2004). Further, the α–keto acids are

20
AGRIEAST 2015 (10) Pagthinathan and Nafees : Cheese ripening

degraded to other compounds by enzymatic or non methyl-2-butanone, 2-methyl propionate, 2-

enzymatic reactions (McSweeney, 2004b) and methyl-1-propanal, 2-methyl butyrate, and 2-
may be involved in a series of biochemical methyl butanal (Rychlik and Bosset, 2001; Yvon
reactions such as oxidative decarboxylation, and Rijnen 2001; Marilley and Casey, 2004;
decarboxylation and reduction as illustrated in McSweeney, 2004). Curtin and McSweeney
Figure 3, releasing volatile flavour compounds (2004) reported that aminotransferase is a rate-
including phenylacetic acid, phenethanol, p-cresol, limiting step for the production of flavour
methane thiol, dimethyl disulphide, 3-methyl compounds in cheese.
butyrate, 3-methyl butanal, 3- methyl butanol, 3-

Figure 3 Catabolism of Branched- Chain Amino Acids

(Source: Marilley and Casey, 2004)

Catabolism of aromatic amino acids catabolised in a series of metabolic pathways to

produce flavor compounds in cheese ripening
The first step of the catabolism of aromatic (McSweeney, 2004b), which includes
amino acids is by transamination (Figure 4), benzaldehyde, phenyl acetaldehyde, phenyl
which produces α–keto acids indole pyruvate, ethanol, phenyl acetate, phenyl propanoate,
phenyl pyruvate and p-hydroxy-phenyl pyruvate indole, skatole and p-Cresol (Moio et al., 2000;
from tryptophan, phenylalanine and tyrosine, Curioni. and Bosset, 2002; McSweeney,
respectively (Yvon and Rijnen 2001; Marilley and 2004b).
Casey, 2004). The α–keto acids are further

21
AGRIEAST 2015 (10) P. 16 - 26

NH3 CO2

Deaminase Decarboxylase

Phenylalanine Phenylethylamine
Phenylpropionate
¢
-Ketoacid
Aminoacid
Aminotransferase

Phenylpyruvate

Phenyllactate
Benzaldehyde
Phenylethanol
Phenylacetate

Figure 4. Catabolic pathways of phenylalanine

(Source: McSweeney, 2004b)

Catabolism of methionine
(McSweeney and Sousa, 2000). The
degradation of sulphur-containing amino acid
Sulphur-containing volatile compounds are
(methionine) is also initiated by a transamination
important flavours in many cheese varieties,
producing 4-methylthio- 2-ketobutyric acid
especially in Cheddar cheese and are major
(Figure 5) (Marilley and Casey, 2004).
contributors to aroma development in cheese
Demethiolation or/ and decarboxylation of 4-

Figure 5. Catabolism of methionine and generation of volatile sulphur compound

(Source: Marilley and Casey, 2004)

