Authentication of Meat and Meat Products

Meat Science 86 (2010) 577587
Contents lists available at ScienceDirect
Meat Science
j o u r n a l h o m e p a g e : w w w. e l s ev i e r. c o m / l o c a t e / m e a t s c i
Review
Authentication of meat and meat products

N.Z. Ballin
Department of Food Chemistry, Regional Veterinary and Food Control Authority, Danish Veterinary and Food Administration, Soendervang 4, DK-4100 Ringsted, Denmark
Department of Food Science, Faculty of Life Sciences, University of Copenhagen, Rolighedsvej 30, DK-1958 Frederiksberg C, Denmark
a r t i c l e
i n f o
Article history:
Received 17 February 2010
received in revised form 30 May 2010
accepted 3 June 2010
Keywords:
Authentication
Authenticity
Fraud
Meat
Meat fraud
Meat products
Mislabelling
a b s t r a c t
In recent years, interest in meat authenticity has increased. Many consumers are concerned about the meat they
eat and accurate labelling is important to inform consumer choice. Authentication methods can be categorised
into the areas where fraud is most likely to occur: meat origin, meat substitution, meat processing treatment and
non-meat ingredient addition. Within each area the possibilities for fraud can be subcategorised as follows: meat
originsex, meat cuts, breed, feed intake, slaughter age, wild versus farmed meat, organic versus conventional
meat, and geographic origin; meat substitutionmeat species, fat, and protein; meat processing treatment
irradiation, fresh versus thawed meat and meat preparation; non-meat ingredient additionadditives and water.
Analytical methods used in authentication are as diverse as the authentication problems, and include a diverse
range of equipment and techniques. This review is intended to provide an overview of the possible analytical
methods available for meat and meat products authentication. In areas where no authentication methods have
been published, possible strategies are suggested.
2010 The American Meat Science Association. Published by Elsevier Ltd. All rights reserved.
Contents
1.
2.
Introduction . . . . . . . . . . . . . . . . . .
Identication of meat origin . . . . . . . . . .
2.1.
Sex . . . . . . . . . . . . . . . . . .
2.2.
Meat cuts . . . . . . . . . . . . . . .
2.3.
Breed . . . . . . . . . . . . . . . . .
2.4.
Feed intake . . . . . . . . . . . . . . .
2.5.
Slaughter age . . . . . . . . . . . . . .
2.6.
Wild versus farmed meat . . . . . . . .
2.7.
Organic versus conventional meat . . . .
2.8.
Geographic origin . . . . . . . . . . . .
3.
Identication of meat substitution. . . . . . . .
3.1.
Meat (species and tissue) . . . . . . . .
3.2.
Fat (vegetable and animal fat) . . . . . .
3.3.
Protein (vegetable protein, animal protein,
4.
Identication of meat processing treatment . . .
4.1.
Irradiation . . . . . . . . . . . . . . .
4.2.
Fresh versus thawed meat . . . . . . . .
4.3.
Meat preparation (baking, cooking etc.) .
5.
Identication of non-meat ingredient additions .
5.1.
Additives . . . . . . . . . . . . . . . .
5.2.
Water . . . . . . . . . . . . . . . . .
Acknowledgements . . . . . . . . . . . . . . . . .
References . . . . . . . . . . . . . . . . . . . . .
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Department of Food Chemistry, Regional Veterinary and Food Control Authority, Danish Veterinary and Food Administration, Soendervang 4, DK-4100 Ringsted, Denmark.
Tel.: +45 2682 8561; fax: +45 7227 6000.
E-mail addresses: nixb@fvst.dk, nicolaiba@hotmail.com.
0309-1740/$ see front matter 2010 The American Meat Science Association. Published by Elsevier Ltd. All rights reserved.
doi:10.1016/j.meatsci.2010.06.001
578
N.Z. Ballin / Meat Science 86 (2010) 577587
1. Introduction
Today, many consumers are concerned about the meat they eat,
and accurate labelling is important to inform consumer choice. The
choice of one product over another can reect aspects of lifestyle (e.g.
vegetarianism and organic food), religion (e.g. absence of pork from
some diets), diet and health concerns (e.g. absence of allergens). In
addition, accurate labelling is important to support fair-trade.
Additional descriptive label information can be added as a consequence of branding, product marketing purposes and regulations.
While regulations enshrined in national and international law
underpin mandatory label information, unfortunately, regulations
are not sufcient to prevent food fraud. To ensure adherence to
regulations, and to enforce punitive measures when needed, robust
analytical tests are required. The prevalence of meat fraud today is
difcult to measure. However, the examples of mislabelling and its
abundance in meat and meat products illustrated in Table 1 make
analytical authentication highly relevant.
This review provides an overview of the possible analytical methods
used for authentication of meat and meat products and identify areas of
future research needs. The analytical methods include polymerase chain
reaction, chromatography, mass spectrometry, microscopy, spectroscopy, electronic spin resonance, and enzymatic assays. An overview of the
analytical methods is provided in Tables 24. In the few cases where no
authentication methods have been published, possible analytical
strategies are suggested. For simplicity, authentication problems with
respect to meat and meat products are grouped into four major
categories: meat origin, meat substitution, meat processing treatment
and non-meat ingredient addition. Within each area the possibilities for
fraud can be subcategorised as follows: meat originsex, meat cuts,
breed, feed intake, slaughter age, wild versus farmed meat, organic
versus conventional meat, and geographic origin; meat substitution
meat species, fat, and protein; meat processing treatmentirradiation,
fresh versus thawed meat and meat preparation; non-meat ingredient
additionadditives and water. Fig. 1 presents an overview of the
different categories and subcategories.
Analytical authentication of food products often requires sample
preparation such as extraction of proteins, DNA, and organic
compounds. In this review, preparation prior to actual analysis is
excluded for simplicity. However, it is often an important step that
must be carefully considered.
2. Identication of meat origin
2.1. Sex
Analysis of sex-specic hormones is a conventional method used
to determine the sex of meat. The level of sex specic hormones varies
not only among individuals, but also within a given individual (Zeleny
& Schimmel, 2002). It is therefore challenging to establish sex
determination methods based on hormone analysis alone. However,
gas chromatography-mass spectrometry (GC-MS) (Hartwig et al.,

