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Francesca Cecchi, Claudia Russo, Giovanna Preziuso, Dario Cianci
Dipartimento di Produzioni Animali. Università di Pisa, Italy.
Corresponding author: Dr. Francesca Cecchi. Dipartimento di Produzioni Animali. Facoltà di Medicina
Veterinaria, Università di Pisa. Viale delle Piagge 2, 56124 Pisa, Italy – Tel. +39 050 3139401 – Fax: +39 050
3139433 – Email: fcecchi@vet.unipi.it
Paper received October 17, 2003; accepted February 2, 2004
Genetic variability
of meat quality traits
in Chianina beef cattle
ABSTRACT
The heritability of quality traits (chemical composition, colour, tenderness and water-holding capacity) of the meat deriv-
ing from three muscles (Triceps brachii, Longissimus dorsi and Semitendinosus) was evaluated in 92 young Chianina
bulls. The animals were raised on two farms with different feeding programs, and were slaughtered at approximately 19
months of age. Single pair correlations were considered in order to evidence the relationships between the same traits
detected in the three muscles and between all the traits of the same muscle.
The h2values range from 0.00 to 0.24 for the chemical composition, from 0.00 to 0.19 for colour parameters, from 0.03
to 0.31 for those observed after 48 h, and from 0.00 to 0.08 for tenderness and water-holding capacity. The values were
not homogeneous for the three muscles. The most heritable trait was yellowness (b*48) in all three muscles, with val-
ues of 0.11 in the Longissimus dorsi, 0.23 in the Semitendinosus, and 0.31 in the Triceps brachii. Conversely, tender-
ness appeared to have a low heritability, as did the parameters relating to the water-holding capacity. Colour traits
observed before and after 48 h of storage generally were significantly and positively correlated, indicating that certain
colour characteristics are maintained even after 48 h of storage. There was not always a close relationship between the
traits of each muscles: significant correlations existed between the Triceps brachii and the Longissimus dorsi muscles,
confirming the similar quality traits of their meat, while there were considerable differences between these muscles and
the Semitendinosus.
Key words: Heritability, Correlations, Meat quality, Chianina breed.
RIASSUNTO
IL MIGLIORAMENTO GENETICO DELLA QUALITÀ
DELLA CARNE DELLA RAZZA CHIANINA: L’EREDITABILITÀ.
Numerosi studi hanno riportato stime di parametri genetici e fenotipici per l’efficienza riproduttiva, l’accrescimento, la
composizione e qualità della carcassa, mentre sono molto scarse le ricerche sulla variabilità genetica della qualità della
carne, per la quale si considerano preponderanti le influenze di origine ambientale. Scopo della presente ricerca è stato
quindi quello di stimare l’ereditabilità dei parametri di qualità della carne della razza Chianina.
Su tre muscoli (Triceps brachii, Longissimus dorsi e Semitendinosus) di 92 vitelloni Chianini, macellati a circa 19 mesi di
età e provenienti da due allevamenti che adottavano diete diverse, sono stati quindi analizzati i parametri di composi-
zione chimica, di colore, di tenerezza e di capacità di ritenzione idrica. Per evidenziare le relazioni esistenti tra tali para-
metri sono state impiegate le correlazioni semplici. I valori di h2ottenuti variano da 0,00 a 0,24 per la composizione chi-
mica, da 0,00 a 0,19 per i parametri di colore, da 0,03 a 0,31 per quelli rilevati dopo la conservazione e da 0,00 a 0,08
per i parametri di tenerezza e di potere di ritenzione idrica. Tali valori non sono risultati omogenei fra i tre muscoli. Il
parametro più ereditabile è risultato l’indice del giallo rilevato dopo 48 ore (b*48) in tutti e tre i muscoli, con valori di
0,11 nel Longissimus dorsi, 0,23 nel Semitendinosus e 0,31 nel Triceps brachii. La tenerezza sembra invece poco eredi-
tabile, così come i parametri relativi alla capacità di ritenzione idrica. Sono da evidenziare correlazioni significative e posi-
ITAL.J.ANIM.SCI. VOL. 3, 191-198, 2004 191
* I.J.A.S. Imp. 02/04 21-12-2005 12:54 Pagina 191
CECCHI et al.
