MEAT

SCIENCE
Meat Science 74 (2006) 231–241
www.elsevier.com/locate/meatsci

Changes in colour characteristics and pigmentation

of subcutaneous adipose tissue and M. longissimus dorsi of heifers
fed grass, grass silage or concentrate-based diets
a,b
P.G. Dunne , F.P. O’Mara b, F.J. Monahan b, A.P. Moloney a,*

a
Teagasc, Grange Research Centre, Dunsany, Co. Meath, Ireland
b
School of Agriculture, Food Science and Veterinary Medicine, University College Dublin, Dublin 4, Ireland

Received 30 September 2005; received in revised form 8 February 2006; accepted 8 February 2006

Abstract

The objective of this experiment was to determine the effect of dietary composition and duration of feeding on subcutaneous (s.c.)
adipose tissue and M. longissimus dorsi (LD) colour and pigment concentrations of heifers. Fifteen heifers were permanently housed
and fed a concentrate diet (PH-CON). Fifty-four heifers were grazed on pasture (PAS) for 90d, housed and offered concentrates
(PAS-CON), 200 g grass silage (GS)/kg dry matter (DM) (PAS-GS20), 500 g GS/kgDM (PAS-GS50) or zero-grazed grass (PAS-
GRA). To facilitate assessment of the temporal pattern of tissue colour, 3 heifers/treatment were slaughtered at housing (following
7d adjustment to diets) and 28, 56, 91 and 120 days thereafter. Yellowness (‘b’ value) of s.c. adipose tissue and lightness (‘L’ value)
and redness (‘a’ value) of LD were recorded 48 h post-mortem. b-Carotene and lutein contents of s.c. adipose tissue and total LD haem
pigments were determined. At housing, s.c. adipose tissue ‘b’ values of the PAS group (mean = 13.47) were higher (P < 0.01) than those
of the PH-CON group (mean = 10.35) but there was no difference in b-carotene or lutein concentrations. The change in s.c. adipose
tissue ‘b’ for each diet following housing was best described by the following equations: PH-CON: y = 0.087 (SE 0.0347)
X + 0.0005 (SE 0.00029) X2 + 10.06 (SE 0.600), MSE 1.94, R2 0.57, P < 0.01. PAS-CON: y = 0.102 (SE 0.0286) X + 0.0006 (SE
0.00024) X2 + 13.32 (SE 0.598), MSE 2.30, R2 0.62, P < 0.001. PAS-GS20: y = 0.106 (SE 0.0296) X + 0.0008 (SE 0.00025)
X2 + 13.10 (SE 0.618), MSE 2.46, R2 0.47, P < 0.01. PAS-GS50: y = 0.077 (SE 0.0269) X + 0.0006 (SE 0.00023) X2 + 13.29 (SE
0.552) MSE 1.95, R2 0.38, P < 0.05. PAS-GRA: y = 0.018 (SE 0.0079) X + 13.77 (SE 0.528), MSE 2.28, R2 0.24, P < 0.05, where
y = ‘b’ value, x = days post-housing (d96–d216) and MSE = mean square error. Treatment had a significant effect on s.c. adipose tissue
b-carotene and lutein (both P < 0.001) with PAS-GRA and PH-CON tending to have the highest and lowest concentrations, respectively.
PH-CON heifers tended (P = 0.058) to have lower LD haem pigments and lighter LD than other heifers. It is concluded that, while
concentrate feeding led to the greatest decrease in s.c. adipose tissue yellowness relative to PAS-GRA, choice of dietary ingredients
and duration of feeding will depend on the stringency of the colour criteria in particular markets.
 2006 Elsevier Ltd. All rights reserved.

Keywords: Colour; Adipose; Muscle; Heifers; Diets

1. Introduction and relatively cheap source of cattle feed (O’Riordan &

The temperate, maritime climate of North-Western Eur-
ope is conducive to grass growth (Lee, 1988). Conse-
quently, fresh and conserved grass are widely used in beef
production in this region, as they represent a renewable
*
Corresponding author. Tel.: +353 46 90 61100; fax: +353 46 90 26154.
E-mail address: amoloney@grange.teagasc.ie (A.P. Moloney).
O’Kiely, 1996). Consumption of the carotenoid pigments
in green leafy forage results in yellow subcutaneous (s.c.)
adipose tissue (carcass fat) of cattle (Strachan, Yang, &
Dillon, 1993; Yang, Larsen, & Tume, 1992) and such beef
is unacceptable in markets of Southern Europe, where car-
casses with ‘white’ carcass fat command a premium price
(Anon, 1999; Dunne, O’Mara, Monahan, & Moloney,
2004). Grass silage (GS), supplemented with concentrates,