1997), high performance liquid chromatography-mass spectrometry/
mass spectrometry (HPLC-MS/MS) (Draisci et al., 2000), and enzymelinked immuno sorbent assays (ELISA) (Simontacchi et al., 1999) have
been found effective to measure bovine sex hormones and thereby
determine the sex. Nevertheless, the most common methodology
used to determine the sex of animals is unquestionably traditional
polymerase chain reaction (PCR) that involves gel electrophoresis of
DNA amplicons. An overview of the analytical methods is provided in
Table 2.
DNA regions that differ between male and female individuals are
essential features in PCR sex determination. Such DNA regions include
the zinc nger genes (ZFX and ZFY), the sex determination region of
the Y chromosomal gene (SRY), and the tooth enamel amelogenin
gene (AMELX and AMELY). Traditional PCR amplication of ZFX and
ZFY genes have been used to determine sex in cattle (Aasen &
Medrano, 1990; Kirkpatrick & Monson, 1993; Zinovieva et al., 1995),
sheep (Aasen & Medrano, 1990), pig (Lockley et al., 1997), and goat
(Aasen & Medrano, 1990). PCR amplication of the Y chromosomal
SRY gene has been reported to be successful in sex determination of
cattle (Lu et al., 2007) and buffalo (Fu et al., 2007), while the AMELX
and AMELY genes have been used to determine sex in cattle (Chen et
al., 1999; Ennis & Gallagher, 1994) and pig (Fontanesi et al., 2008).
Different bovine sex determination methodologies have been comprehensively described (Zeleny & Schimmel, 2002) and evaluated
(Zeleny, Bernreuther, Schimmel, & Pauwels, 2002).
Currently the focus seems to have shifted from traditional PCR
towards real time PCR. Real time PCR based TaqMan technology
directed toward the specic Y chromosomal SRY and the X chromosomal proteolipid protein gene has been used in sex determination of
cattle (Parati et al., 2006). Furthermore, the real time PCR based SYBR
Green technology combined with the melting curve analysis of
amplied AMELX and AMELY genes has been used in bovine sexing
(Ballin & Madsen, 2007), and a similar assay that amplies genes
encoding chromodomain-helicase-DNA binding protein (CHD) was
described in avian sexing (Chang et al., 2008).
2.2. Meat cuts
Staff educated with this purpose in mind can easily differentiate
between primary beef cuts such as chuck, brisket, sirloin, and shank
through visual inspection. Steaks, as a result of secondary meat cuts,
are more difcult to visually authenticate than primary cuts.
Authentication is further complicated since the names of primary
and secondary meat cuts vary across countries. In addition, meat cuts
vary considerably over time because of various consumer demands
and market trends. These factors make it difcult to establish visual
objective criteria for authentication of specic meat cuts. To my
knowledge, no analytical authentication methods have yet been
published. However, the fact that some chemical constituents in meat
differ between different meat cuts might aid in authentication. The
Table 1
Examples of results from authenticity investigations of meat products.
Investigated product
Country of investigation
Authentication problem
Percentage of mislabelling cases

(number of analysed samples)
Reference
Hamburgers
Brazil
Undeclared soy protein
30.8% (39)
Hamburgers
Mexico
Undeclared animal species
39% (23)
Sausages
Meat products
Mexico
United States of America

Meat products
Meat products
Meat products
Turkey
Switzerland
United Kingdom

Thawed meat declared as fresh
Thawed meat declared as fresh
29% (17)
15.9% raw samples
22.9% cooked samples
22% (100)
15% (43)
8% (534)
Macedo-Silva, Shimokomaki, Vaz, Yamamoto,

and Tenuta-Filho (2001)
Flores-Munguia, Bermudez-Almada, and
Vzquez-Moreno (2000)
Flores-Munguia et al. (2000)
g (902)
Hsieh, Woodward, and Ho (1995)

Ayaz, Ayaz, and Erol (2006)
Anon (2001)
Anon (1996)

Table 2
Examples of analytical techniques applicable in authentication of sex, meat cuts, breeds,
feed intake, slaughter age, wild vs. farmed meat, organic vs. conventional meat, and
geographic origin in meat and meat products.
Authenticity
problem
Meat origin
Sex
Analytical
technique*
Reference
GC-MS
HPLC-MS-MS
Hartwig, Hartmann, and Steinhart (1997)

Draisci, Palleschi, Ferretti, Lucentini, and
Cammarata (2000)
Simontacchi, Marinelli, Gabai, Bono, and
Angeletti (1999)
Aasen & Medrano, 1990; Chen et al.,
1999; Ennis & Gallagher, 1994; Fontanesi,
Scotti, & Russo, 2008; Fu et al., 2007;
Kirkpatrick & Monson, 1993; Lockley, Bruce,
Franklin, & Bardsley, 1997; Lu, Rawlings,
Zhao, & Wang, 2007; Zinovieva, Palma,
Mller, & Brem, 1995
Chang et al., 2008; Ballin & Madsen, 2007;
Parati, Bongioni, Aleandri, & Galli, 2006
ELISA
PCR
Meat cuts
Breed
Real time
PCR
Visual
inspection
SNP
AFLP
Genotyping
Feed intake
RAPD
Infrared
reectance
spectroscopy
HPLC-UV
GC-FID
Head space
GC-MS
Spectroscopy
Dalvit et al., 2008; Sasazaki et al., 2004

Alves, Castellanos, Ovilo, Sili, and Rodrguez
(2002)
Garcia et al., 2006; Vega-Pla, Martinez,
Cabello, Rodriguez-Gallardo, & Delgado, 2003
Martinez and Yman (1998)
Alomar, Gallo, Castaeda, and Fuchslocher
(2003)
Dunne, O'Mara, Monahan, & Moloney, 2006;
Prache, Priolo, & Grolier, 2003
Duckett, Wagner, Yates, Dolezal, & May, 1993;
French et al., 2000
Vasta, Ratel, and Engel (2007)
Priolo, Prache, Micol, & Agabriel, 2002;
Moreno et al., 2006
Slaughter age
See text
regarding feed
intake.
Wild vs.
GC
Almeida & Franco, 2007; Karapanagiotidis,
Farmed meat
Bell, Little, Yakupitiyage, & Rakshit, 2006
Multi element Thomas et al. (2008)
isotopic
analysis
Organic vs.
Fluorescence Kelly, Tarbin, Ashwin, and Sharman
Conventional meat microscopy
(2006)
GC
Angood et al., 2008; Castellini, Mugnai,
& Dal Bosco, 2002; Husak, Sebranek, &
Bregendahl, 2008; Kim et al., 2009
Schmidt et al. (2005)
Stable ratio
mass
spectrometry
Geographic origin
MS
Franke, Koslitz, et al., 2008; Franke,
Haldimann, et al., 2008; Nakashita et al.,
2008; Schmidt et al., 2005
SNP
Negrini et al., 2008; Sasazaki, Mutoh,
Tsurifune, & Mannen, 2007
*Abbreviations used: AFLP, amplied fragment length polymorphism; ELISA, enzymelinked immuno sorbent assay; FID, ame ionization detector; GC, gas chromatography;
HPLC, high performance liquid chromatography; MS, mass spectrometry; PCR,
polymerase chain reaction; RAPD, random amplied polymorphic DNA; SNP, single
nucleotide polymorphism; UV, ultraviolet.
difference in the amounts of collagen present in different meat cuts is