192 ITAL.J.ANIM.SCI. VOL. 3, 191-198, 2004
tive tra i parametri relativi al colore, prima e dopo conservazione, indicando che alcune caratteristiche colorimetriche si
mantengono inalterate anche dopo 48 ore di conservazione in frigorifero. In questa ricerca viene inoltre confermata la
differenza qualitativa tra il muscolo Semitendinosus e gli altri muscoli analizzati. I risultati ottenuti evidenziano che non
è possibile disporre, attualmente, di uno o pochi parametri di semplice determinazione, che permettano di esprimere un
giudizio complessivo sulla qualità della carne, ma che sarebbe necessario identificare un indice globale che prenda in con-
siderazione più parametri qualitativi.
Parole chiave: Ereditabilità, Correlazioni, Qualità carne, Razza Chianina.
Introduction
Meat from Chianina cattle is of excellent quali-
ty (Lucifero et al., 1991; Poli, 1997; Russo and
Preziuso, 2000) and is highly appreciated on the
domestic market (Lucifero and Giorgetti, 1980;
Tartari and Benatti, 1990). However, quality traits
are extremely variable and operations regarding
genetic improvement as well as rearing technique
should be considered. The difficulties of any steps
taken in this direction are compounded by the high
number of traits that determine the quality, and the
need to choose the ones that are most pertinent to
the evaluations. An initial approach to the problem
concerns the establishing of a successful program of
genetic improvement with an acceptable hereditary
base for the chosen parameters. Numerous studies
have reported values of genetic or phenotypic traits
for reproductive efficiency, growth, composition and
carcass traits (Sartore and Di Stasio, 1990;
Mohiuddin, 1993; Koots et al., 1994), but very little
research has been carried out on the genetic vari-
ability of quality traits, which are considered to be
influenced mainly by environmental factors.
Up until now therefore, the estimation of the
genetic aspect of variability has not been
addressed in a decisive manner, and little infor-
mation is available in literature. Heritability stud-
ies have been performed almost exclusively for
tenderness because it is a critical aspect of beef
palatability (Morgan et al., 1991; Brooks et al.,
2000), however they offer varying results, ranging
from low to moderately high (Shackelford et al.,
1994; Gregory et al., 1995; Barkhouse et al., 1996;
Wheeler et al., 1996; O’Connor et al., 1997; Elzo et
al., 1998). Johnston et al. (2001) reports values for
heritability coefficients ranging from 0.19 to 0.39
for shear force, observed on the Longissimus dorsi
muscle in three cattle breeds (Belmont Red,
Brahman, and Santa Gertrudis). Marshall (1994)
reports heritability values for Warner-Blatzler
shear force ranging from 0.09 to 0.71, with an
average of 0.37; similar values (h2=0.26-0.31) were
summarized by Koch et al. (1982), Koots et al.
(1994) and Splan et al. (1998).
Recent estimates carried out on a large number
of carcasses (Burrow et al., 2001) showed that h2
values for meat tenderness were 0.19 in Bos taurus
cattle and 0.23 in both Bos taurus and Bos indicus
straightbred and their crossbreds. Wheeler et al.
(2001) reports 0.22 for tenderness in crossbred cat-
tle, while Riley et al (2003) reported h2values of
0.14, 0.14 and 0.03 for shear force measured after
7, 14 and 21 days respectively. Instead, for param-
eters relative to colour (L*, a* and b*), Aass (1996)
reports values ranging from 0.08 to 0.27.
As far as Chianina cattle are concerned, Franci
et al. (1996) reports non-homogeneous values in
the ten muscles analysed, with low heritability
coefficients for the meat colour traits ranging from
0.00 for L* and b* to 0.07 for a* and medium-low
coefficients for shear force on cooked meat (0.12)
and free water (0.26).
Considering these uncertainties, together with
the inherent risk in transferring results obtained
from different breeds and breeding conditions, the
aim of this work is to contribute to the knowledge
of the hereditary transmission of the Chianina
breed meat quality traits, by estimating heritabil-
ity and correlations.
