doi: 10.1111/1471-0307.

12755

REVIEW
The cheese matrix: Understanding the impact of cheese
structure on aspects of cardiovascular health – A food
science and a human nutrition perspective
EMMA L FEENEY,1,2 PRABIN LAMICHHANE3 and
JEREMIAH J SHEEHAN*2,3
1
Institute of Food and Health, University College Dublin, 2.16a Science Centre South, Dublin 4, 2Food for Health
Ireland (FHI), S2.09 Science Centre South, Belfield, Dublin 4, and 3Teagasc Food Research Centre, Moorepark,
Fermoy, Cork P61 C996 Ireland

Health guidelines recommend limiting saturated fat consumption due to adverse associations with
low-density lipoprotein cholesterol, a marker for cardiovascular disease risk. Recently, this advice
is being questioned, since it does not account for the diversity of fatty acids present in different
foods and may be overly simplistic. Current research suggests that for dairy foods and cheese in
particular, a matrix effect exists, whereby the other components present interact with the overall
structure, leading to health benefits. This review examines how factors in cheese production and
processing impact this matrix, and considers how they affect biological function, and the potential
impact on human health.
Keywords Cheese matrix, Structure, Composition, Saturated fat, Cardiovascular disease, Human
health.

within the cheese matrices resulting in develop-

INTRODUCTION
ment of the characteristic texture, aroma and fla-
The current global cheese market is valued at ~ vour. Other cheeses are prepared by acid
*Author for
correspondence. E-mail:
$100 bn, and global cheese consumption is coagulation (e.g. Cottage, Quarg) or by a combi-
diarmuid.sheehan expected to increase by ~13.8% between 2019 nation of heat and acid (e.g. Queso blanco,
@teagasc.ie and 2029 (OECD/FAO, 2020). Although cheese Paneer), and are normally consumed fresh. All
All authors contributed has been part of the European diet for 6000– of the above categories of cheese are referred to
equally to this work. 8000 years (Salque et al. 2013), it is now also as natural cheeses, alternatively other cheese
© 2021 The Authors.
gaining popularity in countries where it was not types may be categorised as ‘processed’ or ‘ana-
International Journal of traditionally part of the national diet, likely due logue’ cheese (Guinee 2016). Processed cheeses
Dairy Technology to the Westernisation of the diet (OECD/FAO are prepared by blending shredded natural
published by John Wiley 2020). Dairy consumption has increased signifi- cheese with other dairy and nondairy ingredi-
& Sons Ltd on behalf of cantly in Asia in the last two decades (Lipoeto ents, including emulsifying salts, which act as
Society of
Dairy Technology
et al. 2012), and although milk still accounts for calcium chelating agents promoting disaggrega-
This is an open access much of the consumption (de Goede et al. tion of the protein in the cheese matrix, and
article under the terms of 2016), it is estimated that the retail value of where the fat is contained as an emulsion within
the Creative Commons cheese in China will increase at an annual rate the matrix (Kapoor and Metzger, 2008; Guinee
Attribution-NonCommerc of 12% during the forecast period of 2019–2024 2016). Cheese analogues are cheese-like prod-
ial-NoDerivs License,
which permits use and
(Euromonitor 2019). ucts in which milk fat, milk protein or both are
distribution in any Approximately 2000 cheese varieties are partially or wholly replaced by non-milk-based
medium, provided the reported, mostly prepared by coagulation of components and prepared by blending various
original work is properly milk by chymosin, and matured for between edible fats/oils, proteins, other ingredients and
cited, the use is non- 2 weeks and 2 years (McSweeney 2004). Dur- water into a smooth homogenous blend with the
commercial and no
modifications or
ing maturation numerous physico-chemical, aid of heat, mechanical shear and emulsifying
adaptations are made. microbiological and biochemical changes occur salts (Cunha et al. 2010; Guinee 2016). Such