a case-in-point.
2.3. Breed
Authentication of breed is primarily based on PCR and a
subsequent analysis of amplicons. An overview of the analytical
methods is provided in Table 2. Microsatellite DNA markers and a
579
Bayesian statistical model identied the Italian cattle breeds Chianina,

Marchigiana, Romagnola, and Piemontese (Dalvit et al., 2008).
Another study that used single nucleotide polymorphism (SNP)
markers directed toward the SRY gene and the mitochondrial NADH
dehydrogenase subunit 5 (ND5) gene identied the cattle breeds
Holstein and Japanese Black (Sasazaki et al., 2004). Amplied
fragment length polymorphism (AFLP) (Alves et al., 2002) and
multilocus genotyping of repetitive sequences (Garcia et al., 2006;
Vega-Pla et al., 2003) successfully distinguished Duroc and the
crossbred IberianDuroc from the purebred Iberian pig. Interestingly,
Iberian pigs are used for the dry-cured Iberian ham, which is a Spanish
speciality. This ham must consist of 100% of the Iberian genome or a
50:50% mixture of the Iberian and Duroc genome. Random amplied
polymorphic DNA (RAPD) has successfully differentiated between
horse breeds (Martinez & Yman, 1998). With regard to RAPD, it is
relevant to consider that electrophoretic proles from mixtures of
breeds might be difcult to interpret, specically for species that can
interbreed. For example, the RAPD pattern of a hybrid is similar to the
5050% mixture of the RAPD patterns of the parental species
(Martinez & Yman, 1998). In addition, a RAPD assay designed with
primers toward mitochondrial DNA might fail in the identication of
hybrids as mitochondrial DNA is primarily maternally inherited
(Adam, 2000; Gyllensten, Wharton, Josefsson, & Wilson, 1991). PCR
and a subsequent amplicon analysis is not the only possibility in
identication of breed and, for example, near infrared reectance
spectroscopy (NIRS) was used to study Friesian and Hereford beef;
spectral differences were observed, especially in the region between
1449 and 1974 nm (Alomar et al., 2003). A larger data set should,
however, be obtained to build a model capable of discriminating
between Friesian and Hereford beef samples.
2.4. Feed intake
There are various means of tracing animal feed intake. Traceability
is possible because different chemical constituents are present in
feeds such as milk, pasture, hay, maize, and concentrate (mixed dried
constituents), which upon consumption, shows up as different
chemical constituents or metabolised forms in the animals' blood
and fat. An overview of the analytical methods is provided in Table 2.
A group of chemical constituents that can be used as authentication
markers is the carotenoids. The carotenoids, xanthophylls and
carotenes, are much more abundant in pasture when compared to
hay and concentrate (Prache et al., 2003). HPLC was used to measure
carotenoid content in sheep's blood (Prache et al., 2003) and in
heifer's fat, and the carotenoid content was dependent on whether
the diet was based on pasture or concentrate (Dunne et al., 2006).
Carotenoid differences also allowed reectance spectroscopy between
400 and 700 nm to distinguish between lambs fed on grass and lambs
fed on concentrate (Priolo et al., 2002). However, intrinsic factors such
as breed, gender, lactation, and rumen environment also affect the
carotenoid content (Dunne, Monahan, O'Mara, & Moloney, 2009) and
must be considered in the development of these methods.
The composition of fatty acids in meat fat is also dependent on an
animal's diet. Gas chromatographic studies describe a higher ratio of
polyunsaturated fatty acids to saturated fatty acids (Duckett et al.,
1993; French et al., 2000), and a lower ratio of n 6 to n 3
polyunsaturated fatty acids (French et al., 2000) in steers fed on grass
compared to steers fed on concentrate. A study on lambs found
differences in saturated fatty acids and mono- and polyunsaturated
fatty acid between lambs fed on milk versus a concentrate and hay
diet (Okeudo, Moss, & Chestnutt, 2004). Differences in saturated fatty
acids (C14:0, C16:0, C17:0, C20:2, and C23:0) and unsaturated fatty
acids (18:1, C18:2, C18:3, and C23:3) were also found in another
study that has successfully been used to discriminate between
weaned and unweaned calves in a multivariate discriminant
procedure (Moreno et al., 2006).
580
Furthermore, head space GC-MS was used to study volatile