22
AGRIEAST 2015 (10) Pagthinathan and Nafees : Cheese ripening

methylthio-2-ketobutyric acid liberate the sulphur- fermentation of lactose mainly to lactic acid is
containing flavour compounds including, dimethyl caused by starter microorganisms. Lipolysis is
disulphide, dimethyl trisulphide, S-methyl quite limited in most cheese. However the
thioacetate, boiled potato-like (methional) and proteolysis is the most complex process, which
cooked cabbage like methanethiol (McSweeney is not possible to characterize in detail. The
and Sousa, 2000; Yvon and Rijnen, 2001; Thierry primary reactions such as increase in pH and
and Maillard, 2002; Marilley and Casey, 2004). hydrolysis of protein matrix are primarily
In addition to ammonia produced by deamination, responsible for changes in cheese texture, but
it contributes to flavour in certain varieties of they are a minor contribution to cheese flavor.
smear cheese. It may also contribute to elevation The catabolism of lactic acid has minor
of cheese pH during ripening. Decarboxylation influence on the flavor of most cheeses. The
of amino acids produce amine, which has strong catabolism of fatty acid to methyl ketones via
and unpleasant aroma, and adversely affects the β-oxidation and decarboxilation are a major
cheese quality. The rate of amine production contribution to flavor characters of some cheese
usually depends on the concentration of precursor (eg. blue molt ripened cheese). While the
amino acids, cheese micro flora (non-starter catabolism of amino acid is a major contribution
lactobacilli and enterococci), ripening temperature, to the flavor of many cheese varieties. Lactic
pH and salt concentration (Curtin and acid metabolism and the ammonia produced in
McSweeney, 2004; McSweeney, 2004a; many of the reactions are contributing to the
McSweeney, 2004b). pH of cheese during ripening, and this change
in the pH affects the texture of the cheese,
salability and activity of the enzyme. Since the
Microbial ripening biochemistry of cheese ripening is responsible
for the development of flavor, texture and
Cheese contains a diversity of microorganisms appearance, these biochemical reactions are a
and enzymes which could cause biological,
prerequisite for controlling and modifying
biochemical and chemical changes in all varieties
cheese ripening. In general, this biochemical
of cheeses. Generally in all cheeses, the enzymes
reactions, particularly secondary reaction of
originating from the milk, coagulant and starter
amino acid catabolism remain as challenges for
are not sufficient to make a ripening (Fox et al.,
future researchers in this area.
2000; Shetty et al., 2006). The action of these
enzymes combined with enzymes from the
secondary microflora induces changes to the
flavour, aroma and texture in cheese during References
ripening. The rate and extent of microbial ripening
is directly dependent on the percentage salt-in- Alewijn, M., E.L. Sliwinski and J.T.M. Wouters
moisture in the in cheese (Fox et al., 2000). (2005). Production of fat-derived (flavour)
Microbial ripening is mainly dependent on pH, compounds during the ripening of Gouda
ripening temperature, water activity, and
cheese. International Dairy Journal 15:
concentration of salt, oxidation reduction potential,
NO 3 , and other compounds (bacteriocins). 733-740.
However, cheese ripening involves the
combination of microbial and biochemical changes Awad, S. (2006). Texture and flavour
in cheese (Fox et al., 2000; Curtin and development in Ras cheese made from
McSweeney, 2004). raw and pasteurized milk. Food
chemistry 97: 394-400.
Summary

Cheese ripening involves a very complex series Awad, S. (2007). Effect of sodium chloride
of biochemical reactions by living organisms or and pH on the rennet coagulation and
by enzymes from several sources. The primary gel firmness. LWT 40: 220–224.
events are glycolysis, lipolysis and proteolysis. The