Vol 0 International Journal of Dairy Technology 1

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diversity in manufacture procedures results in cheeses as called for a revisit of the guidelines. This debate continues,
complex dairy products, having broad ranges in composition with a recent group noting that the 2018 draft WHO guide-
and structural characteristics. lines could even inadvertently promote a less healthful diet
Cheeses are generally nutrient-dense foods and are a valu- through unnecessary reduction of nutrient-dense foods, such
able source of high-quality proteins, lipids, vitamins (e.g. as cheese (Astrup et al. 2019). Saturated fats are not a
vitamin A, B2 and B12) and minerals (particularly calcium homogenous group, but rather they vary considerably, from
and phosphorus). In the Irish diet, as well as in the UK and the type and length of the SFA, to the nature of the food in
the United States, they make a significant contribution to which they are contained, including other nutrients, as well
nutrient intake and diet quality (Feeney et al. 2016). In as the overall food structure (Thorning et al. 2017), and
addition to macro and micronutrients, some matured cheeses specific fatty acids have different effects on health out-
contain bioactive components (e.g. bioactive peptides), comes. This demonstrates a need to distinguish the individ-
which have health benefits, while beneficial bacteria present ual food source of the saturated fat in dietary guidelines
in the cheese matrix can potentially improve human gut (Astrup et al. 2019; Wu et al. 2019). Further, Astrup et al.
health by producing short-chain fatty acids (Santiago-L
opez (2020) note that while SFA intake is associated with total
et al. 2018). However, cheese also contains relatively high LDL-c levels, LDL-c particle size is a much greater indica-
levels of saturated fatty acids (SFAs), which are commonly tor of CVD risk and the small dense LDL particles are con-
perceived as negatively impacting the healthfulness of the sidered more atherogenic and thus confer a greater CVD
diet, and have been associated with increased blood LDL-c- risk than large LDL-c particles. Specifically, they note these
holesterol levels, often considered a marker for cardiovascu- smaller LDL-c particles are not reduced via dietary SFA
lar disease (CVD) risk (Ference et al. 2019). restriction in most individuals. Therefore, SFA restriction
Current dietary guidelines recommend that the daily can result in a disproportionately greater reduction of the
intakes of saturated fat (SFA) should be as low as possible large LCL-c particles, which are much less associated with
(EFSA, 2010), and should exceed no more than 10% of CVD risk. For this reason, they highlight the pressing need
total energy intake (USDA, 2015). Population intakes of to develop more reliable markers to monitor the effects on
SFA are considerably higher than this, 13%–14% in Ireland CVD risk from dietary SFA.
(Tierney et al. 2011) and 12.6% in the UK (PHE, 2014). There has been particular interest in dairy foods as a
Recent draft guidelines published by the WHO (2018) rec- source of saturated fat in recent times, and specifically in
ommend a reduction in population SFA intakes, and cheese. In one cross-sectional study of over 18 000 subjects,
replacement with poly- and monounsaturated fatty acids in cheese consumption was correlated with beneficial meta-
order to reduce rates of chronic disease and related deaths. bolic health markers, including higher circulating HDL-c
Dairy foods are a major contributor to population SFA levels (Hostmark et al. 2009), while a later meta-analysis of
intakes, as circa 60% of the fats in dairy fat are in the satu- 15 prospective cohort studies found a (nonlinear) reduced
rated form, contributing approximately 1/5 of SFA on aver- risk of CVD risk with increasing cheese consumption (Chen
age, according to data from UK, Irish and US food et al. 2016). Further, several randomised controlled trials
consumption databases (Feeney et al. 2016). Therefore, have demonstrated that the consumption of fat from cheese
dairy foods, and cheese in particular, are a considerable tar- specifically, does not increase LDL-c compared to the same
get for reduction on a population intake level. However, the amount of fat from some other dairy sources (Biong et al.
supporting evidence for a link between SFA intake and 2004; Nestel et al. 2005; Hjerpsted et al. 2011; Hjerpsted
health is more nuanced and indicates that the food source of and Tholstrup, 2011; de Goede et al. 2015; Feeney et al.
the saturated fatty acids may be of particular importance. A 2018).
number of recent meta-analyses suggest that while SFA Since the fat contained in different dairy foods share a
from meat and processed meat sources are associated with similar fatty acid profile, this further highlights the impor-
detrimental health effects (de Oliveira Otto et al. 2012), tance of the food source and structure, and the interaction
SFA intake from dairy sources on the other hand has con- with the other nutrients contained within that food (i.e. the
trasting health outcomes and is associated generally with food matrix; Thorning et al. 2017) in the study of saturated
either neutral health effects (Benatar et al. 2013) or with fat consumption, and health effects. Although equally the
beneficial health associations (Elwood et al. 2010; Chen validity of LDL-c as a marker of CVD risk is under debate
et al. 2012; de Oliveira Otto et al. 2012; Kratz et al. 2013). (Astrup et al. 2020), the existence of a food matrix effect is
For a recent review of these effects from association studies now well accepted.
and randomised controlled trials, examining the individual The aim of this review is to examine the cheese matrix
products milk, cheese and yoghurt, please see Timon et al. from a both a physico-chemical and biological perspective,
(2020). and to determine how interactions between both phases may
Lawrence (2013) concluded that the scientific evidence exert beneficial effects arising from cheese consumption on
did not support a blanket minimisation of SFA intakes, and aspects of cardiovascular health.

2 © 2021 The Authors. International Journal of Dairy Technology published by John Wiley & Sons Ltd on behalf of Society of Dairy Technology
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various types of microscopy) differs markedly (Figure 1)

CHEESE COMPOSITION AND STRUCTURE
due to a number of factors, such as composition and pre-
Cheese compositions vary according to variety (Table 1) treatment of cheese milk (e.g. homogenisation and heat
including for the major components: moisture, protein treatment), cheese manufacturing conditions (e.g. method of
(mainly casein) and fat. For example, Cottage cheese con- coagulation), maturation conditions (e.g. duration and matu-
tains high levels of moisture and low levels of protein and ration temperatures) and preparation before consumption
fat, whereas Parmesan contains low level of moisture and (e.g. cooking, baking, grilling). For example, cream cheese
high levels of protein and fat (Table 1). Additionally, microstructure is composed of compact fat and protein
cheeses also contain varying quantities of other numerous aggregates with large spaces filled with whey (Figure 1a),
minor components, including calcium (calcium phosphate), which is mainly due to its composition (fat: protein ratio,
and salt (sodium chloride). moisture content) as well as the prior homogenisation of
The cheese matrix itself is a complex assembly of the cheese milk. The ‘cheddaring’ steps during Cheddar cheese
individual components/nutrients. Protein, especially casein, manufacture and cooking of curds to high temperatures (50–
hydrated with water forms networks (within the cheese 55 °C) during Emmental manufacture contributes to the coa-
matrix) in which fat globules/pools, minerals, bacteria and lescence of fat globules and formation of large irregular fat
dissolved solutes such as lactic acid, potentially residual lac- pools within the cheese matrix (Figure 1b,e). Similarly, the
tose, soluble salts and peptides are all interspersed (Hickey stretching of curd in hot water or cooling in brine during
et al. 2015; Lamichhane et al. 2018b). The manner in which the production of Mozzarella cheese results in the orienta-
the individual components/nutrients assemble within the tion of protein stands and aggregated fat globules/pools in
cheese matrix and their interactions determines the structure the direction of stretching (Figure 1d). The specific manu-
of cheese. The structural organisation of the major con- facturing steps of process cheese results in entrapment of
stituents of various cheeses (which can be observed under spherical fat globules within the homogenous protein matrix

Table 1 Approximate composition of selected cheese varieties.a

Moisture Protein Fat MNFS FDM Calcium Calcium Salt S/M Cholesterol
(mg/ (mg/g
Cheese type (g/100 g) (g/100 g) (g/100 g) (%) (%) 100 g) protein) pH (g/100 g) (%) (mg/100 g)
Extra hard
Parmesan 30.6 34.9 29 43 41.79 1200 34.38 5.2–5.3 1.89 6.18 88
Hard
Cheddar 34.5–37.5 24.7–25.1 32.8–33.9 51–56 51.5–52.5 780– 28.24 5.2–5.3 1.6–2.2 4.5–5.8 100
830
Emmental 35.7 28.7 29.7 51 46.19 970 33.80 5.5 0.70 1.96 90
Gruyere 35 27.2 33.3 52 51.23 950 34.93 5.5 1.48 4.23 100
Semihard
Mozzarella 49.8 25.1 21 63 41.83 590 23.51 5.1–5.3 1.40 2.81 65
Gouda 40.1 24 31 58 51.75 740 30.83 5.2–5.4 2.30 5.74 100
Edam 43.8 26 25.4 59 45.20 770 29.62 5.2–5.4 1.45 3.31 80
Semisoft
Danish 41.3 19.7 32.9 62 56.05 530 26.90 5.3 4.10 9.93 90
blue
Stilton 45.3 20.1 29.6 64 54.11 500 24.88 6.6 3.30 7.28 75
Roquefort 38.6 22.7 35.5 60 57.82 320 14.10 6.0–6.50 2.20–2.70 5.70–6.99 105
Soft
Cottage 79.1 13.8 3.9 82 18.66 73 5.29 4.8 0.50–0.70 0.63–0.88 13
Brie 48.6 19.3 26.9 66 52.33 540 27.98 7.5 1.40–2.10 2.88–4.32 100
Fromage 77.9 6.8 7.1 84 32.13 89 13.09 4.5–4.8 0.15 0.19 25
frais