compounds in lambs' fat, wherein 33 compounds were identied as
being relevant in identication of meat from pasture and concentrate
fed animals (Vasta et al., 2007). Another possibility is to investigate
the content of vitamins and terpenes that are found in higher
concentrations in meat from animals fed on grass compared to
animals fed on concentrate (Descalzo et al., 2005; Larick et al., 1987).
2.5. Slaughter age
Slaughter age is important as meat sold from young animals is
often more highly valued than meat from older animals. Examples are
the higher priced veal compared to beef and the higher priced lamb
compared to mutton. The denitions of veal and lamb meat compared
to beef and mutton meat differ among countries. This difference
complicates analytical testing. In the European Union, veal must
origin from an animal not older than 12 months. A limit of 12 months
is, however, impossible to measure with analytical methods. If the
veal label is also related to feed intake, such as the milk-fed veal calves
and the grain-fed calves, analytical testing becomes feasible as specic
constituents from the feed can be found in the meat (see previous
section Feed intake).
2.6. Wild versus farmed meat
Wild pig (Sus scrofa scrofa), for example, differs genetically from
domesticated pig (Sus scrofa domestica) hence differentiation between
wild and domesticated animals can in this case be done by (sub)
species identication, see section Meat (species and tissue). Game,
such as crocodile and ostrich, is farmed in some countries and though
fraud is a possibility, the scientic community has given this little
attention as this is not done on a large scale. In sh, gas chromatographic measurement of differences in fatty acid composition
(Almeida & Franco, 2007; Karapanagiotidis et al., 2006) in combination
with multi element isotopic analysis (Thomas et al., 2008) was used to
differentiate wild and farmed sh. This strategy might also be
applicable to meat.
2.7. Organic versus conventional meat
One restriction in production of organic meat is the use of
veterinary drugs and rules dictate how and when such drugs may
be used (Anon, 2008a). Fluorescence microscopy (Kelly et al., 2006) of
cross sectional bone cuts can estimate the number of tetracycline
doses administered to pig and chicken, and illegal serial or
prophylactic dosing of tetracyclines can help verify whether the
animal complies with the specied organic rules. This strategy is not
conclusive as absence of tetracyclines is not a guarantee of organically
produced meat.
One strategy is to explore the differences in animal fat from
organically and conventionally raised animals. A gas chromatographic
study of fatty acid methyl esters showed an increase in polyunsaturated fatty acids in organically produced lamb (Angood et al., 2008),
pig (Kim et al., 2009), and broiler (Castellini et al., 2002; Husak et al.,
2008) as compared to conventionally produced meats. However, fatty
acid composition is also dependent on specic dietary intake such as
pasture and concentrate (Prez-Palacios, Ruiz, Tejeda, & Antequera,
2009). Again, this strategy is not conclusive. A high level of
polyunsaturated fatty acids could result from the specic dietary
intake and not in from organically produced feed alone.
Another strategy could be analysis of isotopic composition. Stable
ratio mass spectrometry of carbon, nitrogen, and sulphur isotopes
succeeded in differentiating between organic and conventional Irish
beef (Schmidt et al., 2005). The differences in isotopic composition in
organic and conventional beef are partly due to the difference in the
feed intake (grass or concentrate) (Schmidt et al., 2005). However, the
higher content of 15N in conventional beef compared with organic

beef (Bahar et al., 2008) might be a result of the mineral fertilizers
applied to the soil where conventionally grown animals are fed
(Watzka, Buchgraber, & Wanek, 2006). An overview of the analytical
methods is provided in Table 2.
2.8. Geographic origin
Inductively coupled plasma mass spectroscopy (ICP-MS) of trace
elements and stable isotope ratios are predominantly used in
authentication of geographic origin, and an overview of the analytical
methods is provided in Table 2. The content of trace elements and
isotopes in animals depend on various factors such as feed intake,
drinking water, pollution, and soil composition, all of which depend
on geographic origin.
Franke, Haldimann, et al. (2008) tested fty elements and 75As,
23
Na, 85Rb, 77Se, 88Sr, and 205Tl for poultry meat and 75As, 10B, 137Ba,
42
Ca, 111Cd, 63Cu, 163Dy, 167Er, 57Fe, 7Li, 55Mn, 104Pd, 85Rb, 77Se, 88Sr,
128
Te, 203Tl, 238U, and 51V for dried beef, were shown to differ
signicantly between countries. Poultry discrimination was possible
among samples from Brazil, France, Germany, Hungary, and
Switzerland. In dried bovine meat, linear discriminant analysis was
applied to a number of elements and used to establish a classication
matrix. The classication matrix was used to predict geographic origin
of bovine meat from Australia, Austria, Switzerland, Canada, Brazil,
and the United States of America. Prediction of geographic origin was
performed with varying degrees of success, and the accuracy might be
improved by combining different approaches (Franke, Haldimann, et
al., 2008). The oxygen isotope ratio (18O/16O) in meat's water fraction
was used to differentiate between beef and chicken when geographic
conditions were clearly different (Australia vs. Europe vs. North
America) (Franke, Koslitz, et al., 2008). A combination of multi trace
elemental analysis (Franke, Haldimann, et al., 2008) and oxygen
isotope analysis (Franke, Koslitz, et al., 2008) would seem appropriate; however, a recent attempt did not improve correct geographic
classication compared to the individual methods (Franke, Hadorn,
Bosset, Gremaud, & Kreuzer, 2008). Measurement of carbon and
nitrogen isotopes has also shown potential in differentiating between
beef originating from Japan, Australia, and the United States of
America (Nakashita et al., 2008), as well as beef originating from
Europe and the United States of America (Schmidt et al., 2005).
Some cattle breeds are country specic and DNA analysis has been
used in indirect identication of geographic origin. Analysis of 24
breeds was used to build a Bayesian statistical model based on single
nucleotide polymorphisms (SNP) markers, and indirect differentiation between Italy, France, Spain, and the United Kingdom was
possible (Negrini et al., 2008). Another study, also using SNP markers,
differentiated between breeds from Japan and Australia (Sasazaki et
al., 2007). For species that have country specic breeds, this strategy
could be useful in combination with elemental analysis. However,
identication of breed alone is not evidence of geographic origin as
individual breeds can be raised in different countries. An overview of
European breeds and their origin is described elsewhere (Dalvit, De
Marchi, & Cassandro, 2007).
3. Identication of meat substitution
3.1. Meat (species and tissue)
Fraudulent substitution of meat can involve both species and
tissue. In species determination, analysis of DNA and protein are
common practices. Detection of protein has traditionally been the
most suitable method for animal species determination. Detection of
animal protein is covered later in the section Protein. An overview of
the analytical methods is provided in Table 3.
581
Table 3
Examples of analytical techniques applicable in authentication of meat, fat, and protein substitution in meat and meat products.
Authenticity problem
Analytical technique*
Reference
Meat substitution
Meat (species)
ELISA
Gonzalez-Cordova, Calderon de la Barca, Cota, & Vallejo-Cordoba, 1998; Koppelman,

