Genetic correlations between wool traits and meat quality traits in Merino sheep1

S. I. Mortimer,*†2 S. Hatcher,†‡ N. M. Fogarty,†‡ J. H. J. van der Werf,†§ D. J. Brown,†#

A. A. Swan,†# R. H. Jacob,†║ G. H. Geesink,†§ D. L. Hopkins,†¶ J. E. Hocking Edwards,† **
E. N. Ponnampalam,† †† R. D. Warner,† †† K. L. Pearce,† ‡‡ and D. W. Pethick† ‡‡
*NSW Department of Primary Industries, Agricultural Research Centre, Trangie, NSW 2823, Australia; †CRC
for Sheep Industry Innovation, University of New England, Armidale, NSW 2351, Australia; ‡NSW Department
of Primary Industries, Orange Agricultural Institute, Orange, NSW 2800, Australia; §School of Environmental and
Rural Science, University of New England, Armidale, NSW 2351, Australia; #Animal Genetics and Breeding
Unit,3 University of New England, Armidale, NSW 2351, Australia; ║Department of Agriculture and Food WA,
Baron Hay Court, South Perth, WA 6151, Australia; ¶NSW Department of Primary Industries, Centre for
Red Meat and Sheep Development, Cowra, NSW 2794, Australia; **South Australian Research and Development
Institute, Naracoorte, SA 5271, Australia; ††Agriculture Victoria, Department of Economic Development,
Jobs, Transport and Resources, Attwood, VIC 3049, Australia; ‡‡Murdoch University, Murdoch, WA 6151, Australia

ABSTRACT: Genetic correlations between 29 wool
production and quality traits and 25 meat quality and
nutritional value traits were estimated for Merino sheep
from an Information Nucleus (IN). Genetic correlations
among the meat quality and nutritional value traits are
also reported. The IN comprised 8 flocks linked geneti-
cally and managed across a range of sheep production
environments in Australia. The wool traits included
over 5,000 yearling and 3,700 adult records for fleece
weight, fiber diameter, staple length, staple strength,
fiber diameter variation, scoured wool color, and visual
scores for breech and body wrinkle. The meat quality
traits were measured on samples from the M. longis-
simus thoracis et lumborum and included over 1,200
records from progeny of over 170 sires for intramus-
cular fat (IMF), shear force of meat aged for 5 d (SF5),
24 h postmortem pH (pH24LL; also measured in the
semitendinosus muscle, pH24ST), fresh and retail meat
color and meat nutritional value traits such as iron and
zinc levels, and long-chain omega-3 and omega-6
polyunsaturated fatty acid levels. Estimated heritabili-
ties for IMF, SF5, pH24LL, pH24ST, retail meat color
lightness (L*), myoglobin, iron, zinc and across the
Key words: eating quality, genetic correlations, meat quality, Merino sheep, nutritional value, wool
range of long-chain fatty acids were 0.58 ± 0.11, 0.10 ±
0.09, 0.15 ± 0.07, 0.20 ± 0.10, 0.59 ± 0.15, 0.31 ± 0.09,
0.20 ± 0.09, 0.11 ± 0.09, and range of 0.00 (eicosapen-
taenoic, docosapentaenoic, and arachidonic acids) to
0.14 ± 0.07 (linoleic acid), respectively. The genetic
correlations between the wool production and meat
quality traits were low to negligible and indicate that
wool breeding programs will have little or no effect on
meat quality. There were moderately favorable genetic
correlations between important yearling wool produc-
tion traits and the omega-3 fatty acids that were reduced
for corresponding adult wool production traits, but
these correlations are unlikely to be important in wool/
meat breeding programs because they have high SE,
and the omega-3 traits have little or no genetic variance.
Significant genetic correlations among the meat quality
traits included IMF with SF5 (-0.76 ± 0.24), fresh meat
color L* (0.50 ± 0.18), and zinc (0.41 ± 0.19). Selection
to increase IMF will improve meat tenderness and col-
or which may address some of the issues with Merino
meat quality. These estimated parameters allow Merino
breeders to combine wool and meat objectives without
compromising meat quality.