MNFS, moisture-in-non-fat substance; FDM, fat in dry matter; and S/M, salt-in-moisture.
a
Data compiled from multiple sources: Lawlor et al. (2002), Lawlor et al. (2003), Govindasamy-Lucey et al. (2004), Guinee (2007), Heino et al.
(2010), Calzada et al. (2014), Guinee (2016), McCarthy et al. (2017), and O’Brien and O’Connor (2017).

© 2021 The Authors. International Journal of Dairy Technology published by John Wiley & Sons Ltd on behalf of Society of Dairy Technology 3
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(a) (b) (c) (d)

(e) (f) (g) (h)

Figure 1 Microstructure of various cheese types: (a) Cream cheese, (b) Cheddar cheese; arrow shows curd granule junction, (c) Processed Cheese,
(d) Mozzarella cheese, (e) Emmental cheese, (f) Camembert cheese, (g) whipped cream cheese and (h) soft cheese made with ultrafiltration technol-
ogy. Micrographs (a) to (d) are adapted from Auty et al. (2001); the protein phase appears red while the fat phase appears blue; scale bar 25 µm.
Micrographs (e) to (h) are adapted from Lopez (2005); fat is coloured in red, proteins are in grey levels. Black areas correspond to serum or gas
holes.

(Figure 1c). In the following sections, factors influencing casein micelles by different means have previously been
the cheese matrix as well as their nutritional importance will reviewed extensively (please see review by Dalgleish and
be discussed. Corredig, 2012). The destabilised casein micelles aggregate
into chains and clusters, leading to formation of a 3-dimen-
Milk pretreatment and coagulation of milk sional gel.
Caseins and whey proteins are the two major groups of pro- It is expected that coagulation methods also alter the
tein found in milk, comprising ~80% and ~20% of total structure of casein networks, as well as the molecular inter-
milk protein, respectively. In milk, the caseins exist as large actions between the caseins within the cheese matrices
colloidal aggregates, known as casein micelles, where the j- (Lucey 2002; Lamichhane et al. 2018b). Such different
casein present on the surface of casein micelles stabilises structures within casein networks will also behave differ-
against aggregation in milk (Dalgleish and Corredig, 2012). ently during digestion, affecting the bioavailability of the
During cheese manufacture, casein micelles are destabilised nutrients contained within. For example, Floury et al.
intentionally, which is achieved through chymosin addition, (2018) showed that milk-based chymosin gels, but not acid
acidification, heat or a combination of acidification and heat. gels formed compact protein aggregates under acidic condi-
The structure of casein micelles is altered differently tions of the stomach. Moreover, the kinetics of protein
depending on the coagulation method used (Dalgleish and hydrolysis was slow in milk-based chymosin gels as com-
Corredig, 2012). For example, in chymosin-induced gela- pared to acid gels. Similarly, Barb
e et al. (2014) in in vivo
tion, destabilisation of casein micelles occurs due to selec- studies, using mini pigs, showed that the digestion of milk
tive hydrolysis of j-casein (present on the surfaces of proteins in chymosin gels was delayed as compared to acid
casein micelles) at the Phe105-Met106 peptide bond by gels.
enzymes present in chymosin, whereas in acid-induced gela- Milk used for the manufacture of most cheese varieties is
tion, destabilisation of casein micelles occurs due to acidifi- generally pasteurised, most commonly at 72 °C for 15 s, to
cation (using starter cultures or food grade acids and/or eliminate pathogenic bacteria (Kelly et al. 2008); such a
acidulants), which decrease the negative charge of the mild heat treatment has minimal impact on the structure of
micelle surface (Lucey 2002; Dalgleish and Corredig, 2012; milk components. However, more severe heat treatments
Guinee 2016). Detailed mechanisms of destabilisation of (e.g. 85 °C for 5 min for the production of Queso blanco;