0309-1740/$ - see front matter  2006 Elsevier Ltd. All rights reserved.
doi:10.1016/j.meatsci.2006.02.003
232 P.G. Dunne et al. / Meat Science 74 (2006) 231–241
is the dominant feedstuff offered to housed cattle during the
winter months in Ireland and much of North-Western Eur-
ope (French et al., 2000; O’Kiely, Power, & O’Donnell,
1993). However, little information is available on the
threshold for inclusion of GS in diets of housed finishing
cattle such that their carcass fat is considered acceptably
pale for Southern European beef markets.
The majority of Irish calves are Spring-born (Drennan,
Keane, & Dunne, 1995) and many beef production systems
involve one or two grazing seasons from March to early
November. Cattle given an opportunity to graze pastures
for extended periods during their production, are reported
to have ‘darker’ muscle than cattle raised in more intensive
systems (Muir, Smith, Wallace, Cruickshank, & Smith,
1998; Priolo, Micol, & Agabriel, 2001). ‘‘Darker’’ muscle
is less desired by consumers and can also result in exclusion
of carcasses from the lucrative markets mentioned above
(Anon, 1999).
The objective of this study was to determine the effect of
dietary composition and duration of feeding on s.c. adipose
tissue and M. longissimus dorsi (LD) colour and pigment
concentrations of heifers and to model changes in the yel-
lowness with a view to predicting the time necessary to
achieve an acceptable s.c. adipose tissue yellowness. We
hypothesised (a) that feeding a non-grass-based diet to
heifers which had been grazing for three months previ-
ously, would reduce yellow colour in s.c. adipose tissue in
a time-dependent manner; (b) that within these diets, a
decrease in the proportion of GS would decrease yellow
colour in s.c. adipose tissue and (c) that removal of the
grazing opportunity would result in paler muscle.
2. Materials and methods
2.1. Experimental design and animal management
Seventy-five continental crossbred (Charolais · Limou-
sin/Friesian) heifers (mean live weight of 387 [SD = 30.6]
kg) were divided into three blocks (n = 25 heifers per
block) on the basis of decreasing age. Within block, heifers
were randomly assigned to 1 of 25 cells. Corresponding
cells from each age block were grouped to give 25 groups
of three heifers each. Groups were then randomly assigned
to treatments (T) and slaughter dates (D). Prior to the com-
mencement of the experiment, six heifers which had been
assigned to a pre-experimental group were slaughtered.
On April 26, fifty-four heifers were turned out to pasture
where they rotationally grazed six paddocks for 90 days.
One heifer, discovered to be pregnant was removed from
the trial during this stage. Fifteen heifers were permanently
housed (PH-CON) from April 26 and offered 1.5 kg con-
centrate and 2 kg straw per head initially, increasing by
0.5 kg concentrate daily until an allowance of 6 kg per head
was reached. The concentrate consisted of rolled barley
(430 g/kg), molassed sugar beet pulp (430 g/kg), soyabean
meal (80 g/kg), molasses (45 g/kg) and a proprietary min-
eral/vitamin mix (15 g/kg). On d90, heifers previously graz-
ing at pasture (PAS) (n = 53) were weighed and allocated
to their pre-assigned T and D and housed in slatted floor
pens, in groups of six according to treatment. Six of these
53 heifers continued to graze a paddock adjacent to the
house for a further 6 days (while the remaining 47 previ-
ously grazing PAS heifers were adjusted to their diets) until
they, and six of the PH-CON heifers were slaughtered on
what was designated as d0 of the indoor phase, coinciding
with full implementation of treatments (diets). The four
additional treatments were 6 kg of concentrate plus 2 kg
of straw (PAS-CON), 6 kg of concentrate plus 1.47 kg of
GS dry matter (DM) (PAS-GS20), 4.2 kg of concentrate
plus 3.5 kg of GS DM (PAS-GS50) and 1.5 kg concentrate
plus 6.4 kg of grass DM (PAS-GRA). Heifers (n = 3/treat-
ment, on each date) were slaughtered on d28, 56, 91 and
120 days thereafter. Due to a limitation on the numbers
of animals available, no PH-CON heifers were assigned
for slaughter at d56 indoors. All heifers were weighed at
intervals of approximately 3 weeks and when necessary,
adjustments to the dietary allowances were made to main-
tain comparable growth rates and carcass fatness across
treatments while avoiding feed refusals.
Concentrates were offered to all treatments in two equal
feeds, once in the morning and again in the late evening.
Fresh forage was offered once daily. Grass was mechani-
cally recovered from pasture on a daily basis using a disc
mower equipped with a flail conditioner (Pottinger, Grie-
skirchen 4710, Austria). To calculate the fresh weight
allowance, forage DM concentration was determined daily
for mechanically-recovered grass and weekly for GS, by
recovering a representative sample of approximately
100 g and drying in a microwave oven to a constant weight.
2.2. Sampling of feedstuffs
For general chemical analysis, concentrate (1 ± 0.1 kg)
and straw samples (0.2 kg) were collected on a weekly basis
throughout the experiment. During the in situ grazing per-
iod, grass samples (1 ± 0.1 kg) were collected from alter-
nate paddocks on the day immediately prior to grazing
by the heifers. After housing, grass and GS samples
(1 ± 0.1 kg) were collected daily. All samples were stored
at 20 C. At the end of the experiment, samples were
composited to represent a 1–2 week period, partially
thawed and chopped in a bowl chopper (Müller, Saarbrüc-
ken, Germany) and re-frozen at 20 C, in the dark, prior
to chemical analysis. Composite straw and concentrate
samples, each representing 3–5 weeks of the experimental
period, were stored in the same manner.
2.3. Slaughter, tissue sample recovery and post slaughter
measurements
On each slaughter date, animals were weighed, trans-
ported 128 km to a commercial slaughter facility and
slaughtered humanely, following stunning by captive bolt,
within 1.5 h of arrival. After slaughter, the kidney and
 P.G. Dunne et al. / Meat Science 74 (2006) 231–241
channel fat (KCF) was excised from both sides of the car-
cass and weighed. A sample of the external KCF surface
measuring approximately 7.5 · 7.5 cm was placed in a foil
tray, covered and stored at 4 C until 24 h post-mortem
when its colour was measured using a Minolta chromame-
ter (model CR300, Minolta Camera Co. Ltd., Osaka,
Japan). Hot carcass weights were recorded, carcass fat
score and conformation were assessed according to EU
guidelines (Anon, 1981) and carcasses were held at 4 C
until 24 h post-mortem. Carcass fat colour was subjectively
assessed for a Southern European beef market by an expe-
rienced industry grader as ‘acceptable’, ‘borderline’ or
‘unacceptable’ (‘excessively yellow’). Carcass right sides
were then cut at the 5th/6th rib interface (‘pistola’ cut)
and the LD muscle, between the 7th and 13th ribs, was
excised and stored at 4 C until 48 h post-mortem. A sec-
tion of s.c. adipose tissue, measuring approximately
10 · 10 cm was removed from over the 10th rib for colour
measurement and carotenoid analysis. The pH of the LD
muscle (pHu) was measured by making a scalpel incision
at the 10th rib and inserting a glass electrode (Model EC-
2010-06, Amagruss Electrodes Ltd., Westport, Co. Mayo,
Ireland) attached to a portable pH meter (Model no.
250 A, Orion Research Inc., Boston, MA) approximately
2.5 cm into the muscle. A steak, 2.5 cm in thickness, was
removed from the 12th rib, trimmed of adhering s.c. adi-
pose tissue, overwrapped with oxygen-permeable PVC film
and permitted to bloom at 4 C, in darkness for 3 h prior to
colour measurement (Strange, Benedict, Gugger, Metzger,
& Swift, 1974). Another steak of 2.5 cm thickness was
removed adjacent to the colour steak and frozen at
20 C, in darkness, for muscle haem pigment analysis.
2.4. Instrumental measurement of tissue colour coordinates
The Minolta chromameter was calibrated for readings of
‘L’ (lightness), ‘a’ (redness) and ‘b’ (yellowness) values using
its standard white calibration tile, overwrapped prior to
measurement with PVC film, where appropriate. On each
date of use, improvised red and yellow cards were used as
standards to ensure reproducibility of readings between
dates. Triplicate measurements were made on three non-
overlapping zones of each tissue and average values calcu-
lated as final readings. All measurements were made in
the Hunter Lab colour space using the D65 illuminant. Tis-
sue ‘H’ value (hue) and ‘C’ value p (saturation or chroma)
were calculated as tan1(b/a) and (a2 + b2), respectively
(McLaren, 1987). Final conversion of hue from radians to
degrees was achieved by multiplying tan1(b/a) by 180/p.
2.5. Chemical analysis
2.5.1. Extraction and measurement of pigments from
subcutaneous adipose tissue and muscle
b-carotene and lutein concentrations in s.c. adipose tis-
sue were determined using modifications of the saponifica-
tion and extraction procedures used by Strachan et al.
233
(1993). During saponification, the water bath was covered
with aluminium foil to prevent the ingress of light, after
which 3 · 1 mL volumes of distilled water at 65 C were
added and the saponified adipose tissue was homogenised
using an Ultra-Turrax T25 homogeniser (Janke & Kunkel
GmbH, IKA Labortechnik, Staufen, Germany) operated
at 8000 rpm for 60 s. Carotenoids were extracted three
times using 4 · 1 mL volumes of diethyl ether (containing
0.004% BHT; Sigma Chem. Co., St. Louis, MO) incorpo-
rating centrifugation (250g; 3 mins) between extractions
to improve separation between the lower and the upper
phase that contained the extracted carotenoids. Anhydrous
sodium sulphate was added to the final ether extract to
remove moisture, then removed by centrifugation (250g;
3 min.) and the ether extract was evaporated to dryness
under N2 at room temperature. The residue was reconsti-
tuted in 0.5 mL of mobile phase (CH3CN (acetonitrile):
CH3OH (methanol) : CH2Cl2 (dichloromethane) at a ratio
of 60:35:5 (Pupin, Dennis, & Toledo, 1999)).
b-Carotene and lutein in samples and standards were
determined by reverse phase high performance liquid chro-
matography, using a modification of the conditions used by
Strachan et al. (1993) whereby separation of carotenoids
was carried out on a C8 column (250 · 4 mm, packing
material Lichrosphere RP-select B, 5 lm particle size;
Phenomenex, Macclesfield, UK). The carotenoids were
eluted using CH3CN: CH3OH: CH2Cl2 (60:35:5) (Pupin
et al., 1999) at a flow rate of 0.5 mL per minute. Detection
was achieved using a Spectra-Physics UV 2000 spectropho-
tometric detector set at 436 nm (Strachan et al., 1993) and
chromatograms were recorded using a Spectra-Physics
SP4600 Chrom-Jet Integrator (Spectra-Physics, Inc., San
José, CA). Samples (40 lL) were injected through a syringe
loading sample injector equipped with a 20 lL loop (Model
7125NS, Rheodyne, Inc., Cotati, CA).
Haem pigments were extracted from LD samples as out-
lined by Warriss (1979). Total haem pigment content of LD
muscle was determined using the method of Krzywicki (1982).
2.5.2. Compositional analysis of feedstuffs and muscle
Compositional analysis of LD muscle (intramuscular
(i.m.) lipid, moisture, protein) was carried out as previously
described by French et al. (2001). Feedstuff DM concentra-
tion was determined by oven drying at 103 ± 2 C for 4 h,
crude protein (CP) was determined by the AOAC approved
method (AOAC, 1990) using a Leco protein analyser
(model FP428, Leco Corp., St. Joseph, MI), ash content
was determined in a muffle furnace at 550 C for 5 h,
DM digestibility (DMD) was determined by the method
of Tilley and Terry (1963). Silage was washed in water
and the pH measured with a glass electrode. Average chem-
ical composition of feedstuffs is presented in Table 1.
2.6. Statistical analysis
Data (carcass characteristics, tissue pigment concen-
trations, pH and colour coordinates for each tissue) were
234 P.G. Dunne et al. / Meat Science 74 (2006) 231–241