Material and methods
Animals and meat instrumental measurements
and chemical analysis
Meat from three muscles (Triceps brachii,
* I.J.A.S. Imp. 02/04 21-12-2005 12:54 Pagina 192
HERITABILITY OF MEAT QUALITY TRAITS
ITAL.J.ANIM.SCI. VOL. 3, 191-198, 2004 193
Longissimus dorsi and Semitendinosus) of 92
young Chianina non-consanguineous bulls
raised in Tuscany on two breeding-farms was
analysed. Neither the sires nor the dams of each
family were related. Animals had been slaugh-
tered at approximately 19 months of age, after
attaining a live weight of about 770 kg. The
farms which supplied the bulls used in the test
followed different feeding systems; the first farm
used corn-silage as the basic fodder, while the
second provided hay.
Animals were chosen according to their genealog-
ical data, in order to form paternal half-sib groups.
In accordance with the commercial ageing peri-
ods for Chianina carcasses, the Triceps brachii (TB)
muscle was excised from the forequarters after
about 10 days of ageing, while the Longissimus dorsi
(Ld) and Semitendinosus (St) muscles were taken
from the hindquarters after 19 days of ageing.
In order to assess meat quality each muscle was
instrumentally analyzed (ASPA, 1991) as follows:
Meat colour, using a Minolta CR300 colourme-
ter (illuminant C), calibrated against a standard
white tile in the CIE L*a*b* system, which mea-
sures the value of lightness (L*), redness (a*), yel-
lowness (b*), Croma (C*) and Hue (H*) coordi-
nates (Renerre, 1982), by making three readings
for each sample consisting of a 2.5-cm thick slice of
meat covered with a polyethylene film and refrig-
erated for 45 min. at 4°C.
Meat colour after 48 h, on the same sample
kept at 4°C for 48 h to show any possible colour
changes during meat storage (L*48, a*48, b*48,
C*48 and H*48).
Water-holding capacity (WHC), determined
using two different methods:
- Drip loss, that is the percentage of water lost
from meat during storage at 4°C for 48 h in a
double-bottomed plastic container;
- Cooking loss, on the meat sample used for drip
loss and then cooked on a metal tray in an oven
at 180°C to an internal temperature of 75°C.
Tenderness measured as the shear force (kg) using
Warner Bratzler Shear applied to an Instron 1011, on
1-inch-diameter cylinders of raw and cooked meat.
Chemical analysis: dry matter (%), ether
extract (%), crude protein (%) and ash (%)
(A.O.A.C., 1990).
Statistical analysis
For the evaluation of the heritability, the
following mixed linear sire model was per-
formed using JMP, ver. 5.0 for PC, of the SAS
Institute (2002):
Yijk= µ+ si+ Hj+ bXijk + εijk where Yijk= consid-
ered parameters; µ= overall mean; si= random
effect of the ith sire (i=1, ..., 6); Hj= fixed effect of
the jth herd (j=1, 2); b= regression coefficient on
slaughter age in days; Xijk= slaughter age in days;
εijk = residual error.
All estimates of variance components for esti-
mates of heritability were obtained with a
derivate-free REML algorithm (JMP of the SAS
Institute, 2002). Because a sire model was used,
estimates of ratios of sire to total variance were
multiplied by four to yield heritability estimates.
Standard errors of heritabilities were approxi-
mated according to Becker (1984).
Single pair correlations were considered in
order to evidence the relationships between the
same trait detected in the three muscles, and
between all the traits of the same muscle, as well
as to estimate the correlations of colour parame-
ters measured before and after 48h storage.
Results and discussion
Heritability
Heritability coefficients varied from 0.00 to
0.24 for chemical composition, from 0.00 to 0.19 for
colour parameters before storage, from 0.03 to 0.31
for colour parameters observed after 48h of stor-
age, and from 0.00 to 0.08 for traits of tenderness
and water-holding capacity (Table 1).
For the three muscles, the heritability coeffi-
cients of the colour observed before storage were in
agreement with those determined by Franci et al.
(1996) for Chianina cattle, but lower than those
reported by Aass (1996). Nonetheless, heritability
coefficients improved after meat storage for 48 h
at +4°C; this was especially evident in the Triceps
brachii, for which there was an increase in values
that ranged from 0.02 to 0.05 for colour parame-
ters measured before storage and from 0.08 to 0.31
for those observed after 48h.
* I.J.A.S. Imp. 02/04 21-12-2005 12:54 Pagina 193
CECCHI et al.