© 2017 American Society of Animal Science. All rights reserved. J. Anim. Sci. 2017.95:4260–4273
doi:10.2527/jas2017.1628
1This project is from the Information Nucleus program of the 2Corresponding author: sue.mortimer@dpi.nsw.gov.au
Cooperative Research Centre for Sheep Industry Innovation which 3Animal Genetics and Breeding Unit (AGBU) is a joint venture

is supported by the Australian Government’s Cooperative Research of NSW Department of Primary Industries and the University of
Centres Programme, Australian Wool Innovation Ltd. and Meat & New England.
Livestock Australia. The authors acknowledge the contributions Received April 12, 2017.
of the many research and technical staff from 7 research agencies Accepted July 27, 2017.
and support provided by Australian sheep breeders.

4260
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 Correlations between wool and meat quality
INTRODUCTION
Quality and the human health attributes of sheep
meat are vitally important to consumers (Pethick et al.,
2011). Continued selection emphasis on leanness may
adversely affect eating quality (Hopkins et al., 2005b;
Hopkins et al., 2007b; Pannier et al., 2014a), intramus-
cular fat content (Hopkins et al., 2005b; Hopkins et al.,
2007b; Pannier et al., 2014c), and iron levels (Pannier
et al., 2014b). Recent research outcomes are also shift-
ing the focus of ram breeders and commercial produc-
ers to a more balanced approach that includes consumer
needs (Pethick et al., 2014). Genetic variation exists for
at least some meat quality traits associated with eat-
ing quality and the nutritional value of meat (see re-
views by Hopkins et al. 2011 and Mortimer et al. 2014).
However, there are few published estimates of genetic
and phenotypic correlations among meat quality traits
or correlations with other production traits, which are
essential for development of effective sheep breeding
programs that also consider meat quality. The Merino is
the major sheep breed in Australia and contributes the
majority of genes to meat production for domestic and
export markets through purebred and crossbred lambs,
as well as being the leading source of fine apparel wool
sold globally. Other breeds, such as the Dohne Merino
and Targhee, perform a similar role in contributing to
both wool and meat production systems worldwide. In
Merinos, there have also been particular concerns with
aspects of meat quality such as high ultimate pH and re-
tail discoloration (Hopkins and Fogarty, 1998; Gardner
et al., 2010; Jacob and Pethick, 2014) that potentially
could be addressed through breeding. This paper re-
ports genetic correlations between wool traits and traits
associated with meat quality including eating quality
and its nutritional value, as well as correlations among
the meat quality traits. Earlier studies have reported
the genetic correlations with carcass composition traits
(Mortimer et al., 2017a; Mortimer et al., 2017b).
MATERIALS AND METHODS
Animals
Data were recorded on the Merino progeny born
during 2007 to 2011 in the Information Nucleus (IN;
Fogarty et al., 2007; van der Werf et al., 2010) breed-
ing program of the Cooperative Research Centre for
Sheep Industry Innovation (Sheep CRC, Armidale
Australia). The IN comprised 8 genetically linked flocks
located in each of the major sheep growing areas of
Australia (Armidale NSW, Trangie NSW, Cowra NSW,
Rutherglen VIC, Hamilton VIC, Struan SA, Turretfield
SA, and Katanning WA). The design of the IN, includ-
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4261
ing procedures used to select the sires for artificial in-
semination of the base ewes and for management, are
described by van der Werf et al. (2010) and Geenty et al.