Table 1
Average chemical composition of the feedstuffs
Sample Compositiona
pH DM (g/kg) CP (g/kg DM) Ash (g/kg DM) DMD (g/kgDM)
Barley-based concentrate – 858.9 127.6 73.5 869.3
Grazed grass – 208.8 214.6 91.0 835.2
Mechanically-recovered grass – 141.6 205.4 133.3 705.4
Grass silage 3.94 213.7 170.9 111.9 719.3
Straw – 884.5 53.0 59.3 454.9
a
DM: dry matter; CP: crude protein; DMD: dry matter digestibility.

subjected to analysis of variance using a model that had
block, T, D and T · D as sources of variation. Where the
F-test was significant, means were separated using Fisher’s
least significant difference test. Data from the pre-experi-
mental group and the two groups slaughtered on d0 were
subjected to analysis of variance using a model that had
block and T as sources of variation. Linear modelling
approaches were used to determine relationships between
relevant tissue colour coordinates and pigment contents
and to determine the pattern of change of tissue yellowness
over time.
3. Results
3.1. Changes in carcass traits, tissue colour and pigment
concentrations due to in situ grazing
Unless stated otherwise, differences were significant at
P 6 0.05. Compared to animals slaughtered at the begin-
ning of the study, grazing for 96 days increased carcass
weight, had little effect on carcass characteristics, increased
s.c. yellowness (higher ‘b’ value and lutein concentration)
while increasing muscle redness and haem pigment concen-
tration. (Table 2). Compared to PAS, the PH-CON group
had less (P < 0.1) and more white KCF (Table 2), more
white s.c. adipose tissue that contained less lutein and less
haem in LD muscle (Table 2).
3.2. Changes in carcass traits, tissue colour and pigment
concentrations during housing
3.2.1. Carcass traits
On average, carcasses from GS50 were heaviest (Table
3). Carcass weight increased as the experiment progressed
(245, 251, 256, 279 and 287 kg, at housing and d28, 56,
91 and 120 thereafter, respectively, SED = 6.4) but there
was no interaction between T and D. There was no effect
of T nor any interaction between T and D for indices of
carcass fatness (KCF weight, as kg or g/kg carcass and
i.m. lipid concentrations) although KCF weight increased
with increasing D (Table 3).
3.2.2. Adipose tissue colour coordinates and pigment
concentrations
Changes in s.c. adipose tissue and KCF ‘b’ and ‘C’ val-
ues during the experiment are presented in Table 4. There
was a T · D interaction for s.c. adipose tissue ‘b’ value
whereby yellowness of PAS-GS20 and PAS-GS50 heifers
decreased from housing to day 28, but then increased to
day 120, significantly so for PAS-GS50. The ‘b’ value of
PAS-CON heifers decreased (P < 0.01) from housing to
day 28, and tended to decrease to day 120, similar to
PH-CON heifers. The ‘b’ value of PAS-CON heifers
remained higher (P < 0.05) than the PH-CON heifers at
each D. The PH-CON heifers had the least yellow s.c. adi-
pose tissue (P < 0.05) on all dates. The s.c. adipose tissue of
the PAS-GRA tended to be most yellow but this depended
on D; PAS-GRA was more yellow than PAS-GS50 on d28
but not on d56 or d91. The ‘b’ and ‘C’ values of KCF were
affected by T (both P < 0.001) whereby PH-CON heifers
had the least yellow KCF and PAS-GRA heifers tended
to have the most yellow KCF, but not significantly more
yellow than the PAS-GS50 heifers. Approximately 40%
of the variation in s.c. ‘b’ was explained by KCF ‘b’. Inclu-
sion of the square of KCF ‘b’ in the model did not improve
the fit (s.c. ‘b’ = 0.55 (SE 0.082) KCF ‘b’ + 2.42 (SE 1.24)
MSE 3.13, P < 0.001.
The change in s.c. adipose tissue ‘b’ for each diet follow-
ing housing of all animals (d96) was best described by the
following equations:
PH  CON : y ¼ 0:087ðSE 0:0347ÞX
þ 0:0005ðSE 0:00029Þ X 2 þ 10:06 ðSE 0:600Þ;
MSE 1:94; R2 0:57; P < 0:01;
PAS  CON : y ¼ 0:102 ðSE 0:0286Þ X
þ 0:0006ðSE 0:00024Þ X 2 þ 13:32ðSE 0:598Þ;
MSE 2:30; R2 0:62; P < 0:001;
PAS  GS20 : y ¼ 0:106 ðSE 0:0296Þ X
þ 0:0008ðSE 0:00025Þ X 2 þ 13:10 ðSE 0:618Þ;
MSE 2:46; R2 0:47; P < 0:01;
PAS  GS50 : y ¼ 0:077 ðSE 0:0269Þ X
þ 0:0006ðSE 0:00023Þ X 2 þ 13:29 ðSE 0:552Þ
MSE 1:95; R2 0:38; P < 0:05;
PAS  GRA : y ¼ 0:018 ðSE 0:0079Þ X
þ 13:77ðSE 0:528Þ; MSE 2:28; R2 0:24; P < 0:05;
where y = ‘b’ value, x = days post-housing (d96 – d216)
and MSE = mean square error.
 P.G. Dunne et al. / Meat Science 74 (2006) 231–241 235