194 ITAL.J.ANIM.SCI. VOL. 3, 191-198, 2004
The most heritable trait was the yellowness
measured after 48 h (b*48) in all three muscles,
with values of 0.11 in the Longissimus dorsi,
0.23 in the Semitendinosus, and 0.31 in the
Triceps brachii.
The “L*” and “L*48” parameters also gave
moderately elevated values, especially in the
Semitendinosus, with coefficients equivalent to
0.19 and 0.30 respectively.
On the other hand, tenderness had low heri-
tability, with coefficients falling within the range
reported by Barkhouse et al. (1996) and Riley et al.
(2003), though not compliant with those indicated
by Burrow et al. (2001), and other Authors (Koch
et al., 1982; Koots et al., 1994; Splan et al., 1998;
Johnston et al., 2001 and 2003).
Parameters relating to the water-holding
capacity (drip loss and cooking loss) also appeared
to have low heritability.
Correlations
Correlations among traits (Tables 2, 3, 4) con-
firm that the dry matter of meat is always posi-
tively linked to the ether extract (P≤0.01) and to
the crude proteins (P≤0.01), while the latter two
parameters are negatively correlated to each other
(P≤0.01), as reported in the literature for other
breeds (Russo and Preziuso, 1999).
As is to be expected, in the Triceps brachii and
Longissimus dorsi muscles the parameters relating
to meat colour measured before and after storage
for 48 h are correlated positively with each other in
a highly significant manner. This is partly
explained by the fact that some of them, such as
Chroma (C*) and Hue (H*), derive from the combi-
nation between the redness (a*) and the yellowness
(b*); the positive correlations between lightness
(L*) and the other traits were also interesting, indi-
cating that paler meat colour is usually accompa-
Table 1. Heritability coefficients and standard error (SE) estimates for meat quality
traits in the three muscles.
Muscle
Triceps brachii Longissimus dorsi Semitendinosus
h2SE h2SE h2SE
Chemical composition:
Dry matter 0.02 0.267 0.01 0.238 0.00 0.233
Ether extract 0.02 0.269 0.00 0.237 0.24 0.289
Ash 0.01 0.282 0.03 0.257 0.00 0.234
Crude protein 0.01 0.267 0.00 0.234 0.12 0.263
Meat colour:
L* 0.02 0.269 0.02 0.260 0.19 0.277
a* 0.02 0.268 0.00 0.234 0.05 0.244
b* 0.04 0.272 0.00 0.233 0.12 0.262
C* 0.02 0.269 0.00 0.233 0.05 0.246
H* 0.05 0.273 0.05 0.246 0.05 0.245
After 48 h of storage:
L* 0.08 0.279 0.03 0.240 0.03 0.301
a* 0.19 0.297 0.04 0.244 0.09 0.255
b* 0.31 0.313 0.11 0.259 0.23 0.285
C* 0.19 0.297 0.03 0.239 0.08 0.252
H* 0.13 0.288 0.12 0.261 0.06 0.247
Shear force:
on raw meat 0.08 0.278 0.00 0.233 0.00 0.232
on cooked meat 0.00 0.265 0.01 0.235 0.03 0.241
Water-holding capacity:
Drip loss 0.05 0.273 0.01 0.246 0.00 0.233
Cooking loss 0.01 0.267 0.01 0.239 0.05 0.245
* I.J.A.S. Imp. 02/04 21-12-2005 12:54 Pagina 194
HERITABILITY OF MEAT QUALITY TRAITS
ITAL.J.ANIM.SCI. VOL. 3, 191-198, 2004 195
Table 2. Significant correlations between parameters revealed on Triceps brachii muscle.