23
AGRIEAST 2015 (10) P. 16 - 26

Bosset, J.O. and R. Gauch (1993). Comparison Deckelbaum, R. J., J.A. Hamilton., A. Moser.,
of the volatile flavour compounds of six G. Bengtsson-Olivecrona., [Link].,
European ‘AOC’ cheeses by using a new Y.A. Carpentier., A. Gutman and T.
dynamic headspace GCMS method. Olivecrona (1990). Medium-chain
International Dairy Journal 3: 359– versus long chain triacylglycerol
377. emulsion hydrolysis by lipoprotein lipase
and hepaticlipase: Implications for the
Bruno, M. A., M.C. Lazza., M.E. Errasti., L.M.I. mechanisms of lipase action.
Lò pez., N.O. Caffini and M.F. Pardo Biochemistry 29: 1136–1142.
(2010). Milk clotting and proteolytic
activity of an enzyme preparation from Deeth, H. C. and Fitz-Gerald, C. H. (1995).
Bromelia hieronymi fruits. LWT - Food Lipolyticenzymes and hydrolytic
Science and Technology 43 (4): 695-701. rancidity in milk and milk products. In
P. F. Fox (Ed.), Advanced dairy
Chalier, P. and J. Crouzet (1992). Production of chemistry-2-Lipids (pp. 247–308).
lactones by Penicillium roqueforti. London: Chapman & Hall.
Biotechnology Letters 14:275-280.
de Figueroa, R., F. Alvarez., A. Pesce de Ruiz
Christensen, J.E., E.G. Dudley., J.A. Pederson and Holgado., G. Oliver and F. Sesma
(2000). Citrate utilization by homo- and
J.L. Steele (1999). Peptidases and amino
heterofermentative lactobacilli.
acid catabolism in lactic acid bacteria.
Microbiologyn Research 154: 313-
Antonie van Leeuwenhoek, 76: 217-246.
320.
Cogan, T. M. and C. Hill (1993). Cheese starter
de Figueroa, R.M., G. Oliver and I.L. Benito de
cultures. In Cheese: Chemistry, Physics
Cardenas (2001). Influence of
and Microbiology, Vol 1: General
temperature on flavour compound
Aspects, 2nd edn, pp 193–256. Fox P F,
production from citrate by
[Link]: Chapman and Hall.
Lactobacillus rhamnosus ATCC 7469.
Microbiologyn Research 155: 257-
Collins, Y. F., P.L.H. McSweeney and M.G. 262.
Wilkinson (2003). Lipolysis and free fatty
acid catabolism in cheese: a review of Dimitrellou, D., Y. Kourkoutas., and A.A.
current knowledge. International Dairy Koutinas and M. Kanellaki (2009).
Journal 13: 841-866 Thermally-dried immobilized kefir on
casein as starter culture in dried whey
Curioni, P.M.G. and J.O. Bosset (2002). Key
cheese production. Food Microbiology
odorants in various cheese types as
determined by gas chromatography– 26: 809-820.
olfactometry. International Dairy
Journal 12: 959-984. Dimitrellou, D., P. Kandylis., A. Mallouchos.,
M. Komaitis., A.A. Koutinas and Y.
Curtin, A. C. and P.L.H. McSweeney (2004). Kourkoutas (2010). Effect of freeze-
Catabolism of amino acids in cheese dried kefir culture on proteolysis in feta-
during ripening. In Cheese: Chemistry, type and whey-cheeses. Food
Physics and Microbiology, Vol 1: General Chemistry 119: 795-800.
Aspects, 3rd edn, pp 435-454. Fox, P. F.,
McSweeney, P. L. H, Cogan, T. M. and Dubey, V.K., M. Pande., B.K. Singh and M.V.
Guinee, T. P., eds. London: Elsevier. Jagannadham (2007). Papain-like
proteases: Applications of their
inhibitors. African Journal of
Biotechnology 6 (9): 1077-1086.

24
AGRIEAST 2015 (10) Pagthinathan and Nafees : Cheese ripening

Dybing, S. T., J. [Link]., S. A. Brudvig., E. Katechaki., E., P. Panas., Y. Kourkoutas., D.

A. Huang and R. C. Chandan (1988). Koliopoulos and A. A. Koutinas (2009).
Effect of processing variables on the Thermally-dried free and immobilized
formation of calcium lactate crystals on kefir cells as starter culture in hard-type
Cheddar cheese. Journal of Dairy Science cheese production. Bioresource
71:1701-1710. Technology 100: 3618-3624.

Fox, P. F., J. Law., P. L. H. McSweeney and J. Magboul, A.A.A., L. B. Larsen., P. L. H.