Table 2 Table 3
Liveweight, carcass traits and tissue colour of heifers prior to the study Carcass weights and indices of carcass fatness throughout the experiment
and after grazing or fed indoors for 96 days
Day (D)a Treatment (T) Carcass (kg) KCF (kg)b Intramuscular
a
Variable Treatment P-value SED lipid (g/100 g)c
PRE PAS PH-CON 124(28) PH-CON 239.8 4.51 1.51
Liveweight (kg) b
417.2 465.7 451.9 * 13.73 PAS-CON 255.5 4.69 2.00
PAS-GS20 265.9 3.99 1.15
Carcass (kg) 211.7 247.6 238.9 * 10.48
PAS-GS50 261.7 6.28 1.84
Conformationc 2.2 2.7 2.6 NS 0.29
PAS-GRA 234.0 3.69 1.42
Fat scorec 2.9 3.0 2.7 NS 0.36
KCF (kg) 6.0 5.5 3.7 NS 0.99 152(56) PH-CON 247.7 4.76 1.71
KCF (g/kg)d 27.9 22.3 15.4 * 3.73 PAS-CON 262.2 4.84 1.94
KCF coloure PAS-GS20 262.0 5.03 1.23
PAS-GS50 260.1 3.68 2.64
‘L’ 67.4 69.4 69.1 NS 2.29
PAS-GRA 250.8 5.22 1.58
‘a’ 4.6 5.8 7.5 NS 1.90
‘b’ 17.7 15.3 14.0 ** 0.60 187(91) PH-CON 266.3 7.26 2.36
‘H’ 75.7 69.4 62.9 NS 6.27 PAS-CON 274.9 6.84 2.04
‘C’ 18.4 16.6 16.3 * 0.78 PAS-GS20 260.2 7.14 2.16
PAS-GS50 309.0 9.97 1.51
s.c. adipose coloure
‘L’ 65.1 67.8 66.7 NS 1.22 PAS-GRA 285.8 7.19 2.27
‘a’ 4.8 4.6 5.4 NS 0.86 216(120) PH-CON 286.3 8.28 2.71
‘b’ 10.9 13.5 10.3 * 0.85 PAS-CON 286.1 6.05 1.57
‘H’ 66.3 71.1 62.5 NS 3.59 PAS-GS20 271.6 5.57 2.89
‘C’ 12.0 14.4 11.6 * 0.87 PAS-GS50 312.6 8.79 3.12
b-carotene, lg/g 0.24 0.37 0.22 NS 0.091 PAS-GRA 278.6 5.93 1.86
Lutein, lg/g 0.16 0.28 0.18 * 0.048 **
T NS NS
e *** ***
LD colour D NS
‘L’ 36.8 35.1 34.6 * 0.61 SED 6.42 0.615 0.324
‘a’ 9.8 13.3 12.9 * 0.90
‘b’ 6.1 4.8 4.6 * 0.41 T·D NS NS NS
‘H’ 31.8 20.0 19.7 ** 0.92 SED 14.35 1.375 0.725
a
‘C’ 11.6 14.1 13.7 * 0.97 Days in parentheses are corresponding days after housing of all heifers,
following 90 days grazing and a 6d dietary adjustment period.
LD composition, g/100 g b
Kidney and channel adipose tissue.
Protein 23.2 23.6 24.0 NS 0.30 c
Intramuscular lipid of M. longissimus dorsi.
Moisture 74.7 74.5 73.9 NS 0.80 **
P < 0.01.
Lipid 2.2 1.9 2.2 NS 0.64 ***
P < 0.001.
pHu 5.63 5.74 5.77 NS 0.059
Haem pigments, mg/g 5.55 6.61 5.64 * 0.369
a
The PH-CON group (n = 5) were housed from the outset of the b-Carotene and lutein concentrations in s.c. adipose
experiment until slaughter at d96. The PAS group (n = 6) were permitted
tissue were affected by T but not by D (Table 4). Thus
to graze a series of paddocks for 96 days prior to slaughter. The PRE
group were slaughtered at the beginning of the study. PAS-GRA heifers had the highest and PH-CON heifers
b
Liveweight at slaughter. had the lowest concentrations of both pigments. There
c
Assessed according to the EU Beef Carcass Classification Scheme for was a tendency for b-carotene concentration to increase
the carcasses of adult bovine animals. Conformation classes E (best), U, as the proportion of forage in the diet increased but of
R, O, P (worst) are assigned numerical values 5, 4, 3, 2 and 1, respectively.
the two GS treatments, only PAS-GS20 was significantly
Fat score classes 1(least fat), 2, 3, 4H, 4L and 5 (most fat) are assigned
numerical values 1, 2, 3, 3.66, 4.33 and 5, respectively. lower than PAS-GRA. There was no difference in b-caro-
d
g/kg carcass. tene concentration between PAS-CON and PH-CON heif-
e
‘L’: lightness. Scale 0 (black) to 100 (white); ‘a’: redness. Scale + ers. PAS-CON heifers, had a higher (P < 0.05) lutein
a (red) to a (green); ‘b’: yellowness. Scale +b (yellow) to b (blue); concentration than PH-CON heifers but were not differ-
‘H’: hue = tan1(b/a)(180/p);‘C’: chroma/saturation/colour intensity =
p 2 ent from PAS-GS20 or PAS-GS50. PAS-GRA heifers
(a + b2).
tended to have a higher lutein concentration than the
GS treatments.

The change of b value in PAS-GRA animals was best
described by a linear model whereas, a quadratic (polyno-
mial) function best described the change for the other diets.
The pattern of change in ‘b’ value for PH-CON from the
beginning of the experiment (216 days) was best described
by the quadratic equation y = 0.014 (SE 0.0129)
X  0.00003 (SE 0.000061) X2 + 11.02, MSE 1.96, R2
0.61, P < 0.001.
3.2.3. Muscle pH, colour coordinates and pigment
concentrations
Muscle tissue pH, pigment and colour coordinates are
presented in Table 5. There was a T · D interaction for
pH whereby between housing and d28, all T, with the
exception of PAS-GS50, tended to decrease. This trend
continued between d28 and d56 but the decrease was
marginal for the PAS-GRA heifers (5.48 to 5.42) which
236 P.G. Dunne et al. / Meat Science 74 (2006) 231–241

Table 4
Yellowness (‘b’) and saturation (‘C’) of subcutaneous (s.c.) adipose tissue and kidney and channel fat (KCF) and adipose tissue pigment concentrations
throughout the experiment
Day (D)a Treatment (T) s.c. adipose tissue KCF Carotenoid concentrations (lg/g)
b C b C b-carotene Lutein
s s
124(28) PH-CON 7.14 8.50 10.24 11.59 0.19 0.14
PAS-CON 10.33uvw 11.73tu 15.36 16.51 0.25 0.21
PAS-GS20 9.39tu 11.50t 13.30 14.95 0.19 0.18
PAS-GS50 10.87uvw 11.78tu 14.96 16.84 0.34 0.28
PAS-GRA 14.46z 15.88w 19.18 20.72 0.70 0.33
152(56) PH-CON 6.81s 8.04s 11.88 13.17 0.12 0.12
PAS-CON 9.58u 11.03t 14.08 15.18 0.24 0.18
PAS-GS20 9.63u 11.22t 15.63 16.57 0.26 0.19
PAS-GS50 11.27uvwx 12.49tuv 14.85 16.61 0.26 0.21
PAS-GRA 12.07vwxy 12.72tuv 16.94 17.51 0.46 0.31
187(91) PH-CON 7.41st 8.75s 13.41 15.27 0.16 0.15
PAS-CON 9.43tu 11.18t 14.42 16.32 0.22 0.16
PAS-GS20 11.47uvwxy 12.25tu 14.93 12.63 0.27 0.15
PAS-GS50 12.37wxyz 15.52w 16.91 17.89 0.45 0.22
PAS-GRA 12.44wxyz 14.34vw 17.78 19.72 0.31 0.22
216 (120) PH-CON 6.76s 8.00s 9.86 10.62 0.13 0.11
PAS-CON 9.43tu 11.06t 14.38 15.52 0.33 0.24
PAS-GS20 11.21uvwx 11.65tu 16.49 17.51 0.28 0.27
PAS-GS50 13.22xyz 13.68uvw 15.25 15.80 0.55 0.22
PAS-GRA 11.46uvwxy 12.36tuv 15.26 16.46 0.42 0.32
*** *** *** *** *** ***
T
*** ***
D NS NS NS NS
SED 0.471 0.464 2.564 0.862 0.062 0.033
* *
T·D NS NS NS NS
SED 1.053 1.037 5.733 1.927 0.138 0.074
s–z
Within columns, letters with different superscripts differ significantly.
a
Days in parentheses are corresponding days after housing of all heifers, following 90 days grazing and a 6d dietary adjustment period.
*
P < 0.05.
***
P < 0.001.