(2014). The data were derived from records of a max-
imum of 9,135 progeny born to 184 Merino sires and
4,614 Merino dams. At least 50% of the sires were rep-
resented at all sites, while all sires were represented at 2
sites (Armidale and Katanning). Depending on the site,
the Merino dams were from pedigreed (Cowra, Trangie,
Struan, Turretfield, and Katanning) and commercial
flocks (Armidale, Rutherglen, and Hamilton). The source
flocks for the Merino dams varied with site, though
the dams were from the same source flocks at Cowra
and Trangie, as was the case for Struan and Turretfield
(Geenty et al., 2014). The lambs were tail docked and the
males castrated at marking (7 to 43 d). After weaning (av-
erage age of 90.7 d [SD 9.7]), the lambs at each site were
managed to achieve target growth rates of 150 g/d. Half
of the wether lambs (balanced for sire) were randomly
allocated to groups for slaughter in their first year after
finishing. Each group was scheduled to be slaughtered
after reaching a target carcass weight of 21.5 kg on aver-
age. Yearling and adult wool production measurements
were recorded on the ewe lambs and the remainder of the
wethers. The lambs usually grazed the extensive pastures
available at the sites, but were supplemented with grain,
hay, or feedlot pellets when the pasture supply was re-
stricted. The sites were managed by Sheep CRC partner
organizations, with research and data collection activi-
ties based on a common protocol at each IN site. All ac-
tivities were approved by the Animal Ethics Committee
of the organizations responsible for managing lambs at
each site. Management of all animals was conducted ac-
cording to the Australian Code for the Care and Use of
Animals for Scientific Purposes (NHMRC, 2013).
Wool Traits
The ewes and wethers were shorn as yearlings (300
to 400 d) and adults (> 540 d), when greasy fleece weight
(GFW) was recorded (i.e. yearling GFW [yGFW] and
adult GFW [aGFW]). Mid-side wool samples were
measured by a commercial testing laboratory (AWTA,
2000) for the following traits: washing yield (YLD),
clean fleece weight (CFW), mean fiber diameter (FD),
FD SD (FDSD), FD CV (FDCV), staple strength (SS),
staple length (SL), and mean fiber curvature (CUR).
Wool color measurements were carried out on the clean
scoured and carded samples by the testing laboratory for
brightness (Y) and yellowness (Y-Z). Within 1 mo after
shearing (yearling and adult), visual traits were scored
for breech cover (BCOV), breech wrinkle (BRWR),
and body wrinkle (BDWR) using an industry standard
1 to 5 diagrammatic scale (1 is the least expression and
 4262 Mortimer et al.

5 is the most expression of the trait; AWI and MLA, Table 1. The number of records, mean, standard devi-
2013). Breech wrinkle was also assessed at marking ation, and number of sires and dams for meat traits1
(mBRWR). Yearling wool production, wool quality, n Mean SD Sires Dams
and wool color traits were recorded on over 5,000 ani- Meat quality
mals while over 3,600 animals were recorded as adults. IMF (%) 1,236 4.6 1.14 177 1110
The number of records available for the visual traits SF5 (N) 1,135 31.04 11.70 172 1024
ranged from over 3,300 for yearling scores to 6,035 for pH24LL 1,292 5.72 0.138 177 1162
pH24ST 1,295 5.84 0.228 177 1164
mBRWR. Over 1,800 records were available for the
ICDH (μmol/min/g tissue) 727 5.21 1.69 107 664
adult scores. Numbers of adult wool measurements
Myo (mg/g tissue) 1,292 7.45 2.22 177 1161
were reduced due to adult traits not being recorded on
cfa* 1,262 18.54 2.32 177 1137
some cohorts of animals, as well as some wethers be- cfb* 1,264 3.64 3.78 177 1139
ing both recorded for yearling wool and visual traits cfL* 1,262 34.07 3.23 177 1137
and slaughtered for recording of carcass measurements reta* 703 15.77 2.10 160 628
(Mortimer et al., 2017b). Further details of the measure- retb* 703 17.03 2.09 160 628
ments including the means and standard deviations and retL* 703 35.55 3.40 160 628