Wallace (1993). Biochemistry of cheese McSweeney and A. L. Kelly (2001).
ripening. In P.F. Fox (Ed.), Cheese: Cysteine protease activity in bovine
Chemistry, Physics and Microbiology. milk. International Dairy Journal 11:
Vol. 1 (2nd ed.) (pp. 389-438). London: 865-872.
Chapman and Hall.
Marilley, L. and M. G. Casey (2004). Flavors
Fox, P. F. and L. Stepaniak (1993). Enzymes in
of cheese products: metabolic
cheese technology. International Dairy
pathways, analytical tools and
Journal 3: 509-530.
identification of producing strains.
International Journal of Food
Fox, P.F., T. K. Singh and P. L. H. McSweeney
Microbiology 90:139-159.
(1995). Biogenesis of flavour compounds
in cheese. Adv. Exp. Med, 367: 59-98.
Nourani, A., E. Dako., A. Morsli., N.
Fox, P. F. and J. M. Wallace (1997). Formation Belhamiche., S. Belbraouet., M. M.
of flavour compounds in cheese. Advances Bellal and A. Dadie (2009).
in Applied Microbiology 45: 17-85. Characterization of the purified
coagulant extracts derived from
Fox, P. F. and McSweeney, P. L. H. (1998). Dairy Artichoke flowers (Cynara scolymus)
Chemistry and Biochemistry. Blackie and from the Fig tree latex (Ficus
Academic and Professional, London. carica) in light of ntheir use in the
manufacturer of traditional cheese in
Fox, P. F., Guinee, T. P., Cogan, T. M. And Mc
Sweeney, P. L. H. (2000). Fundamental Algeria. Journal of Food Technology 7
of Cheese Science. Gaithesburg, (1): 20-29.
Mayyland, An Aspen Publisher, Inc.
Olivecrona, T., Vilaro, S. and Bengtsson-
Hannon, J. A., K. N. Kilcawley., M. G. Olivecrona, G. (1992). Indigenous
Wilkinson., C. M. Delahunty and T. P. enzymes in milk. II. Lipases in milk. In
Beresford (2007). Flavour precursor P. F. Fox (Ed.), Advanced dairy
development in Cheddar cheese due to chemistry. 1. Proteins (pp. 292-310).
lactococcal starters and the presence and London: Elsevier Applied Science.
lysis of Lactobacillus helveticus.
International Dairy Journal 17: 316-327. Parente, E. and T. M. Cogan (2004). Starter
cultures: general aspects. In Cheese:
Hill, A. R. (1995). Chemical species in cheese and Chemistry, Physics and Microbiology,
their origin in milk components. In Vol 1: General Aspects, 3rd edn, pp
Chemistry of Structure Function 123–148. Fox P F, McSweeney P, L,
Relationships in Cheese, Malin, E.L. and H., Cogan, T. M. and Guinee, T. P.
Tunick, M.H. Editors. Plenum Press, NY.
eds. London: Elsevier.
Hurley, M.J., L. B. Larsen., A. L. Kelly and R. L.
H McSweeney (2000). The milk acid
proteinase, cathepsin D: a review.
International Dairy Journal 10: 673-681.

25
AGRIEAST 2015 (10) P. 16 - 26

Pastorino, J., C. L. Hansen and D. J. McMahon Tavaria, F. K., M. J. Sousa., A. Domingos., F.

(2003). Effect of sodium citrate on X. Malcata., A. Brodelius., A. Clemente
structure-function relationships of cheddar and M. S. Pais (1997). Degradation of
cheese. Journal of Dairy Science 86: caseins from different milk types by
3113-3121. extracts of Centaurea calcitrapa.
Journal of Agricultural and Food
Rychlik, M. and J. O. Bosset (2001). Flavour and Chemistry 45: 3760-3765.
off-flavour compounds of Swiss Gruyere
cheese. Identification of key odorants by Thierry, A. and Maillard, M.B. (2002).
quantitative instrumental and sensory Production of cheese flavor
studies. International Dairy Journal 11: compounds derived from amino acid
903-910. catabolism by Propionibacterium
freudenreichii. Lait 82: 17-32.
Shetty, K., Paliyath, G., Pometto, A. and Levin,
R. E.(2006). Food biotechnology. Ed 2nd, Walstra, P., J. T. M. Wouters and T. J. Geurts
Taylor & Francis Group: CRC Press. (2006). Dairy Science Technology. Ed
2nd, Taylor & Francis Group, CRC
Smit, G., J. E. T. van Hylckama Vlieg., B. A. Smit., Press. Yvon, M. and Rijnen, L (2001).
E. H. E. Ayad and W. J. M. Engels Cheese flavor formation by amino acid
(2002). Fermentative formation of flavor catabolism. International Dairy Journal
compounds by lactic acid bacteria. 11: 185-201.
Australian Journal of Dairy Technology
7: 61-68.

Sousa, M. J., Ardo, Y. and P. L. H. McSweeney

(2001). Advances in the study of
proteolysis in cheese during ripening.
International Dairy Journal 11: 327-345.

26