still tended to have lower pH than other heifers. The pH
of all T increased between d56 and d91 but the increase
was greatest for PAS-GS50 and PAS-GRA heifers, which
like PH-CON and PAS-CON heifers decreased in the
final 30 days, in contrast to PAS-GS20 which increased.
All pH values were between 5.32 and 5.82.
There was a tendency for LD of PH-CON heifers to
have a higher ‘L’ value (P = 0.087), i.e., for LD muscle
to be lighter or ‘paler’ and to have a lower haem pigment
concentration (P = 0.058) than other T. Both ‘L’ and ‘a’
values increased as the experiment progressed.
3.2.4. Relationships between tissue colour and pigment
concentrations
b-Carotene and lutein concentrations separately
explained 42% (P < 0.001) and 36% (P < 0.001) of the var-
iation in s.c. adipose tissue yellowness, respectively, while
total carotenoid concentration explained 46% (P < 0.001)
of the variation. There was a low but significant negative
correlation between haem pigments and muscle lightness,
(r = 0.3; P = 0.009) but haem pigments were not correlated
with muscle redness (r = 0.09; P = 0.45).
3.2.5. Subjective assessment of carcass fat colour
The six pre-experimental heifers, which were fed whole-
crop wheat silage had acceptable carcass fat colour. At
housing following the grazing period, four of the six
PAS- heifers were assessed as bordering on excessive yel-
low colour, but the accompanying PH-CON heifers were
deemed to have acceptable carcass fat colour. On d28
after housing, all carcasses other than the PAS-GRA heif-
ers were acceptable. On d56 after housing, two of the
three heifers in each of PAS-GRA and PAS-GS50 were
assessed as borderline. On d91 after housing, two of the
three PAS-CON and all three of the PH-CON heifers
were acceptable. On the final day, all of the PH-CON
and PAS-CON carcasses were accepted as well as two
of the three PAS-GS20 heifers and one of the three
PAS-GRA carcasses. Thus, the PH-CON were the only
carcasses with a 100% acceptability on every D. When
all carcass fat ‘b’ values from the experiment were
arranged in ascending order and assigned their respective
grades as either ‘acceptable’, ‘borderline’ or ‘unaccept-
able’, the lowest ‘b’ value to be rejected was 10.5 although
the lowest ‘b’ value to be assigned ‘borderline’ status was
 P.G. Dunne et al. / Meat Science 74 (2006) 231–241 237

Table 5
Muscle pH, haem pigment concentration and colour coordinates (lightness (‘L’), redness (‘a’) and saturation (‘C’)) of M. longissimus dorsi (LD)
throughout the experiment
Days (D)a Treatment (T) pHu Total haemb LD colour coordinates
L a C
wxyz
124 (28) PH-CON 5.69 6.03 35.17 12.78 13.57
PAS-CON 5.63vwxyz 5.91 35.14 13.46 14.34
PAS-GS20 5.66vwxyz 6.87 35.31 11.76 12.47
PAS-GS50 5.82z 7.13 35.00 12.73 13.49
PAS-GRA 5.48uvw 5.89 34.18 12.94 13.78
152 (56) PH-CON 5.42u 5.80 35.53 12.90 14.93
PAS-CON 5.49uvwx 7.02 35.04 12.75 16.81
PAS-GS20 5.46uv 6.51 33.67 12.24 12.93
PAS-GS50 5.40u 6.50 34.92 13.57 14.38
PAS-GRA 5.42u 6.26 35.12 13.92 14.77
187 (91) PH-CON 5.67vwxyz 5.77 37.28 12.71 15.16
PAS-CON 5.70xyz 6.84 33.63 12.59 13.24
PAS-GS20 5.52uvwxy 6.41 34.37 13.33 14.05
PAS-GS50 5.79z 7.01 35.31 14.08 15.00
PAS-GRA 5.71yz 6.96 35.53 14.04 14.93
216 (120) PH-CON 5.33u 6.13 37.66 14.61 15.59
PAS-CON 5.49uvwx 6.72 36.79 15.75 16.70
PAS-GS20 5.79z 7.26 34.97 14.10 14.97
PAS-GS50 5.40u 6.27 36.61 15.68 16.72
PAS-GRA 5.32u 6.30 37.23 14.13 14.95
T NS 0.058 0.087 NS NS
*** *** ***
D NS *
SED 0.047 0.333 0.463 0.464 0.653
**
T·D NS NS NS NS
SED 0.106 0.745 1.035 1.038 1.460
u–z
Within columns, letters with different superscripts differ significantly (P < 0.05).
a
Days in parentheses are corresponding days after housing of all heifers, following 90 days grazing and a 6d dietary adjustment period.
b
Total haem pigments, as mg/g muscle tissue (fresh weight basis).
**
P < 0.01.
***
P < 0.001.