4 © 2021 The Authors. International Journal of Dairy Technology published by John Wiley & Sons Ltd on behalf of Society of Dairy Technology
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Farkye 2004) are applied in the manufacture of several other added components (such as milk fat or cream) is con-
acid-heat coagulated cheeses (e.g. Queso blanco and Paneer) centrated by evaporation to achieve the final product (Jelen
and may also be used in the manufacture of reduced or low- 2002).
fat cheeses. High heat treatment of milk (greater than pas- In vitro and in vivo studies have reported that the whey-
teurisation) results in high levels of whey protein denatura- based cheese showed different digestion and absorption
tion. For example, ~34 % of total whey protein has been behaviour as compared to casein-based cheese (Lorieau
reported to denature in milk heated at 87 °C for 26 s et al. 2018; Lorieau et al. 2019). For example, in an in vivo
(Rynne et al. 2004). The denatured whey protein interacts study using pigs, Lorieau et al. (2019) observed a higher
with caseins, MFGM proteins or both via a number of amino acid bioavailability in whey-based cheeses as com-
molecular forces, including disulphide bonds (Kelly et al. pared to casein-based cheeses, and those authors suggested
2008; Michalski 2009), and there is current interest in that cheese based on whey proteins rather than those based
understanding how such process-induced modifications of on caseins are potentially more suitable for muscle synthesis
structures or nutrients influence their disintegration or in elderly people.
release patterns in the gastrointestinal tract under digestion.
To date, there is little published data on the digestion Cheese manufacture processes
behaviour of cheese made from high heat-treated milk (e.g. Although acid or acid-heat coagulated cheese types undergo
Queso blanco and Paneer). In cheese made from unheated minimal curd handling processes after curd formation, chy-
(or minimally heated) milk, caseins are responsible for net- mosin-coagulated curd is subjected to several post-curd for-
work formation, whereas in cheese made from high heat- mation processes, such as cutting, cooking, ‘cheddaring’ (in
treated milk, both denatured whey proteins and caseins are Cheddar cheese) and hot-water stretching (in pasta-filata type
responsible for network formation. Moreover, denatured cheeses), and moulding and pressing. Such processes con-
whey protein associated with the surface of casein micelles tribute to the characteristic structure, texture and rheological
may restrict the rearrangements and/or fusion of casein par- properties of the final cheese. For example, draining of whey
ticles in cheese made from high heat-treated milk (Lucey at higher curd pH, such as 6.4–6.5 in Emmental and Maas-
et al. 2001). Therefore, cheeses made from high heat-treated dam cheeses, results in higher levels of micellar/insoluble
milk are more likely to show different digestion behaviours, calcium (i.e. calcium associated with casein), which is
and further research on this is recommended. responsible for their rubbery or elastic texture. Levels of
A number of in vitro studies however investigated the insoluble calcium in ripened Gouda, Emmental and Maas-
impact of heat treatment of milk on its digestion behaviour. dam cheese are ~24 mg/g protein (Lamichhane et al. 2018b;
For example, Ye et al. (2016a; 2016b) observed a faster rate Lamichhane et al. 2019), higher than those in Cheddar
of proteolysis by pepsin in high heat-treated (90 °C for cheese (~18 mg/g protein), as whey is drained at lower pH
20 min) milk samples as compared to unheated milk. Simi- (i.e. 6.15–6.30) during Cheddar cheese manufacture. In acid
larly, Mulet-Cabero et al. (2019) observed an accelerated coagulated cheeses, such as Cottage cheese, the level of
protein hydrolysis in ultra-high-temperature (140 °C for 3 s) insoluble calcium is very low (i.e. <5 mg/g protein), most
processed milk. Moreover, apart from protein hydrolysis, probably due to solubilisation of the colloidal calcium phos-
studies have also reported that high temperature processing phate associated with the caseins at low pH (i.e. ~4.6). Dur-
of milk increased the free fatty acid release during in vitro ing Cheddar cheese manufacture, the curd granules are
digestion (Tunick et al. 2016; Ye et al. 2016a). In addition, allowed to fuse by stacking ‘loaves’ of curd on top of one
during in vitro digestion high heat-treated milk formed frag- another, called ‘cheddaring’, which results in coalescence of
mented and soft coagula, whereas unheated milk formed fat globules into large fat pools and development of the char-
dense-structured coagula (Ye et al. 2016a; 2016b; Mulet- acteristic fibrous structure (Lucey et al. 2003). Similarly, the
Cabero et al. 2019), which may potentially be the reason stretching of curd in hot water or brine during the production
for different rates of protein hydrolysis as well as different of Mozzarella cheese results in orientation of protein stands
rates of fat globule release from the coagula (Ye et al. in the direction of stretching, resulting in fibrous structure
2016a). and anisotropy, that is physical and mechanical properties
Whey is also used as a starting material for manufacture dependent on the direction of examination (Bast et al. 2015;
of some cheese varieties (also called whey-based cheeses), Sharma et al. 2018). The specific manufacturing conditions
such as Ricotta, Anari and Mysost (Phelan et al. 1993; Jelen of Parmesan-type cheeses, such as cutting of coagula into
2002). Ricotta and Anari are manufactured by coagulation small curd granules, high scalding temperatures (55 °C for
of whey proteins, which is achieved by high heat treatment 30 min) and long brining times, is responsible for a very dry
of whey (~90 °C for few minutes) and often addition of cheese with a very hard and brittle texture and evinced by
organic acids (e.g. acetic, citric) and/or mineral salts (e.g. its low moisture and also moisture-in-non-fat substance
calcium), to the whey (Smithers 2008). For manufacture of (MNFS) levels, 43 % in comparison with 82 % in Cottage
the Norwegian-style whey cheese, Mysost, whey with some cheese. Levels of curd hydration are also influenced by curd