Lakemond, Vlooswijk, & Hee, 2004
Ashoor, Monte, & Stiles, 1988; Ashoor & Osman, 1988; Chou et al., 2007
Skarpeid, Kvaal, and Hildrum (1998)
Vallejo-Cordoba and Cota-Rivas (1998)
Calvo, Rodellar, Zaragoza, & Osta, 2002; Colgan, O'Brien, Maher, Shilton, McDonnell,
& Ward, 2001; Hopwood, Fairbrother, Lockley, & Bardsley, 1999; Pascoal, Prado, Calo,
Cepeda, & Barros-Velazquez, 2005; Rodriguez et al., 2003; Sun & Lin, 2003
Brodmann & Moor, 2003; Chrisholm, Conyers, Booth, Lawley, & Hird, 2005; Fumiere,
Dubois, Baeten, von Holst, & Berben, 2006; Hird, Chisholm, & Brown, 2005; Laube,
Zagon, & Broll, 2007; Lopez-Andreo, Garrido-Pertierra, & Puyet, 2006; Martn et al.,
2009; Mendoza-Romero et al., 2004
Verkaar, Nijman, Boutaga, & Lenstra, 2002; Wolf , Rentsch, and Hubner (1999)
Calvo, Zaragoza, & Osta, 2001; Martinez & Yman, 1998; Rastogi et al., 2007
Bartlett & Davidson, 1992; Forrest & Carnegie, 1994; Iijima et al., 2006; Imaizumi,
Akutsu, Miyasaka, & Yoshino, 2007
Tartaglia et al. (1998)
Rastogi et al. (2004)
Al Jowder, Defernez, Kemsley, and Wilson (1999)
Chen, Hsieh, & Bridgman, 2002; Kim et al., 2004, 2005; Muldoon, Onisk, Brown, &
Stave, 2004
Colgrave, Allingham, and Jones (2008)
Nair, Kanfer, and Hoogmartens (2006)
Szucs, Sarvary, Cain, and Adany (2006)
Lembcke et al. (2005)
Precht (1992a,b)
Precht (1992a,b)
Castro, Garca, Rodrguez, Rodrguez, and Marina (2007)
Gonzalez-Cordova et al., 1998; Koppelman et al., 2004
Haasnoot and du Pre (2007)
Ashoor et al., 1988; Ashoor & Osman, 1988; Chou et al., 2007
Chen et al., 2002; Kim et al., 2004, 2005; Muldoon et al., 2004
Frick, Dubois, Chaubert, and Ampuero (2009)
Varelis and Jeskelis (2008)
Frick et al. (2009)
LC
Isoelectric focusing
Capillary gel electrophoresis
Traditional PCR
Real time PCR
RFLP
RAPD
Sequencing
Meat (tissue)
Fat (vegetable)
Fat (animal)
Protein (vegetable)
Protein (animal)
Protein (melamine and urea)
SSCA
CSGE
Mid-infrared spectroscopy
ELISA
LC-MS/MS
HPLC
GC-MS
APPI LC-MS/MS
GC-FID#
GC#
HPLC
ELISA
Microsphere-based ow cytometric immunoassay
LC
ELISA
Head space GC-MS
LC-MS/MS
Head space GC-MS
*Abbreviations used: APPI, atmospheric pressure photoionization; CSGE, conformation sensitive gel electrophoresis; ELISA, enzyme-linked immuno sorbent assay; FID, ame
ionization detector; GC, gas chromatography; HPLC, high performance liquid chromatography; LC, liquid chromatography; MS, mass spectrometry; PCR, polymerase chain reaction;
RAPD, random amplied polymorphic DNA; RFLP, restriction fragment length polymorphism. #These methods are described for dairy products but the same methodology might be
applicable to meat.
Rapidly evolving DNA based techniques have resulted in a change

in species determination from protein to DNA analysis. Degeneracy of
DNA has the advantage that differentiating among different animal
species can be done solely using DNA analysis. Furthermore, when
compared to proteins, DNA has a higher thermal stability, is present in

the majority of cells, and potentially enables identical information to
be obtained from the same animal regardless of tissue of origin
(Lockley & Bardsley, 2000). PCR is capable of amplifying very few
Fig. 1. Potential meat authenticity problems.
582
copies of DNA and the limit of detection is therefore often lower than
what is observed with protein based assays. Amplied PCR products
can either be visualised on a gel (end point PCR) or in real time PCR,
sometimes referred to as qPCR (quantitative PCR). Real time PCR data
can be collected at the early exponential phase of amplication, which
allows quantitative results to be obtained. In contrast, species
determination by end point PCR is only qualitative (Calvo et al.,
2002; Colgan et al., 2001; Hopwood et al., 1999; Pascoal et al., 2005) or
in some cases semi-quantitative (Rodriguez et al., 2003; Sun & Lin,
2003). Currently, real time PCR is generally the method of choice and a
number of qualitative (Brodmann & Moor, 2003; Fumiere et al., 2006;
Hird et al., 2005; Laube et al., 2007; Mendoza-Romero et al., 2004) and
quantitative (Chrisholm et al., 2005; Lopez-Andreo et al., 2006; Martn
et al., 2009) species determination methods have been published.
Other common PCR based techniques are restriction fragment
length polymorphism (RFLP) (Verkaar et al., 2002; Wolf et al., 1999)
and random amplied polymorphic DNA (RAPD). RAPD analysis takes
advantage of short arbitrary primers that are able to produce a range
of PCR products. RAPD is a powerful technique where little or no
information on the DNA sequence is available as long as reference
material for comparison is available. This technique is relevant to
differentiate not only between domestic animals (Calvo et al., 2001;
Martinez & Yman, 1998) but also among rare species (Rastogi et al.,
2007) since no prior knowledge of the DNA sequences is required. For
more information, the applications of RAPD analysis in livestock
species are reviewed elsewhere (Cushwa & Medrano, 1996).
Characterisation of animal species by sequencing depends on
availability of known sequences used for comparison. Much information is present in databases, such as the National Center for
Biotechnology Information, (http://www.ncbi.nlm.nih.gov/), which
contains a large number of sequences from common animal species,
breeds, and other genetic variations. Sequencing can therefore be
used to identify species in unknown samples, even if no reference
material is available. It only requires that the species have a unique
DNA sequence and that data are available for comparison. Sequenced
PCR amplicons and identication through database comparison has in
a number of cases established the identity of the species (Bartlett &
Davidson, 1992; Forrest & Carnegie, 1994; Iijima et al., 2006; Imaizumi
et al., 2007). In addition, single-strand conformational analysis (SSCA)
(Tartaglia et al., 1998) and conformation sensitive gel electrophoresis
(CSGE) (Rastogi et al., 2004) helps provide post-PCR speciation
evidence. Species determination with special focus on quantication
is reviewed elsewhere (Ballin, Vogensen, & Karlsson, 2009). General
DNA based authentication methods including species determination
have also been reviewed (Lockley & Bardsley, 2000; Mafra, Ferreira, &
Oliveira, 2008).
Besides substitution of one species with another, fraudulent
substitution of tissue with collagen and offal might also be protable
to the food industry. In collagen, the natural amino acid, 4hydroxyproline (hyp) is a major component and therefore useful in
determination of collagen content. Collagen contains about 8% hyp,
which is signicantly more than other proteins e.g. elastin contains
about 1% hyp (Etherington & Sims, 1981). Different amounts of hyp in
different meat parts can be used to calculate the content of collagen in
meat and meat products (Anon, 2008b). The EU reference method for
quantication of hyp (Anon, 1978a) is based on a simple spectroscopic method. Other more advanced methods are also available and
include chromatographic techniques such as LC-MS/MS (Colgrave et
al., 2008). Offal has successfully been discriminated from muscle
tissue with use of mid-infrared spectroscopy (Al Jowder et al., 1999).
Another way to discriminate between tissues is the use of bovine hcaldesmon, which is present in smooth muscles (Sobue, Tanaka,
Kanda, Ashino, & Kakiuchi, 1985) but absent in cardiac and skeletal
muscles (Bretscher & Lynch, 1985). This makes ELISA useful for tissue
differentiation (Chen et al., 2002; Kim et al., 2004, 2005; Muldoon et
al., 2004).
3.2. Fat (vegetable and animal fat)