10.57. The highest ‘b’ value to be assigned ‘borderline’
status was 11.92.
4. Discussion
4.1. Colour of subcutaneous adipose tissue
The design used was chosen to facilitate determination
of the temporal changes in tissue colour. The grazing strat-
egy was chosen to facilitate accumulation of carotenoids
and concomitant development of yellow colour in adipose
tissue depots, and particularly in the s.c. adipose tissue,
since it is the colour of this depot that is ultimately assessed
by meat graders. Adipose tissue colour data from PAS-
(grazing) heifers slaughtered at housing acted as a bench-
mark for the PAS-CON, PAS-GS20, PAS-GS50 and
PAS-GRA heifers slaughtered subsequently. The ‘b’ value
was chosen to evaluate differences in s.c. adipose tissue col-
our between treatments as the colour of this tissue is most
often described in the literature by its ‘b’ value (Muir et al.,
1998; Strachan et al., 1993; Walker, Warner, & Winfield,
1990; Yang et al., 1993). The ‘C’ values are also presented
as it is claimed that they represent s.c adipose tissue colour
more accurately, as it is perceived visually (Rigg, 1987).
The first hypothesis tested was that feeding a non-grass-
based diet would decrease yellow colour in a time-depen-
dent manner. The decrease in yellowness of s.c. adipose
tissue from PAS-GRA animals upon housing, albeit small,
likely reflects the dilution of carotenoid supply due to con-
centrate supplementation of a restricted grass supply since
the concentration of b-carotene in feeds can be ranked as
follows: grass > grass silage > concentrates (McDonald,
Edwards, Greenhalgh, & Morgan, 1995). This strategy
was necessary since the decline in nutritional value of grass
in autumn would not have allowed adequate growth in this
group (French et al., 2001) and so would have introduced a
confounding effect of differences in carcass weight/fat
deposition. The decline in yellowness over time may also
reflect a change in the carotenoid supply from the grass
as the experiment progressed (Park, Anderson, Walters,
& Mahoney, 1983) but feed carotenoid concentrations were
not measured in this study. The data support the first
hypothesis with respect to PAS-CON and PAS-GS20 in
that adipose tissue from PAS-GRA animals tended to be
238 P.G. Dunne et al. / Meat Science 74 (2006) 231–241
more yellow throughout. In general, feeding of concen-
trates produces s.c. adipose tissue which is less yellow than
that from cattle fed diets containing green forage (Forrest,
1981; Knight, Death, Lambert, & McDougall, 2001; Stra-
chan et al., 1993) even when carcass weight/fatness is sim-
ilar (Moloney, Mooney, Drennan, O’Neill, & Troy, 2000)
reflecting the different carotenoid concentrations in the dif-
ferent feedstuffs. In addition, the fresh grass was an old
permanent pasture with a mixture of grass (Lolium perenne,
Poa trivialis, Poa annua) and clover (Trifolium repens) spe-
cies, while the silage used (typical of GS made in Ireland)
was from grass harvested from a predominantly perennial
ryegrass sward in mid May and conserved immediately in
a concrete pit. Therefore, botanical differences between
the swards could contribute to the observed differences
between the grass and GS based-diets (Bauernfeind,
Adams, & Marusick, 1981; Park et al., 1983) together with
the differing proportions of forage in the rations. The dif-
ferences might also reflect dilution of carotenoids accumu-
lated during grazing by subsequent fat deposition (Knight
et al., 2001) and possible mobilisation of stored carotenoids
(see below).
The second hypothesis tested was that a decreased pro-
portion of GS in the diet would lead to a decreased yellow
colour in the s.c. adipose tissue in a time-dependent man-
ner. Averaged across slaughter dates from the end of the
grazing period, ‘b’ value was linearly (P < 0.05) related to
the proportion of GS in the diet (0, 200 and 500 g/kg
DM) supporting the first part of this hypothesis. With
respect to the temporal pattern, the crucial period in rela-
tion to the GS treatments was between housing and d28
indoors since s.c. adipose tissue ‘b’ value of both PAS-
GS50 and PAS-GS20 decreased in this interval. Due to
the positive relationship between carcass fat ‘b’ value/yel-
lowness and carotenoid concentration (Knight, Death,
Boom, & Litherland, 1998; Strachan et al., 1993; Zhou,
Yang, & Tume, 1993) and confirmed in this study, it would
be expected that the carotenoid concentration would also
decrease, as was seen in this study. Both PAS-GS50 and
PAS-GS20 were fed the same silage and the decline in
s.c. adipose tissue ‘b’ value coincident with a decline in
carotenoid concentration during the initial 28 days after
housing and diet implementation, may reflect differences
in carotenoid intake per se.
Diets were designed to produce similar growth rates and
carcass fatness across each T so it is unlikely that differ-
ences in carcass fatness played a major role alone in influ-
encing the whiter s.c. adipose tissue colour at a particular
slaughter time. Covariate adjustment of s.c. adipose tissue
colour coordinates using KCF weight, as g/kg carcass, also
corroborated this as no colour coordinate was significantly
affected. Carcass weight increased throughout the study
and since carotenoids are distributed within adipose tissue,
any increase in the relative proportion of adipose tissue
resulting from accumulation of triacylglycerols, without a
concomitant increase in carotenoid intake, would be
expected to dilute carotenoids already present and decrease
yellowness. This clearly occurred with PAS-CON in the
present study. In contrast, the ‘b’ value of the GS-based
diets increased subsequent to d28 post-housing.
Little information is available on the mobilisation of
carotenoids from adipose tissue. Knight, Wyeth, Ridland,
& Death (1994), based on the carotene concentrations in
blood plasma of heifers after removal from pasture, con-
cluded that there were two ‘pools of carotene’ in the body;
one which was depleted rapidly in the blood and liver and
one in the adipose tissue which was depleted much more
slowly. The change in carotenoid concentrations and yel-
lowness of adipose tissue for PAS-CON animals, in partic-
ular, supports the latter conclusion. The model describing
the change in ‘b’ value predicts that 26 days approximately
of concentrate consumption would be required to undo the
yellowing effect of grazing. However, that adipose tissue
from PAS-CON never achieved the same ‘b’ value as that
from PH-CON indicates that once adipose tissue caroten-
oid concentration has declined to some threshold, either
no further mobilisation occurs or the rate of mobilisation
becomes extremely slow. Since the ‘b’ value of PAS-GRA
cattle did not decrease post-housing, the change in diet
composition may have triggered carotenoid mobilisation
in the other three diets. After the threshold of mobilisation
had been reached, carotenoid repletion seemed to occur for
the GS-based diets which reflected the degree of carotenoid
consumption.
The reason for this is unclear but Knight et al. (1994)
observed a decrease in plasma carotenoids after heifers
were removed from pasture, despite feeding of a pelleted
diet containing 500 mg added carotene per kg, comparable
to the carotenoid content of pasture. Such a decrease might
facilitate mobilisation of adipose tissue carotenoids. It is
possible that the resulting increase in plasma carotenoid
concentration may inhibit absorption of dietary carote-
noids and carotenoid accumulation can not resume until
the mobilised carotenoids have been catabolised. In sup-
port of this Knight, Death, Muir, Ridland, & Wyeth
(1996) suggested that vitamin A supplementation had no
effect on fat colour when plasma concentrations were high.
Similarly, Knight et al. (1996) reported that feeding a low
carotenoid diet to steers reduced fat carotenoid concentra-
tion and colour after 70 days, but continuing the diet for a
further 42 days had no additional effect on carotenoid con-
centration in fat, despite a 3.1 mm increase in fat depth.
According to Knight et al. (1996) these results indicated
that after the initial loss of carotenoids from the fat, no fur-
ther loss occurred, either because the plasma carotenoid
concentration was too high to allow further loss from the