© 2021 The Authors. International Journal of Dairy Technology published by John Wiley & Sons Ltd on behalf of Society of Dairy Technology 5
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pH, such as the swelling of the outer layers of the para-ca- the disintegration of the food bolus (Kong and Singh, 2008;
sein matrix in Brie and Camembert as pH increases during Singh et al. 2015).
maturation and by salt-in-moisture levels. Such initial cheese Disintegrated and partially digested food from the stom-
characteristics may have an impact on the breakdown/disinte- ach is then passed to the small intestine, where many of the
gration behaviour in the mouth during mastication and in the digestive reactions occur, such as hydrolysis of protein and
gastrointestinal tract during digestion, which may affect the fat into amino acids and free fatty acids, and absorption of
kinetics of release of nutrients. Matrix disintegration is one most of the nutrients occurs (Kong and Singh, 2008; Norton
of the important events for digestion of solid food matrix, et al. 2014). Insoluble and indigested food materials from
such as cheese, which facilitates release of nutrients to the the small intestine then pass to the large intestine, where
digestion medium. microorganisms perform fermentation producing gas and
The first step of the food digestion process occurs in the other compounds, such as short-chain fatty acids (Norton
mouth, where food is subjected to a complex series of oral et al. 2014). A schematic overview of physico-chemical,
manipulations, including ingestion, size reduction and mix- structural and biochemical transformations of cheese as it
ing with saliva, to form a bolus for safe swallowing. Solid passes through the oral cavity, the stomach, and the small
foods, including cheese, are broken down into number of and large intestines is shown in Figure 2. More detailed
pieces into the mouth during oral manipulation. Breakdown information regarding transformation of food in the human
processes (i.e. number of fragments and their size) are lar- digestive systems has been reviewed by Kong and Singh
gely dependent on the mechanical properties of food and (2008), Norton et al. (2014), Singh et al. (2015) and Somar-
their interaction with individual oral processing parameters atne et al. (2020).
such as level of mastication (Chen 2009). For example, Guo Studies have reported that the cheese matrix disintegration
et al. (2013) reported that hard whey protein gel with an was correlated with textural properties of cheese (Sharma
inhomogeneous microstructure had a fast-crack propagation Khanal et al. 2020). For example, texture parameters
pattern and a high degree of fragmentation as compared to springiness, cohesiveness, chewiness and hardness were
soft gels of homogeneous microstructure. The mechanical negatively correlated to the rate of cheese disintegration dur-
properties of cheese may vary between cheese types, such ing in vitro gastric digestion (Fang et al. 2016a; Guinot
as the elastic and rubbery texture of Emmental cheese, and et al. 2019). Moreover, the rate of protein/peptide release
the firm and brittle texture of Parmesan (Guinee 2016). was influenced by the textural properties of cheese (Fang
Moreover, the mechanical properties of cheese also largely et al. 2016a; Sharma Khanal et al. 2020).
depend on the level of maturation; cheese usually becomes Textural properties of cheese largely depend on its com-
softer as maturation increases (Lamichhane et al. 2019). position and the structural organisation of its structural com-
Apart from mechanical processes, cheese is exposed to bio- ponents and their interactions. Casein network within cheese
chemical processes in the oral cavity, such as interactions matrix contribute for the rigidity of cheese matrix, whereas
between components of cheese and the active components moisture contribute for softening of cheese texture (Guinee
of saliva (e.g. enzymes and glycoprotein; Norton et al. 2016; Lamichhane et al. 2018b). Solid milk fat in cheese is
2014). Fat globules within the natural cheese matrices have known to act as reinforcing fillers, contributing to elastic
been reported to be held weakly within the structure properties of unheated cheese, whereas liquid fat acts as a
(Lamichhane et al. 2020); therefore, it is expected that the plasticiser between casein strands, making cheese more soft
same fat globules may be released and subsequently mixed and flexible (Rogers et al. 2010; Shima and Tanimoto,
with saliva. Moreover, the level of water-soluble nitrogen 2016). Age-related changes, such as proteolysis and solubili-
(due to hydrolysis of caseins) within the cheese matrices sation of micellar calcium, as well as hydration of the casein
increases with the level of maturation, and is likely to be strand reduce the rigidity of casein network.
solubilised in the saliva during oral manipulation. Thus, it
can be assumed that the bolus characteristics may depend Ripening
on the cheese composition, structure, mechanical properties The length of the maturation period can vary between
and their level of maturation. Bolui with different character- cheese types, such as instant freezing for certain Pizza/Moz-
istics are most likely to behave differently in the following zarella cheese, ~2–4 weeks for some surface-ripened mould
digestion steps (Singh et al. 2015). cheese, 2–3 months for Maasdam and more than 1–2 years
The bolus formed in mouth passes to the stomach through for mature Cheddar or Parmigiano-Reggiano (McSweeney
the oesophagus (Kong and Singh, 2008). In the stomach, 2004). Cheese maturation is a complex process and various
the bolus is further disintegrated due to diffusion of gastric physico-chemical, microbiological and biochemical changes
juice which is a complex mixture of gastric acid, bile salts (such as proteolysis, lipolysis and the metabolism of resid-
and digestive enzymes secreted by the gastric glands (Kong ual lactose and of lactate and citrate) occur within the
and Singh, 2008). In addition, mechanical force generated cheese matrices resulting in the texture and flavour charac-
by contraction of the stomach also play a significant role on teristic of a particular variety (McSweeney 2004).

6 © 2021 The Authors. International Journal of Dairy Technology published by John Wiley & Sons Ltd on behalf of Society of Dairy Technology
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Figure 2 A schematic overview of physico-chemical, structural and biochemical transformations of cheese as it passes through the oral cavity, the
stomach, and the small and large intestines. Abbreviation: LCFA, long-chain fatty acids. Adapted from Norton et al. (2014) and Singh (et al. 2015).

Hydrolysis of intact caseins into small and medium size was positively correlated with the hydrolysis of b-casein.
peptides and free amino acids is one of the important bio- Moreover, rapid solubilisation of calcium associated with
chemical events, which contribute to weakening of protein casein at early stages of maturation is another important
network. aS1-casein and b-casein are the two important age-related change within the cheese matrix, which also
caseins within the cheese matrix, and these undergo varying contributes to weakening of cheese structure as the calcium
degree of hydrolysis during maturation in different cheese associated with casein enhances the cross-linking of casein
varieties through the action of various proteolytic agents, within the cheese matrix (O’Mahony et al. 2005; Lamich-
such as residual coagulant and plasmin (Sheehan 2013; hane et al. 2019).
Lamichhane et al. 2018b; Lamichhane et al. 2019). For Other numerous changes occur within the cheese matrix
example, the extent of aS1-casein hydrolysis in Cheddar and during maturation. For example, changes in the composition
Maasdam cheeses is considerably higher than b-casein of serum phase due to solubilisation of minerals associated
(O’Mahony et al. 2005; Lamichhane et al. 2019), whereas with caseins (primarily calcium and phosphate) and accumu-
in blue-vined cheeses both caseins are completely hydrol- lation of breakdown products of caseins and lipids, such as
ysed at the end of maturation (McSweeney 2004). Some peptides, free amino acids and free fatty acids. In addition,
studies have indicated that the specific hydrolysis patterns the pH of the cheese matrix changes due to metabolism of
of casein can influence melting and fracture properties of residual lactose and lactic acid, and proteolytic liberation of
cheese (Bogenrief and Olson, 1995; Lamichhane et al. basic compounds. For example, in Swiss, Dutch and related
2019). For example, Lamichhane et al. (2019) observed that eye-type cheeses pH increase from 5.2–5.3 at first week of
the rigidity or strength of the Maasdam cheese matrix was maturation to 5.8–6.0 at the end of maturation (Govin-
negatively correlated with the hydrolysis of aS1-casein, dasamy-Lucey et al. 2011; Lamichhane et al. 2018a), and in
whereas the brittleness or shortness of the cheese matrix Camembert-type cheeses, pH increase dramatically from