the genetic variances for each of the yearling and adult retOxy/Met 703 3.19 0.698 160 628
wool traits are reported by Mortimer et al. (2017a). reth 702 47.20 3.21 160 627
retC* 703 23.29 2.70 160 628
Nutritional value
Meat Quality Traits
IRON (mg/kg wet tissue) 1,289 22.1 3.92 177 1159
ZINC (mg/kg wet tissue) 1,290 25.9 4.75 177 1160
Lambs were slaughtered at commercial abattoirs and
EPA (mg/100g tissue) 1,277 14.7 8.81 177 1147
all carcasses were subjected to medium voltage electrical
DPA (mg/100g tissue) 1,279 24.5 8.84 177 1149
stimulation and trimmed according to AUS-MEAT spec- DHA (mg/100g tissue) 1,279 7.3 3.50 177 1149
ifications (AUS-MEAT, 2006). At slaughter, hot carcass EPA+DHA (mg/100g tissue) 1,277 22.1 11.76 177 1147
weight was recorded and a sample (1 g) taken from the EPA+DPA+DHA (mg/100g tissue) 1,277 46.6 19.86 177 1147
M. longissimus thoracis et lumborum (LL) and frozen LA (mg/100g tissue) 1,277 161.2 41.28 177 1147
in liquid nitrogen within 2 h of death for subsequent as- ARA (mg/100g tissue) 1,279 55.3 16.64 177 1149
say of isocitrate dehydrogenase activity (ICDH) accord- LA+ARA (mg/100g tissue) 1,277 216.4 52.96 177 1147
ing to a procedure described by Gardner et al. (2006). 1IMF = intramuscular fat; SF5 = shear force after 5 days ageing; pH24 =
Carcasses were chilled overnight (3°C–4°C) and then pH at 24 h after slaughter for LL and ST muscles; ICDH = isocitrate de-
cut between the 12th and 13th rib. The LL was exposed hydrogenase activity; Myo = myoglobin concentration; cfa* = fresh meat
redness; cfb* = fresh meat yellowness; cfL* = fresh meat lightness; reta* =
to the air at an ambient temperature for 30–40 min and retail display meat redness; retb* = retail display meat yellowness; retL* =
the meat color was measured (Warner et al., 2010) us- retail display meat lightness; retOxy/Met = retail display meat oxy/met value;
ing Minolta Chroma meters (Models CR-300 and CR- reth = retail display meat hue [h = tan-1(b*/a*)]; retC* = retail display meat
chroma (C* = [a*2 + b*2]0.5); IRON = iron content; ZINC = zinc content;
400) set on the L*, a*, b* system, where L* (cfL*) mea- EPA = eicosapentaeonic acid content; DPA = docosapentaeonic acid content;
sures relative lightness, a* (cfa*) relative redness, and DHA = docosahexaeonic acid content; EPA+DHA = sum of EPA and DHA;
b* (cfb*) relative yellowness. Three replicate measure- EPA+DPA+DHA = sum of EPA, DPA and DHA; LA = linoleic acid content;
ARA = arachidonic acid content; LA+ARA = sum of LA and ARA.
ments were taken at different positions on the exposed LL
surface and an average value used for analysis. The pH
of the LL (pH24LL) and of the M. semitendinosus (ST; (LA; 18:2n-6), and arachidonic acid (ARA; 20:4n-6;
pH24ST) were measured at approximately 24 h after Ponnampalam et al., 2010). Myoglobin content (Myo)
slaughter using a number of different pH meters linked was measured on a sample (1 g) taken from the loin using
to pH electrodes calibrated at chiller temperatures (3°C methods described by Trout (1991). A sample (50 g) was
–4°C; Pearce et al., 2010). The lumbar portion of the LL also taken for measurement of intramuscular fat (IMF)
muscle was excised from the carcass 24 h postmortem using a Technicon Infralyser 450 and near infrared proce-
and after removal of subcutaneous fat and epimysium, dure (Perry et al., 2001). For shear force testing (SF5), a
two 40 g samples of diced muscle were collected, frozen, section of the LL (65 g) was taken 5 cm from the cranial
and stored at -18°C. One sample was used to measure end (at random from a side of the carcass), aged for 5 d
iron levels (IRON) and zinc levels (ZINC; Pannier et at 3°C–4°C, and stored frozen after allocation at random
al., 2010). The other sample was used to measure a range (balanced for sire) for transport to 1 of 2 testing labo-
of fatty acids including the long-chain omega-3 fatty ac- ratories managed by Sheep CRC partner organizations.