Chemical composition Meat colour After 48 h of storage Shear force Water-holding capacity
Dry Ether Ash Crude L* a* b* C* H* L*48 a*48 b*48 C*48 H*48 raw cooked Drip Cooking
matter extract protein meat meat loss loss
Chemical composition:
Dry matter 1.00
Ether extract 0.56** 1.00
Ash - 0.28** 1.00
Crude protein 0.56** - 0.37** 1.00
Meat colour:
L* 1.00
a* 0.31** 1.00
b* 0.52** 0.92** 1.00
C* 0.37** 0.99** 0.95** 1.00
H* 0.66** 0.55** 0.83** 0.63** 1.00
After 48 h of storage:
L* - 0.22* 0.63** 0.23** 0.37** 0.27** 0.45** 1.00
a* 0.47** 0.35** 0.45** 0.34** 1.00
b* 0.46** 0.38** 0.45** 0.20* 0.52** 0.92** 1.00
C* 0.46** 0.35** 0.44** 0.39** 0.99** 0.95** 1.00
H* 0.20* 0.33** 0.22* 0.25** 0.55** 0.48** 0.21* 1.00
Shear force:
on raw meat - 0.29** 0.22* 0.22* 0.20* 1.00
on cooked meat 0.22* 0.21* 0.52** 1.00
Water-holding capacity:
Drip loss 0.24* - 0.31** - 0.26** - 0.31** 1.00
Cooking loss - 0.21* 0.20* 0.19* 0.20* 0.21* 0.40** 0.41** 0.39** 1.00
*: P<0.05; **: P<0.01
Table 3. Significant correlations between parameters revealed on Longissimus dorsi muscle.
Chemical composition Meat colour After 48 h of storage Shear force Water-holding capacity
Dry Ether Ash Crude L* a* b* C* H* L*48 a*48 b*48 C*48 H*48 raw cooked drip cooking
matter extract protein meat meat loss loss
Chemical composition:
Dry matter 1.00
Ether extract 0.72** 1.00
Ash 1.00
Crude protein 0.41** - 0.33** 1.00
Meat colour:
L* 1.00
a* 0.35** 1.00
b* 0.21* 0.20* 0.61** 0.90** 1.00
C* 0.21* 0.42** 0.98** 0.94** 1.00
H* 0.78** 0.41** 0.75** 0.51** 1.00
After 48 h of storage:
L* 0.83** 0.27** 0.56** 0.36** 0.76** 1.00
a* 0.55** 0.46** 0.52** 0.23* 1.00
b* 0.40** 0.50** 0.57** 0.54** 0.44** 0.54** 0.91** 1.00
C* 0.20* 0.54** 0.51** 0.59** 0.24** 0.32** 0.97** 0.93** 1.00
H* 0.73** 0.42** 0.69** 0.82** 0.49** 0.21* 1.00
Shear force:
on raw meat 0.44** 1.00
on cooked meat - 0.18* 1.00
Water-holding capacity:
Drip loss - 0.24** - 0.20* 1.00
Cooking loss - 0.22* 0.20* 0.21* 0.23* 0.23* 0.27** 1.00
*: P<0.05; **: P<0.01
* I.J.A.S. Imp. 02/04 21-12-2005 12:54 Pagina 195
Table 4. Significant correlations between parameters revealed on Semitendinosus muscle.
Chemical composition Meat colour After 48 h of storage Shear force Water-holding capacity
Dry Ether Ash Crude L* a* b* C* H* L*48 a*48 b*48 C*48 H*48 raw cooked drip cooking
matter extract protein meat meat loss loss
Chemical composition:
Dry matter 1.00
Ether extract 0.34** 1.00
Ash 1.00
Crude protein 0.79** - 0.32** 1.00
Meat colour:
L* 0.29** 0.33** 1.00
a* 0.25** - 0.22* 1.00
b* 0.38** 0.70** 1.00
C* 0.22* 0.98** 0.82** 1.00
H* 0.25** 0.31** 0.78** 0.59** 1.00
After 48 h of storage:
L* 0.26** 0.88** 0.38** 0.73** 1.00
a* - 0.24** 0.42** 0.23* 0.44** -0.23* 1.00
b* 0.44** 0.49** 0.48** 0.20* 0.31** 0.81** 1.00
C* - 0.22** - 0.20* 0.49** 0.29** 0.44** 0.95** 0.89** 1.00
H* 0.56** 0.24* 0.40** 0.54** - 0.50** - 0.47** 1.00
Shear force:
on raw meat - 0.25** 1.00
on cooked meat 1.00
Water-holding capacity:
Drip loss 0.26** 0.26* - 0.25** - 0.27** 0.35** 1.00
Cooking loss 1.00
*: P<0.05; **: P<0.01
CECCHI et al.