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4.5–4.6 at 1 day to 7.0–8.0 after 3 months of maturation with ACE-inhibitory properties are Valine-Proline-Proline
(Sousa and McSweeney, 2001). and Isoleucine-Proline-Proline (VPP and IPP), first isolated
Studies have reported that the rate of component release from fermented dairy products from Lactobacillus helveticus
and cheese matrix disintegration behaviour during in vitro (Nakamura et al. 1995). In cheeses, when L. helveticus was
digestion is significantly influenced by the level of matura- included in starter culture (Meyer et al. 2009) VPP and IPP
tion of cheese. Fang et al. (2016b) found a positive effect levels were notably high. While levels of VPP and IPP dif-
of proteolysis on disintegration of cheese during in vitro fer across cheese varieties and starter cultures and peak at
digestion. Similarly, Lamothe et al. (2012) found a rapid different maturation points in some cheeses compared to
disintegration of aged Cheddar cheese in the gastric phase others, generally the peptide levels increase with maturation,
as compared to mild Cheddar cheese; moreover, the free oil with peak activity in cheese observed at 4–7 months in age
released from aged Cheddar was higher than the mild Ched- (Meyer et al. 2009).
dar cheese at the end of gastric phase. In another study,
Asensio-Grau et al. (2019) observed a higher extent of
LIPIDS
lipolysis in matured cheeses (ripened for 240 days) as com-
pared to mild cheeses (ripened for 30 days) during in vitro In milk, lipid fractions are dispersed in the form of fat glob-
intestinal digestion. Rapid disintegration, and higher extent ules ranging from 0.2 to 15 lm with an average diameter of
of free oil release and lipolysis of matured cheese have been 4 lm (Lopez 2005). The milk lipid fraction is mainly com-
attributed to weakening of protein network due to age-re- posed of triacylglycerols (representing ~98 % of total
lated changes within the cheese matrix, primarily proteolysis lipids), with the remainder comprising di- and mono-acyl-
and solubilisation of colloidal calcium, which facilitates the glycerols, phospholipids, cholesterol, free fatty acids and
release of fat globules/pools (entrapped within the protein other lipophilic molecules (e.g. carotenoids and vitamins)
network) to the digestion medium and subsequently increase (Alothman et al. 2019). Milk fat contains more than 400
their accessibility to lipases (Lamothe et al. 2012; Asensio- different fatty acids (FA), the levels and composition of
Grau et al. 2019). which are influenced by a number of factors, such as stage
Overall, age-related changes within the cheese matrix of lactation, breed of cow, genetics and diet composition
seem to have a great influence on the digestion behaviour (Lindmark M ansson 2008). As stated above, the major pro-
of cheese. However, to date, little information is available portion of FA found in milk are saturated fatty acids
regarding the impact of maturation on the digestion beha- (SFAs), accounting for 60%–70% of the total milk FA, fol-
viour of different cheese varieties. A better understanding of lowed by monounsaturated fatty acids (MUFA) and polyun-
the roles of maturation on digestion behaviour of cheese saturated fatty acids (PUFA) (Jensen 2002; Alothman et al.
may contribute to establish dietary recommendations for 2019). The fat globules are surrounded by a thin membrane,
general population or specific population groups. For exam- called the milk fat globule membrane (MFGM), which pre-
ple, easily digestible matured cheeses may be more suitable vents the globules from coalescence and enzymatic degrada-
for those individuals who have diseases, such as cystic tion (Dewettinck et al. 2008).
fibrosis, and other pancreatic deficiencies (Asensio-Grau A number of studies have investigated the in vitro diges-
et al. 2019). tion behaviour of fat in different cheese varieties. In cheese,
Maturation may also impact the biological activity of fat globules/pools are entrapped within the protein network
cheeses. The proteolytic changes that occur with ageing can which has to be disintegrated to release fat globules/pools
result in the release of latent bioactivity of the cheese pro- from the cheese matrix to the digestion medium, where the
teins. Bioactive peptides, typically 2–20 amino acids in lipolytic enzymes carry out lipolysis. In vitro studies have
length, have been identified from a wide range of cheeses reported that the release of fat from the cheese matrix and
including Mozzarella, Parmesan, goat’s cheese, Gouda and the extent of lipolysis are closely related to the extent of the
Cheddar (for an extensive review please refer to the review cheese matrix disintegration (Lamothe et al. 2012; Guinot
by Santiago-L
opez et al. 2018). Antioxidant, anti-microbial et al. 2019; Calvo-Lerma et al. 2020).
and angiotensin converting enzyme (ACE)-inhibitory activ- Moreover, cheese manufacturing processes (e.g. scalding,
ity (antihypertensive) are the most commonly reported func- cheddaring, hot water stretching and pressing) greatly affect
tionalities in cheese (Santiago-L
opez et al. 2018). the structure of the native milk fat globules and their mem-
Antihypertensive peptides in particular have been observed brane. In cheese, milk fat can exist as individual fat glob-
in many cheese varieties and have been used from food ules, aggregated, coalesced (spherical but larger than typical
sources, to successfully lower blood pressure (Martinez- milk fat globules), elongated (especially in pasta-filata
Maqueda et al. 2012). Gamalost, a Norwegian cheese, has cheese types) or even nonglobular forms (Lamichhane et al.
also shown some potential for lowering of blood pressure in 2018b). Such cheese manufacture induced modifications of
human feeding studies when consumed simply as cheese the fat globules/pools are likely to influence digestion beha-
(Nilsen et al. 2016). Two well-recognised lactotripeptides viour.