In meat products, vegetable fat might fraudulently substitute
animal fat. However, vegetable fat often contains phytosterols such as
stigmasterol and -sitosterol, which are absent in animal fat. Thus, the
occurrence of stigmasterol and -sitosterol in meat products is
indicative of the presence of vegetable fat. Techniques, such as HPLC
(Nair et al., 2006), GC-MS (Szucs et al., 2006), and atmospheric
pressure photoionization (APPI) LC-MS/MS (Lembcke et al., 2005) can
be used to detect phytosterols in various matrices. It should be
stressed that the ratio between total vegetable fat and stigmasterol or
-sitosterol varies and quantitative determination of vegetable fat is
hence impractical.
In meat products, animal fat from one species might fraudulently
be used to substitute animal fat from another species. However, both
animal fat and vegetable fat contain species-specic relative amounts
of fatty acids (Precht, 1992a). Gas chromatographic-ame ionization
detection (GC-FID) methods based on these relative amounts of fatty
acids have so far been used to quantify foreign animal and vegetable fat
in milk (Precht, 1992b); the same principles might also be applicable
to meat products.
3.3. Protein (vegetable protein, animal protein, and organic compounds)
Vegetable protein such as the cheap and readily available soy is
probably one of the most commonly used proteins for fraudulent
substitution of animal protein. Soy protein can be detected by ELISA
methods (Gonzalez-Cordova et al., 1998; Koppelman et al., 2004). In
addition, commercial immunoassays for soy protein detection are also
available; these includeBioKits Soya Protein Assay (Tepnel,
Stamford, Conn) and Alert for Soy Flour Allergen (Neogen Corporation). Other proteins might also be used as adulterants and a triplex
immunoassay for simultaneous detection of pea, wheat, and soy
protein has been described for milk, and this method might be relevant
to meat and sausage as well (Haasnoot & du Pre, 2007). The triplex
immunoassay is based on microsphere-based ow cytometric immunoassay with MultiAnalyte Proling, which allows measurement of up
to 100 different bio molecular reactions (Haasnoot & du Pre, 2007).
Also HPLC has shown potential in determination of soy even in highly
processed meat mixtures (Castro et al., 2007). Analysis of soy protein
in meat products is reviewed elsewhere (Belloque, Garca, Torre, &
Marina, 2002; Koppelman et al., 2004).
Cheap animal protein might be fraudulently used to substitute more
expensive animal protein. Animal protein can be detected by ELISA
methods. Commercial ELISA methods are available and include Reveal
for Ruminant in MBM (Meat and Bone Meal; Neogen Corporation),
MELISA-Tek (ELISA Technologies), FeedChek (Strategic Diagnostics
Inc.), and the Tepnel BioSystems BioKit (Stamford, Conn.). All four kits
were evaluated with focus on selectivity, sensitivity, ruggedness, and
specicity but failed to meet acceptance criteria set up by the US Food
and Drug Administration's Centre for Veterinary Medicine Ofce of
Research, and hence were regarded inadequate for regulatory use
(Myers et al., 2007; Myers, Yancy, Farrell, Washington, & Frobish, 2005).
Hopefully, other ELISA methods will perform better in future evaluations. Normally, ELISA tests do not differentiate between proteins from
animal derived substances such as milk, whey, casein, cheese, eggs, and
meat. It is therefore difcult to identify whether animal protein has been
fraudulently substituted in meat products that contain labelled animal
derived substances other than meat. In this regard, one option would be
to develop tissue specic ELISA methods. Interestingly, bovine hcaldesmon present in smooth muscle (Sobue et al., 1985), is absent in
body uids, cardiac and skeletal muscles (Bretscher & Lynch, 1985).
Practically, distribution of h-caldesmon enables ELISA test methods to
differentiate between added bovine meat and other bovine derived
substances (Chen et al., 2002; Kim et al., 2004, 2005; Muldoon et al.,
2004). Detection of animal protein is not restricted to ELISA. Liquid
chromatographic (LC) methods that focus on protein proles for