fat or the carotenoid pool in the fat was non-labile. Further
research is required to elucidate the underlying mechanism
of carotenoid mobilisation and repletion.
When subjectively examining sections of s.c. adipose tis-
sue it was clear that even within a 10 · 10 cm section there
could sometimes be considerable variation in the visual
appearance with localised areas of intense yellow colour
which may have introduced some random variation. More-
 P.G. Dunne et al. / Meat Science 74 (2006) 231–241
over, while the carotenoids represent the constituents
which have the largest single influence on adipose tissue
colour, other chemical and structural components of the
adipose tissue contribute to perception of colour also. Irie
(2001) discovered that absorbance of deoxy- and met-
haemoglobin (presumably from peripheral adipose tissue
capillarisation) as well as carotenoids affected spectropho-
tometer absorbance, in addition to the reflectance, trans-
mittance and fluorescence of lipids and the reflectance
and fluorescence of connective tissue and cell membranes.
Despite this, there was a clear distinction between T in
terms of s.c. adipose tissue carotenoid (both b-carotene
and lutein) concentrations.
Comparison of objective ‘b’ values with corresponding
subjective grades in the present study revealed that ‘b’ val-
ues above 12 were regarded as ‘unacceptable’ or ‘exces-
sively yellow’, values below 10.5 were ‘acceptable’ and
those between 10.5 and 12 were borderline. This assessment
was more stringent than in a previous study in the same
factory when 50% of accepted carcasses had a ‘b’ value
>12 (Dunne et al., 2004). This likely reflects different mar-
ket requirements in the course of the two studies. None of
the PAS-GS50 heifers slaughtered at d28 displayed exces-
sively yellow carcass fat, although those slaughtered on
subsequent dates did. This comparative approach also sup-
ported the 100% acceptability of the PH-CON heifers since
all had ‘b’ values below 12 (two-thirds had ‘b’ value below
9), thus confirming that the appropriate strategy to guaran-
tee ‘white’ carcass fat was to omit grazing and to feed con-
centrates without silage inclusion.
The rationale underlying the modelling approach and
upon which the experimental design was based, was to
develop the ability to predict the yellowness of the s.c. adi-
pose tissue depot on any given day or alternatively, to esti-
mate the number of days required to reach any particular
yellowness value. The inconsistencies in subjective colour
assessment compared with objective thresholds, emphasise
the importance and usefulness of this approach. Such a
novel approach has been unreported in the literature. As
an example, most cattle finished in Ireland are offered a diet
based on GS and finishing (of steers) typically lasts for
between 80 and 180 days to ensure adequate carcass weight
(Keane & Drennan, 1991). The models predict that in a fin-
ishing period of 150 days, heifers on either PAS-GS50 or
PAS-GS50 would have a ‘b’ value >15.0 and would not
be acceptable for the ‘‘white fat’’ market. It is recognised
that the large standard errors for the intercept values some-
what diminish the predictive ability of the models.
4.2. Colour of muscle
Muscle colour is affected by the pigment concentration
but also by light scatter at its surface (MacDougall, 1982)
and pH is critical in this regard. Above a pH of 5.8, beef
becomes progressively darker and beef with a pH in excess
of 5.8–5.9 is considered to be ‘dark-cutting’ or ‘dark, firm
and dry’ (Pethick et al., 2000). As pH values were normal,
239
‘dark-cutting’ beef was not an issue with the heifers in this
study.
The third hypothesis tested in this study was that
removal of the grazing opportunity would result in paler
meat in the beef production system used. The PH-CON
heifers, being housed throughout, did not engage in the for-
aging activity of the grazing animals. The increase in LD
haem of the grazing groups may have been caused by the
sustained, low intensity physical activity involved in forag-
ing. A similar trend was seen as a result of the exercise
model applied by Dunne, O’Mara, Monahan, & Moloney
(2005). Physical activity has been proposed as an explana-
tion for darker muscle colour in young bulls that were per-
mitted to graze compared with their counterparts confined
indoors (Vestergaard, Oksbjerg, & Henckel, 2000),
although Moloney, Fallon, Mooney, & Troy (2004) sug-
gested that the observed results could equally have been
attributed to either age or pH differences. In the present
study, the difference in haem pigment concentration during
the grazing period was not reflected in differences in muscle
colour at the end of the grazing period. This supports the
conclusions of French et al. (2001, 2000) that grazed grass
and concentrates result in similar muscle colour. The lack
of an effect of slaughter date indicates that the effect of
grazing on haem pigment concentrations was maintained
throughout the study. The tendency towards lighter muscle
in PH-CON compared to PAS-CON is not sufficiently
strong to support the hypothesis above.
In contrast to the poor relationships between muscle
lightness, redness and haem pigments in this study, Gil
et al. (2001) reported significant negative correlations
between haem pigments and muscle lightness in five of
seven native Spanish cattle breeds and significant positive
correlations between haem pigments and redness. It would
be expected that as haem pigments increased, muscle would
become darker (‘L’ value would decrease) and more red (‘a’
value would increase) since an increase in haem pigments
would result in an increase in haem iron, as iron is com-
plexed in the haem moiety of these pigments, of which
myoglobin is dominant (Oellingrath, Iversen, & Skrede,
1990). The poor relationships between muscle pigments
and lightness and redness in the present study may have
been due to pigment analysis being performed on a
2.5 cm thick core from the centre of the steak and not on
a slice from the steak surface, where colour was measured.
Thus, while the oxygenated pigments in a thin slice from
the surface would be expected to relate closely to surface
colour, the un-oxygenated pigments within the muscle
may not.
5. Conclusions
While concentrate feeding after a three month period at
pasture led to a time-dependent decrease in s.c. adipose tis-
sue yellowness relative to grass feeding, inclusion of GS in
the diet of heifers, led to a significant reduction in yellow-
ness in the first 28 days after housing and but not to a con-
240 P.G. Dunne et al. / Meat Science 74 (2006) 231–241
sistent reduction in yellowness of this tissue thereafter.
Removal of the grazing opportunity (and forage) from
the diet of PH-CON heifers led to lower haem pigment
concentrations but did not have a definite effect on muscle
colour.
6. Implications
This research demonstrates that grass feeding tends to
produce the most yellow, and concentrate feeding the least
yellow, carcass fat. It is possible to house continental cross-
bred heifers, at an average age of 16 months, following a
three month grazing period and to feed GS, at levels of
up to 50% of dietary DM for 28 days after housing without
causing excessive yellow colour in s.c. adipose tissue, when
subjectively assessed. For short-term feeding this is a more
economical strategy than feeding concentrates. For longer
finishing periods, GS is best removed from the diet but lit-
tle additional benefit was seen beyond 56 days consump-
tion of concentrates. Since heifers that were permanently
housed and fed concentrates always had whiter fat than
animals that grazed and were then fed concentrates, this
strategy would be most appropriate for markets that have
a particularly stringent requirement for white fat. The data
indicate that a pre-housing grazed period has little impact
on muscle colour when animals are offered a concentrate
diet subsequently.
Acknowledgement
This research was funded under the National Develop-
ment Plan of the Irish Government, 2000–2006. Financial
support to P.G. Dunne under the Walsh Fellowship Pro-
gramme of Teagasc, the Irish Agriculture and Food Devel-
opment Authority, is acknowledged. The authors
acknowledge the assistance and cooperation of manage-