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Although homogenisation of milk is not common for the G
omez-Cort
es et al. 2018). Some fatty acids found in milk,
manufacture of most natural cheeses, homogenisation of including butyric acid (4:0), conjugated linoleic acid (CLA)
milk and cream is practised for cream cheese manufacture as well as branched-chain FA, have been reported to have
and for some blue mould (to increase lipolysis) and of a positive health outcomes, such as maintenance of gut micro-
portion of the milk for reduced-fat cheeses (to improve tex- biota, weight control, gut health at birth and the prevention
tural properties). In raw milk, fat globule size commonly of chronic inflammatory diseases (Bruen et al. 2017;
ranges from  0.2–15 lm, and homogenisation decreases G
omez-Cort
es et al. 2018). Moreover, the components of
fat globule size (0.2–0.5 µm of volume mean diameter; MFGM are reported to have several health promoting
Lopez 2005) and greatly increases the surface area of fat effects, including reducing the risk of cardiovascular disease
globules, but also changes the surface composition of the (CVD), anti-inflammatory activity, cholesterol-lowering and
fat globules, through adsorption of milk casein micelles and anticarcinogenic activity (Spitsberg 2005; Dewettinck et al.
whey proteins (Kelly et al. 2008). For this purpose, two- 2008; Lordan et al. 2017).
stage valve homogenisers are commonly used, which oper- Dairy fat also contains significant levels of lauric acid
ate at pressure of  20 MPa (Kelly et al. 2008). Moreover, (C12:0), myristic acid (C14:0) and palmitic acid (C16:0).
microstructural analysis of the milk-based chymosin gels Intake of these saturated fatty acids in isolation has been
revealed that the milk fat globules are embedded in the associated in the past with increased blood LDL-cholesterol
casein matrix of the gel made from homogenised milk, sug- levels, increasing CVD risk. However, recent studies have
gesting that the homogenised fat globules interact with the suggested that food/dairy matrix in which these fatty acids
casein matrix of the gel, whereas the fat globules in gel are contained may influence health outcomes (Thorning
from nonhomogenised milk were weakly held or entrapped et al. 2017; Feeney et al. 2018), with strong evidence from
within the casein matrix of the gel (Figure 3; Ong et al. dietary intervention trials for cheese as having either neutral
2010; Lamichhane et al. 2020). or beneficial CVD impact (Timon et al. 2020). It is hypoth-
Cheeses made from homogenised milk showed different esised that other components/nutrients present in the cheese
fat digestion behaviour as compared to cheeses made from matrix, such as protein, peptides, phospholipids derived
nonhomogenised milk. For example, Lamothe et al. (2017) from milk fat globule membrane, calcium and phosphorous,
observed a faster release of free fatty acids from cheese are likely to influence the absorption of SFAs in the human
made from homogenised milk than from nonhomogenised body (Thorning et al. 2017).
milk during in vitro intestinal digestion and suggested that In addition to the impact on texture and rheological
the degree of lipid distribution within the cheese matrix is a properties of the cheese matrix, high levels of calcium
more important factor than the breakdown of the matrix for also appear to impact the absorption of the fat following
modulation of lipid digestion kinetics. consumption. A 2009 meta-analysis of RCTs examining
The health benefits of milk constituents, including some calcium intake (from dairy and nondairy sources) and fat
fatty acids and components of MFGM, have been exten- excretion observed that higher dairy calcium intakes
sively reviewed (Spitsberg 2005; Dewettinck et al. 2008; specifically were associated with higher fat excretion, and

(a) (b)

Figure 3 Microstructure of rennet induced gel made from (a) non-homogenized and (b) homogenized milk. Reprint from Ong et al. (2010). The
protein phase appears green while the fat phase appears red.

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calculated that 1241 mg of dairy Ca resulted in an average process. Buttermilk powder (BMP) is the dehydrated form,
additional excretion of 5.2 g of fat per day (Christensen obtained from BM (Vanderghem et al. 2010; Hickey et al.
et al. 2009). Hjerpsted et al. (2011) in a crossover inter- 2017), and both these BM and BMP are used in the indus-
vention study using dairy fat from cheese vs butter where try to add MGFM-derived phospholipids to various dairy
13% of energy was replaced noted that in the cheese products, such as cheese, yoghurt and infant milk formulas,
group (with a higher Ca intake vs the butter group), the enhancing both the sensory and nutritional properties
LDL-c levels were reduced vs the butter group after (Romeih et al. 2014; Hickey et al. 2017; Lopez et al. 2017;
6 weeks. They observed that the faecal fat excretion was Hickey et al. 2018).
11.6% higher in the cheese period, but this was not statis- The content of the plasma phospholipids within MFGM,
tically significant. The study assigned volunteers to low, and particularly sphingomyelin, may have health implica-
medium and high energy diets depending on their individ- tions, specifically on aspects of cholesterol metabolism.
ual requirements, with the medium diet providing approx. Rosqvist et al. (2015) tested the impact of dairy fat con-
1200 mg of dairy Ca per day in the cheese period, and sumption with and without MGFM on these aspects, with
0 mg in the butter period. A later study by Soerensen scones made using dairy fat in the form of cream (i.e. with
et al. (2014) using milk and cheese delivering similar MFGM; Figure 4a), or with butteroil (no MFGM; Fig-
amounts of dairy Ca (1700 mg per day) vs the control ure 4b). Participants consumed 40 g of dairy fat in either of
(500 mg per day) day each resulted in a significant these two forms, in a randomised, parallel-arm study design.
increase in the faecal fat compared to the control (Soeren- Following 8 weeks of daily consumption, the butteroil
sen et al. 2014). It is reported that calcium in cheese can group (no MGFM), differences were observed in both LDL-
produce insoluble calcium soaps with long-chain fatty and total cholesterol, both being significantly higher in the
acids at intestinal pH conditions (Jenkins and Palmquist, butteroil group compared to the cream. Additionally, there
1982; Ayala-Bribiesca et al. 2017), limiting its absorption were differences observed in expression levels for 19 genes,
in the human body, and this could be one of the possible some of which were associated with cholesterol metabolism
reasons for higher faecal excretions of fatty acids in and fatty acid synthesis. The expression levels were greater
humans who consumed a high-calcium diet than a low- in the butteroil group and reduced in the MGFM group fol-
calcium diet (Thorning et al. 2017). More research is lowing the 8-week intervention, suggesting a phospholipid-
needed to better understand the impact of calcium con- induced mechanism. More recently, Vors et al. (2019)
sumed in different dairy matrices on faecal fat and blood added milk polar lipids (PL) to cream cheese in a dose-de-
lipid profiles. pendent manner up (0, 3 and 5 g per day) in a study of 58
As mentioned earlier, the fat globules are surrounded by a post menopausal females. The authors observed that the PL
thin membrane, called the MFGM. Buttermilk (BM) is a increased total lipid beta-oxidation in the study, and reduced
particularly rich source of MFGM, being the serum fraction a range of markers including the ratios of total-c/HDL-c,
released during the churning of cream in the butter-making and of ApoB /ApoA1. 5 g/day of PL reduced LDL-c levels

(a) (b)

Figure 4 Confocal laser scanning microscopy micrographs of (a) milk fat globules from whipping cream (40% fat) and (b) milk fat globules in an
emulsion made from butter oil, purified water, and sodium dodecyl sulfate (15% fat); fat appears red while the milk fat globule membrane appears
green. Adapted from Rosqvist et al. (2015).