qualitative detection of a variety of meats, which include beef, pork,
and chicken from raw and cooked products have been published
(Ashoor et al., 1988; Chou et al., 2007). Furthermore, protein proles
from LC analysis of unheated chicken and turkey mixtures can give
quantitative results (Ashoor et al., 1988; Ashoor & Osman, 1988).
Examination of the dipeptides, carnosine, anserine, and balenine can
qualitatively detect pork, beef, lamb, and chicken as long as different
species are not mixed (Aristoy & Toldra, 2004). Isoelectric focusing
(Skarpeid et al., 1998) and sodium dodecyl sulfate (SDS) capillary gel
electrophoresis (Vallejo-Cordoba & Cota-Rivas, 1998) of proteins also
successfully differentiates between beef, pork, and turkey in meat
mixtures.
Detection of vegetable and animal protein is dependent on the
nature of proteins. However, detection of protein might be impossible
if proteins are degraded or severely altered during the processing of
meat. In that case, species-specic methods such as PCR can assist in
identication of protein-adulterated meat. The advantage of DNA
based methods as compared to protein based methods is that DNA is
more stable as compared to protein under most conditions. However,
since the amount of residual DNA in a formulation varies from
detectable to non-detectable levels; PCR results should be considered
indicative of qualitative character.
In addition to substitution with vegetable and animal protein,
melamine and urea have been used in fraudulent substitution of
protein (Frick et al., 2009; Sivaraman, 2007). Melamine and urea are
effective adulterants as they contain a high level of nitrogen. Melamine
and urea contain 67% and 43% nitrogen, by mass, respectively,
compared to meat protein that contains about 16% nitrogen, by mass
(Benedict, 1987). Kjeldahl analysis (Wiles, Gray, & Kissling, 1998) or
combustion techniques such as Dumas are often used to measure the
total nitrogen content, which is then converted to protein content.
However, neither Kjeldahl nor Dumas differentiate between nitrogen
atoms derived from protein or chemical compounds. Thus, melamine
and urea can fraudulently substitute a fraction of the meat protein
without being detected through these analyses. Determination of the
triazides melamine and its hydrolysed cyanuric acid is primarily done
using chromatographic techniques. One study describes melamine
and cyanuric acid synthesized with stabile isotopes and used as
internal standards in liquid chromatographic tandem mass spectrometry, applicable to chicken, pork, catsh, and pet food (Varelis &
Jeskelis, 2008). Head space GC-MS (Frick et al., 2009) has been used to
study food ingredients adulterated with urea. An overview of the
analytical methods is provided in Table 3.
4. Identication of meat processing treatment

4.1. Irradiation
Electron spin resonance (ESR) spectroscopy has been found to be
useful in detection of irradiated poultry meat (Marchioni et al., 2005a;
Marchioni, Horvatovich, Charon, & Kuntz, 2005b) while gas chromatography can be used to measure volatile hydrocarbons and 2alkylcyclobutanones present in irradiated poultry meat. (Horvatovich
et al., 2000). Another possibility is the Comet assay (single-cell gel
electrophoresis) (Ostling & Johanson, 1984; Singh, McCoy, Tice, &
Schneider, 1988), which is based on electrophoresis of lysed cells
embedded in agarose on a microscopic slide. The intensity of the
Comet tail relative to the Comet head reects DNA damage (Collins,
2004). The Comet assay nds application in the study of irradiationinduced DNA degradation (Klaude, Eriksson, Nygren, & Ahnstrom,
1996); however, the Comet assay cannot be used as a conrmatory
tool as different treatments such as freeze-thaw cycles also cause DNA
damage (Park et al., 2000). Analytical methods for determination of
irradiation have been reviewed elsewhere (Delincee, 2002).
583
Table 4
Examples of analytical techniques applicable in authentication of irradiation, fresh vs.
thawed meat, and preparation in respect to meat and meat products.
Authenticity
problem
Analytical
technique*
Reference
Meat processing
treatment
Irradiation
Electronic spin
Marchioni, Horvatovich, Charon,
resonance spectroscopy and Kuntz (2005a)
GC
Horvatovich, Miesch, Hasselmann,
and Marchioni (2000)
Comet assay
Park et al. (2000)
Fresh vs.
Enzymatic
Ellerbroek, Lichtenberg, & Weise,
Thawed meat
1995; Gottesmann & Hamm, 1983;
Toldra, Torrero, & Flores, 1991
Comet assay
Park et al. (2000)
Infrared spectroscopy
Al Jowder, Kemsley, and Wilson (1997)
NMR
Evans et al., 1998; Guiheneuf, Parker,
Tessier, & Hall, 1997
Electron microscopy
Sen and Sharma (2004)
Preparation
HPLC
Eerola, Hollebekkers, Hallikainen, &
Peltonen, 2007; Paleologos &
Kontominas, 2007
LC-MS/MS
Granby and Fagt (2004)
*Abbreviations used: GC, gas chromatography; HPLC, high performance liquid
chromatography; LC, liquid chromatography; MS, mass spectrometry; NMR, nuclear
magnetic resonance.
4.2. Fresh versus thawed meat

One method capable of distinguishing between fresh and thawed
meat is the enzymatic -hydroxyacyl-CoA-dehydrogenase (HADH)
method (Gottesmann & Hamm, 1983). The HADH method has been
widely used and discussed (Anon, 1996; Billington, Bowie, Scotter,
Walker, & Wood, 1992; Chen, Yang, & Guo, 1988; Toldra et al., 1991).
Originally, one spectrophotometric and one colour test method based
on the reaction (1) were published (Gottesmann & Hamm, 1983).
HADH
1 AcetoacetylCoA + NADH + H hydroxybutyrylCoA + NAD