196 ITAL.J.ANIM.SCI. VOL. 3, 191-198, 2004
nied by greater luminosity, which contributes to
conferring a “brilliant” aspect to the product.
In all muscles (Tables 2, 3, 4) the colour parame-
ters measured before storage were correlated in a
positive and significant manner with those evaluat-
ed after 48 h of conservation, in particular lightness
(r= 0.63, 0.83 and 0.88 respectively in Triceps brachii,
in Longissimus dorsi and in Semitendinosus).
Moreover, in the Triceps brachii and
Longissimus dorsi muscles, a*, b*, and C* mea-
sured after 48 h are significantly and positively cor-
related with water loss after cooking.
Significant correlations between tenderness
and redness (a* and a*48) and between tenderness
and Croma (C*48) are only observed in the Triceps
brachii muscle; furthermore, there is a significant
and positive correlation between shear force on
raw and cooked meat (r=0.52; P≤0.01).
Table 5 illustrates the correlations for each
quality trait among the three muscles: as can be
observed, the only parameters always correlated
in a significant manner were a*48, C*48 and drip
loss, while no significant correlations are recorded
for b*48, H* 48, and cooking loss.
The majority of significant correlations
between the Triceps brachii and Longissimus
dorsi muscles confirms the similar quality traits in
meat deriving from these two muscles, also in
accordance with the findings of another study
(Russo and Preziuso, 2000).
Conclusions
An examination of the results of heritability
coefficients and correlations between quality
traits led us to draw the following conclusions:
- many of the quality traits of Chianina meat are
affected by somatic variability factors: only the
heritability coefficients of the colour parameters
observed after storage are moderately-high, the
most heritable trait being the yellowness mea-
sured after 48 h (b*48) in all three muscles. The
heritability values for tenderness parameters
are low, indicating that their selective improve-
ment would be slow;
- in order to improve meat quality there are diffi-
culties involved in reducing the number of traits
to be analysed: correlations are normally
* I.J.A.S. Imp. 02/04 21-12-2005 12:54 Pagina 196
HERITABILITY OF MEAT QUALITY TRAITS
ITAL.J.ANIM.SCI. VOL. 3, 191-198, 2004 197
observed in each of the three muscles within the
same group of parameters (i.e. composition
parameters, colour parameters);
- the improvement of tenderness for the consumer
is not easy either since the shear force measured
on raw meat is not related to the shear force
measured on cooked meat; only the Triceps
brachii presents a positive relationship between
tenderness both before and after cooking.
These results underline the fact that, at the
moment, for improving quality traits of Chianina
meat it is important to pay particular attention to
the management problems in order to optimise the
heritable fraction of the variability. Moreover, it is
not possible to utilise one or two easily identified
traits alone for obtaining a complex evaluation of
meat quality, it is instead necessary to consider a
variety of different traits.
It would be interesting to identify a comprehen-
sive index capable of outlining the qualitative
aspects, but which also shows an acceptable heri-
tability, thus furthering prospects of genetic progress.
Research supported by “ARSIA”: “Plan for the
increase in value of the Tuscan bovine genetic
material and meat production”.
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Table 5. Correlation for each parameter between muscles.
Tb 1vs Ld 2Tb 1 vs St 3Ld 2vs St 3
Meat colour:
L* 0.330** 0.332** 0.199
a* 0.291** 0.115 0.321**
b* 0.258* 0.009 0.181
C* 0.272** 0.083 0.259*
H* 0.221* 0.154 0.170
After 48 h of storage:
L* 0.330** 0.196 0.121
a* 0.283** 0.315** 0.268**
b* 0.096 0.145 0.108
C* 0.246* 0.268** 0.252*
H* 0.143 0.165 0.096
Shear force:
on raw meat 0.137 0.162 0.235*
on cooked meat 0.153 0.215* 0.198
Water-holding capacity:
Drip loss 0.246* 0.302** 0.368**
Cooking loss 0.150 0.055 0.021
*.P<0.05; **.P<0.01.
1 Tb:Triceps brachii;
2 Ld:Longissimus dorsi;
3 St:Semitendinosus.
* I.J.A.S. Imp. 02/04 21-12-2005 12:54 Pagina 197
CECCHI et al.
198 ITAL.J.ANIM.SCI. VOL. 3, 191-198, 2004
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