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by 8.7%. They also observed reduced intestinal absorption and maturation regime, among many other factors. As a
of cholesterol, and greater levels of excreted sphingomyelin, concentrated source of saturated fats, traditionally cheese
although there was no difference in the profile of excreted has been perceived as an unhealthy food and may be
short-chain fatty acids. actively avoided in the diet, in the believe that cheese con-
sumption may result in negative health outcomes, particu-
larly on CVD risk. However, recent studies show that foods
PREPARATION BEFORE CONSUMPTION
are more than the sum of their nutrients. This is particularly
(COOKING/BAKING/GRILLING)
true for cheese, where growing evidence suggests that the
Cheese is consumed in both unheated (e.g. mixed with nutrients present appear to work in concert to reduce mark-
salad) and heated form (e.g. toppings on pizzas and ers of CVD risk compared to other dairy products when
lasagne). The application of heat results in various physico- matched for fat content. However, a detailed mechanistic
chemical changes in the structural components of cheese, understanding on how the cheese matrix influences health
such as: (i) increase in proportion of liquid fat, aggregation outcomes is still lacking.
and coalescence of fat and subsequent formation of free oil; In particular, it may be necessary not just to consider
(ii) contraction of casein networks due to temperature-in- cheeses as a distinct product group when considering health
duced increases in the strength of hydrophobic interactions impacts, but rather as a diverse group of products with
and simultaneous release of moisture; and (iii) evaporation potentially different digestive traits and biological and health
of moisture from the cheese matrices (Guinee 2016; properties. For example, whether cheeses are predominantly
Lamichhane et al. 2018b). Such heat-induced changes on casein or whey derived, whether a chymosin or acid gela-
the structural components of cheese result in melting, flow- tion and/or high heat treatments have been used will impact
ing and softening of cheese. Although most mature chy- significantly on the in vivo digestion patterns of the protein
mosin-coagulated cheeses flow on heating, some cheese phase, while factors such as use or otherwise of homogeni-
types, such as Paneer, Queso blanco and Halloumi, greatly sation or emulsification will impact on fat digestion. Further
resist flow on heating, frying and baking, due to their speci- research should employ both in vitro model digestion sys-
fic manufacturing processes (Guinee 2016). In summary, the tems and in vivo clinical trials in this area.
cheese matrix (structural) properties change substantially on Similarly, there is no consensus on which cheese con-
heating. stituents and interactions between constituents are key to the
Although the digestion behaviour of unheated cheese has cheese matrix effect. For example, some studies suggest cal-
been studied frequently, surprisingly, no published data are cium soap formation for reduced cholesterol absorption,
available on the digestion behaviour of heated cheese. It is while others have focused on binding of cholesterol with
expected that heat-induced modification of cheese structure sphingomyelin. These mechanisms in particular require elu-
may impact its digestion behaviour. Thorning et al. (2017) cidation, again using in vitro model digestion systems and/
point out that it is not yet known how melting will impact or in vivo clinical trials where appropriate.
the overall cheese matrix structure, and human intervention Finally, other factors meriting further investigation, which
studies using cheese have specifically noted that volunteers to date have received less attention in this context, include
consumed cheese that has not been melted, due to this the role of herd diet (particularly pasture fed vs total mixed
unknown factor (e.g. Feeney et al. 2018). ration) on the cheese fatty acid profile and the mode of
Several studies have reported a significant impact of cook- cheese consumption. The increasing consumption of cheese
ing processes on the in vitro digestion behaviour of food in a heated format (food service applications, pizza, toasted
products. For example, Kong et al. (2013) observed a posi- sandwiches, etc.) using different heating methods, requires
tive impact of different heating/cooking processes (i.e. fry- specific research to determine whether the resultant physico-
ing, roasting and boiling) on the gastric disintegration chemical changes to the cheese matrix influence its diges-
behaviour of peanuts as compared to raw peanuts. Kaur tion and ultimately, its health effects.
et al. (2014) reported that the prolonged cooking of beef Overall, an enhanced knowledge of the cheese matrix
meat at 100 °C reduced the protein digestibility. A better from both physico-chemical and biological perspectives may
understanding of the role of cooking processes on the diges- contribute to the engineering of ‘healthier’ food structures in
tion behaviour of cheese may help to select cooking pro- the future.
cesses to attain specific digestive outcomes.
ACKNOWLEDGEMENTS
CONCLUSIONS
The authors wish to acknowledge the Food for Health Ireland
Cheeses are a very diverse category of dairy product, the project, funded by Enterprise Ireland, grant number
composition and structure of which can vary greatly depen- TC20180025, and a further significant in-kind contribution by
dent on type, milk pretreatment and manufacture process, author Lamichhane.

© 2021 The Authors. International Journal of Dairy Technology published by John Wiley & Sons Ltd on behalf of Society of Dairy Technology 11
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AUTHOR CONTRIBUTION Bruen R, Fitzsimons S and Belton O (2017) Atheroprotective effects of

conjugated linoleic acid. British Journal of Clinical Pharmacology
Emma Feeney: Conceptualization, Investigation, Methodol- 83 46–53.
ogy, Project administration, Writing-original draft, Writing- Calvo-Lerma J, Asensio-Grau A, Heredia A and Andr
es A (2020) Les-
review & editing. Prabin Lamichhane: Investigation, Writ- sons learnt from MyCyFAPP Project: Effect of cystic fibrosis factors
ing-original draft, Methodology, Writing-review & editing. and inherent-to-food properties on lipid digestion in foods. Food
Jeremiah J Sheehan: Conceptualization, Investigation, Research International 133 109198.
Methodology, Project administration, Writing-original draft, Calzada J, del Olmo A, Picon A, Gaya P and Nu~ nez M (2014) Effect of
Writing-review & editing. high-pressure-processing on the microbiology, proteolysis, texture
and flavour of Brie cheese during ripening and refrigerated storage.
DATA AVAILABILITY STATEMENT International Dairy Journal 37 64–73.
Data sharing is not applicable to this article as no new data Chen J (2009) Food oral processing—A review. Food Hydrocolloids 23
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Chen M, Pan A, Malik V S and Hu F B (2012) Effects of dairy intake
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