ment and staff at Meadow Meats, Rathdowney, Co. Laois,
Ireland. The skilled technical assistance of V. McHugh is
appreciated. M. Cottrill is thanked for feeding and care
of the animals. N. Tonge and staff at Grange Laboratories
are thanked for analysis of feedstuffs.
Statistical advice from Dr. F. Bannon, Department of
Statistics, University College Dublin is gratefully
acknowledged.
References
AOAC (1990). Official Method 990.03. Determination of the crude protein
content of animal feedingstuffs. Arlington, VA: AOAC.
Anon (1999). Bord Bia Meat News. Lower Mount St., Dublin, Ireland:
Bord Bia (Irish Food Board), Clanwilliam Court.
Anon (1981). Community scale for the classification of carcasses of adult
bovine animals. E.C. No. 1208/81 and 2930/81. Luxembourg: Office for
Official Publications of the European Communities.
Bauernfeind, J. C., Adams, C. R., & Marusick, W. I. (1981). Carotenes
and other vitamin A precursors in animal feed. In J. C. Bauernfeind
(Ed.), Carotenoids as colourants and vitamin A precursors. New York:
Academic Press Inc.
Drennan, M. J., Keane, M. G., & Dunne, L. (1995). Outline of beef
production in Ireland. In: Extensification of Beef and Sheep Produc-
tion on Grassland. Occasional Publication No.2 of Concerted Action
AIR3-CT93-0947, Paris, Nov 22–24, p. 87–94.
Dunne, P. G., O’Mara, F. P., Monahan, F. J., & Moloney, A. P. (2004).
Colour of subcutaneous adipose tissue and muscle of Irish beef
carcasses destined for the Italian market. Irish Journal of Agricultural
and Food Research, 43, 217–226.
Dunne, P. G., O’Mara, F. P., Monahan, F. J., & Moloney, A. P. (2005).
Colour of muscle from 18-month-old steers given long-term daily
exercise. Meat Science, 71, 219–229.
Forrest, R. J. (1981). Effect of high concentrate feeding on the carcass
quality and fat colouration of grass reared steers. Canadian Journal of
Animal Science, 61, 575–580.
French, P., O’Riordan, E. G., Monahan, F. J., Caffrey, P. J., Mooney, M.
T., Troy, D. J., et al. (2001). The eating quality of meat of steers
offered grass and/or concentrates. Meat Science, 57, 379–386.
French, P., O’Riordan, E. G., Monahan, F. J., Caffrey, P. J., Mooney, M.
T., & Moloney, A. P. (2000). Meat quality of steers finished on autumn
grass, grass silage or concentrate-based diets. Meat Science, 56,
173–180.
Gil, M., Serra, X., Gispert, M., Ángels Oliver, M., Sañudo, C., Panea, B.,
et al. (2001). The effect of breed-production systems on the myosin
heavy chain *1, the biochemical characteristics and the colour
variables of Longissimus thoracis from seven Spanish beef cattle
breeds. Meat Science, 58, 181–188.
Irie, M. (2001). Optical evaluation of factors affecting appearance of
bovine fat. Meat Science, 57, 19–22.
Keane, M. G., & Drennan, M. J. (1991). Two-year old beef production for
Friesian and Friesian cross steers. Beef Series No. 9, Teagasc, Dublin,
Ireland.
Knight, T. W., Death, A. F., Lambert, M. G., & McDougall, D. B. (2001).
The rate of reduction in carotenoid concentration in fat of steers fed a
low carotenoid ration, and the role of increasing carcass fatness.
Australian Journal of Agricultural Research, 52, 1023–1032.
Knight, T. W., Death, A. F., Boom, C. J., & Litherland, A. J. (1998). The
relationship between carotenoid concentration and fat colour in beef
carcasses. Proceedings of the New Zealand Society of Animal Produc-
tion, 58, 256–258.
Knight, T. W., Death, A. F., Muir, P. D, Ridland, M., & Wyeth, T. K.
(1996). Effect of dietary vitamin A on plasma and liver CCs and fat
colour in Angus and Angus crossbred cattle. New Zealand Journal of
Agricultural Research, 39, 281–292.
Knight, T. W., Wyeth, T. K., Ridland, M., & Death, F. (1994). Effects of
dietary carotene content on mean values and rankings of heifers for
plasma carotene concentrations. New Zealand Journal of Agricultural
Research, 37, 159–165.
Krzywicki, K. (1982). The determination of haem pigments in meat. Meat
Science, 7, 29–36.
Lee, J. (1988). Forages. Livestock Production Science, 19, 13–46.
MacDougall, D. B. (1982). Changes in the colour and opacity of meat.
Food Chemistry, 9, 75–88.
McDonald, P., Edwards, R. A., Greenhalgh, J. F. D., & Morgan, C. A.
(1995). Animal Nutrition (Fifth ed.). Harlow: Prentice Hall.
McLaren, K. (1987). Colour space, colour scales and colour difference. In
R. McDonald (Ed.), Colour physics for industry (pp. 107). Bradford,
UK: Dyers Company Publications Trust.
Moloney, A. P., Fallon, R. J., Mooney, M. T., & Troy, D. J. (2004). The
quality of meat and fatness of bulls offered ad libitum concentrates,
indoors or at pasture. Livestock Production Science, 87, 271–276.
Moloney, A. P., Mooney, M. T., Drennan, M. J., O’Neill, B., & Troy, D.
J. (2000). Fat colour and the quality of meat from beef cattle offered
grass silage or concentrate-based rations. Irish Journal of Agricultural
and Food Research, 39, 152.
Muir, P. D., Smith, N. B., Wallace, G. J., Cruickshank, G. J., & Smith, D.
R. (1998). The effect of short-term grain feeding on liveweight gain and
beef quality. New Zealand Journal of Agricultural Research, 41,
517–526.
 P.G. Dunne et al. / Meat Science 74 (2006) 231–241
Oellingrath, I. M., Iversen, A., & Skrede, G. (1990). Quantitative
determination of myoglobin and haemoglobin in beef by high-
performance liquid chromatography. Meat Science, 28, 313–320.
O’Kiely, P., Power, R., & O’Donnell, C. (1993). Grass conservation,
storage and feeding in Ireland. Farm & Food, 3, 4–7.
O’Riordan, E. G., & O’Kiely, P. (1996). Potential of beef production
systems based on grass. Irish Grassland Animal Production Association,
30, 185–217.
Park, Y. M., Anderson, M. J., Walters, J. L., & Mahoney, A. W. (1983).
Effects of processing methods and agronomic variables on carotene
contents in forages and predicting carotene in alfalfa hay with near-
infra-red-reflectance spectroscopy. Journal of Dairy Science, 66,
235–245.
Pethick, D. W., Cummins, L., Gardner, G. E., Jacobs, R. H., Knee, B. W.,
McDowell, M., et al. (2000). The regulation of glycogen level in the
muscle of ruminants by nutrition. Proceedings of the New Zealand
Society of Animal Production, 60, 94–98.
Priolo, A., Micol, D., & Agabriel, J. (2001). Effects of grass feeding
systems on ruminant meat colour and flavour. Animal Research, 50,
185–200.
Pupin, A. M., Dennis, M. J., & Toledo, M. C. F. (1999). HPLC analysis of
carotenoids in orange juice. Food Chemistry, 64, 269–275.
Rigg, B. (1987). Colorimetry and the CIE system. In R. McDonald (Ed.),
Colour Physics for Industry (pp. 63–96). Bradford, UK: Dyers
Company Publications Trust.
Strachan, D. B., Yang, A., & Dillon, R. D. (1993). Effect of grain feeding
on fat colour and other carcass characteristics in previously grass-fed
241
Bos indicus steers. Australian Journal of Experimental Agriculture, 33,
269–273.
Strange, E. D., Benedict, R. C., Gugger, R. E., Metzger, V. G., & Swift, C.
E. (1974). Simplified methodology for measuring meat colour. Journal
of Food Science, 39, 988–992.
Tilley, J. N. A., & Terry, R. A. (1963). A two-stage technique for the in-
vitro digestion of a forage crop. Journal of the British Grassland
Society, 18, 104–111.
Vestergaard, M., Oksbjerg, N., & Henckel, P. (2000). Influence of feeding
intensity, grazing and finishing feeding on muscle fibre characteristics
and meat colour of semitendinosus, longissimus dorsi and supraspinatus
muscles of young bulls. Meat Science, 54, 177–185.
Walker, P. J., Warner, R. D., & Winfield, C. G. (1990). Sources of
variation in subcutaneous fat colour of beef carcasses. Proceedings of
the Australian Society of Animal Production, 18, 416–419.
Warriss, P. D. (1979). The extraction of haem pigments from fresh meat.
Journal of Food Technology, 14, 75–80.
Yang, A., McLennan, S. R., Armstrong, J., Larsen, T. W., Shaw, F. D., &
Tume, R. K. (1993). Effect of short-term grain feeding on bovine body
fat colour: a cautionary note. Australian Journal of Agricultural
Research, 44, 215–220.
Yang, A., Larsen, T. W., & Tume, R. K. (1992). Carotenoid and retinol
concentrations in serum, adipose tissue and liver and carotenoid
transport in sheep, goats and cattle. Australian Journal Agricultural
Research, 43, 1809–1817.
Zhou, G. H., Yang, A., & Tume, R. K. (1993). A relationship between bovine
fat colour and fatty acid composition. Meat Science, 35, 205–212.