The spectrophotometric method measures the conversion rate of
NADH to NAD+ by monitoring the decrease in absorption at 340 nm.
Gottesmann and Hamm (1983) reported threshold HADH enzyme
activity values for beef, veal, pork, mutton/lamb, game, and poultry;
surpassing of threshold values indicates a freeze-thaw treatment. The
HADH method is applicable to press juice from whole meat, and
measures -hydroxyacyl-CoA-dehydrogenase released from the freezing induced disruption of mitochondria. A grinding procedure of whole
meat also induces disruption of mitochondria, whereby the HADH
content is released to the press juice. In this case, a signicant HADH
difference between fresh ground meat and thawed meat is absent and
discrimination therefore impossible. It is also important to note that
differentiation between fresh and thawed meat is only possible with the
HADH method if the freezing temperature has been 12 C or below
(Gottesmann & Hamm, 1983). Enzymatic methods have not been
restricted to the HADH method. Other research groups have used the
API-ZYM system, which is based on a semi-quantitative method for
simultaneous measurement of 19 different enzymes (Ellerbroek et al.,
1995; Toldra et al., 1991). Toldra et al. (1991) investigated press juice
from pork and showed that enzymatic activity of esterase-lipase, glucuronidase, and -glucosidase differed signicantly between fresh
and thawed pork. A freeze-thaw regimen also induces structural
alteration in DNA and protein. Altered DNA was correlated with freezing
conditions of vacuum packed whole beef with use of the Comet assay
(Park et al., 2000). Alterations in proteins (haemoglobin) alter the colour
(Deman, 1999), as well the capacity to bind water. Furthermore, water
distribution is changed during a freeze-thaw regimen. Different
spectroscopic techniques detect these changes. Infrared spectroscopy
584
(Al Jowder et al., 1997) was used to study ground meat from pork,
chicken, and turkey after storage at 30 C. Successfully, a principal
component analysis was performed subsequently on spectroscopic data
in the region 10001800 cm 1. Nuclear magnetic resonance (NMR)
(Evans, Nott, Kshirsagar, & Hall, 1998; Guiheneuf et al., 1997) and
electron microscopy (Sen & Sharma, 2004) have also shown potential in
fresh vs. thawed discrimination. An overview of the analytical methods
is provided in Table 4. A review on analytical methods to determine
fresh vs. thawed meat is available (Ballin & Lametsch, 2008).
The use of sensory methods for detection should not be forgotten.
Storage of meat either unwrapped or wrapped in vapour permeable
material at subzero temperatures might result in freezer burn. Freezer
burn is caused by sublimation of ice from the surface region. An
opaque dehydrated surface is easily noticed in freezer burns (Kaess &
Weideman, 1967) and clearly indicates meat stored poorly at subzero
temperatures. Another obvious visual feature is decolouration
induced by freezing (Deman, 1999). Studies on bone marrow and
meat showed a red colour decrease and a brown colour increase after
freezing, while little or no changes occurred for refrigerated samples
(MacDougall, 1982; Nicolalde, Stetzer, Tucker, Mc Keith, & Brewer,
2006). A colour change might therefore indicate whether meat is fresh
or thawed but also odour and tenderness of meat are changed
(Jakobsson & Tsson, 1973; Khan & Berg, 1967; MacDougall, 1982).
5.2. Water
If water is added fraudulently to meat it effectively amounts to
selling water for the price of meat. Addition of water is therefore a real
problem (Elliot, 2007) and regulations dictate the permitted amount
of extraneous water in meat. A standard method to determine
extraneous water in meat is to study the water/protein ratio. For
instance, added water can be determined from a plot of water/protein
ratio against extraneous water in boneless, skinless chicken breast
(Anon, 2005). Water content can be determined by ISO 1442 (Anon,
1997), which describes the measure of mass before and after drying of
meat, and protein content can be determined by ISO 937 (Anon,
1978b), which describes an indirect protein determination method
based on Kjeldahl analysis. If water is added, the water/protein ratio
may to be too high and serve as a clear indicator of addition of water.
However, protein and phosphate can be added to meat products to
increase water binding, leaving the water/protein ratio close to the
natural ratio. In this case, detection of foreign protein or phosphate
might provide proof of fraudulent practice.
Acknowledgements
I would like to thank Kirsten Halkjr Lund for critical comments
on the manuscript.
4.3. Meat preparation (baking, cooking etc.)

With regard to preparation of meat products, a low economic
benet from fraudulent behaviour could perhaps be the reason why
scientic discussions are limited. No analytical methods have to my
knowledge been published in this area. However, production of
Maillard compounds (Skog, Johansson, & Jagerstad, 1998) during
heating could perhaps distinguish among different procedures used in
boiled, fried, and grilled meat. Maillard products form above certain
temperatures, which, for example, make it possible to distinguish
between long-term cooking at a low temperature from short term
cooking at a high temperature. For instance, acrylamide is formed
above 120 C (Mottram, Wedzicha, & Dodson, 2002) and could be
used as an indicator of products exposed to temperature above 120 C.
HPLC (Eerola et al., 2007; Paleologos & Kontominas, 2007) and LC-MS/
MS (Granby & Fagt, 2004) can be used to quantify acrylamide.
5. Identication of non-meat ingredient additions
5.1. Additives
The vast number of organic compounds used as additives makes it
difcult to present a detailed description of each. These organic
compounds might be colorants, aromas, or preservatives. Colorants
and some reducing chemicals can be used to increase fresh meat
appearance. Aromas, such as smoke aroma can fraudulently be used
instead of natural smoking of meat. Preservatives can be used to
preserve meat and make it appear fresh much longer than normal. The
nature of the chemical compound determines the appropriate
analytical technique. Since colour, aroma, and preservative relate to
organic compounds added to the meat products both HPLC and GC
methods may be appropriate.
Another group of additives are enzymes that take part in blood
clotting. These can be used as blood-based binding agents and added
to meat cuts or minced meat to form portions of desired mass and
shape. In the binding process, thrombin cleaves brinogen to
brinopeptides A and B. LC-MS/MS can be used to detect bovine
(Grundy et al., 2007) and porcine (Grundy et al., 2008) brinopeptides A and B in concentrations down to 5%. As the brinopeptides A
and B are species-specic the methods are capable of discriminating
between blood-based binding agents of bovine and porcine origin.
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Meat Science 86 (2010) 577587

Contents lists available at ScienceDirect

Authentication of meat and meat products

N.Z. Ballin / Meat Science 86 (2010) 577587

gas chromatography-mass spectrometry (GC-MS) (Hartwig et al.,

Percentage of mislabelling cases

Undeclared soy protein

Undeclared animal species

Undeclared animal species

Undeclared animal species

Macedo-Silva, Shimokomaki, Vaz, Yamamoto,

Hsieh, Woodward, and Ho (1995)

N.Z. Ballin / Meat Science 86 (2010) 577587

Hartwig, Hartmann, and Steinhart (1997)

Dalvit et al., 2008; Sasazaki et al., 2004

difference in the amounts of collagen present in different meat cuts is

Bayesian statistical model identied the Italian cattle breeds Chianina,

N.Z. Ballin / Meat Science 86 (2010) 577587

Furthermore, head space GC-MS was used to study volatile

higher content of 15N in conventional beef compared with organic

N.Z. Ballin / Meat Science 86 (2010) 577587

Gonzalez-Cordova, Calderon de la Barca, Cota, & Vallejo-Cordoba, 1998; Koppelman,

Real time PCR

Protein (melamine and urea)

Rapidly evolving DNA based techniques have resulted in a change

compared to proteins, DNA has a higher thermal stability, is present in

Fig. 1. Potential meat authenticity problems.

N.Z. Ballin / Meat Science 86 (2010) 577587

3.2. Fat (vegetable and animal fat)

N.Z. Ballin / Meat Science 86 (2010) 577587

chromatographic (LC) methods that focus on protein proles for

4. Identication of meat processing treatment

4.2. Fresh versus thawed meat

1 AcetoacetylCoA + NADH + H hydroxybutyrylCoA + NAD

N.Z. Ballin / Meat Science 86 (2010) 577587

4.3. Meat preparation (baking, cooking etc.)

N.Z. Ballin / Meat Science 86 (2010) 577587

N.Z. Ballin / Meat Science 86 (2010) 577587

N.Z. Ballin / Meat Science 86 (2010) 577587

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