ids eicosapentaenoic acid (EPA; 20:5n-3), docosapen- These samples were then cooked from frozen for 35 min
taenoic acid (DPA; 22:5n-3), and docosahexaenoic acid in plastic bags at 71°C in a water bath and tested using a
(DHA; 22:6n-3) and omega-6 fatty acids, linoleic acid Lloyd texture analyzer (Model LRX; Lloyd Instruments,

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 Correlations between wool and meat quality
Hampshire, UK) with a Warner-Bratzler-type shear blade
fitted as described by Hopkins et al. (2010).
Meat color stability under simulated retail display
for 2 d was measured on a 3-cm slice from the cra-
nial end of the LL which had been vacuum packed and
aged for 5 d, with further details provided by Jacob et
al. (2014). These data included color measurements
of L* (retL*), a* (reta*), b* (retb*), and oxymyoglo-
bin/metmyoglobin value (retOxy/Met). Psychometric
hue angle (reth) was calculated as tan-1 (b*/a*) and
psychometric chroma (retC*) as (a*2 +b*2)0.5 (Hunt
et al., 1991). Meat color stability traits were recorded
on samples from animals at 5 sites only.
The number of records for each of the meat quality
traits ranged from 703 to 1,295, with the number of re-
cords, their means. and SD together with the number of
sires and dams represented in the data shown in Table 1.
Statistical Analysis
Fixed effects, variance components, and genetic
parameters were estimated using general linear mixed
models and restricted maximum likelihood methods with
ASReml software (Gilmour et al., 2015). Mixed linear
sire models were fitted initially to data for each of the
wool traits and meat quality traits to identify those fixed
effects influencing each trait. The fixed effects included:
site (8 or 5 [meat color stability traits only] levels), year
of birth (5 levels), sex (2 levels, wool traits only), sheep
type (3 levels, ultra/super-fine, fine-fine/medium, and
medium/strong to account for sires being from different
types of Merino; Swan et al., 2016), type of birth and
rearing (6 levels: 11, 21, 22, 31, 32, or 33 for numbers of
lambs born and reared, respectively), and dam age (7 lev-
els: 2 to greater than or equal to 8 yr of age; see Mortimer
et al. 2017a for more details). Age of the lamb at mea-
surement was fitted as a linear covariate to the data for
meat quality and yearling wool traits only. For the meat
quality traits, slaughter group was also fitted, and testing
laboratory (2 levels) was fitted to the data for SF5 only.
No other covariates, such as carcass weight or pH, were
fitted to the meat quality data. Significant (P < 0.05) two-
way interactions were included in the final model.
Univariate mixed animal models, as described by
Swan et al. (2016), were then fitted to the data for each
trait to estimate variance components. Random effects
of animal and genetic group were included in all models,
with the genetic group effect defined by Merino flock of
origin (bloodline) or sheep type (Swan et al., 2016). The
genetic groups for each trait were derived from the ped-
igrees of animals having observations for that trait and
consisted of 113 to 134 genetic groups as appropriate to
each trait. Random effects of sire x site interaction and
dam (representing a maternal effect comprising both
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4263
maternal genetic and maternal environmental effects)
were then added to the model to assess the importance
of these effects in explaining variation in each trait. If a
significant increase in the log-likelihood value occurred
after inclusion of an additional random effect from that
of a reduced model, the random effect was retained.
However, there was limited ability to estimate the size
and importance of maternal effects for the meat quality
traits due to the structure of the data, where most dams
had only a single progeny recorded for these traits and a
maternal effect was significant only for pH24ST.
Variance ratios, including heritabilities, for each trait
were estimated as described by Mortimer et al. (2017a).
The phenotypic variance was the sum of the additive
genetic, maternal (pH24ST only), sire x site (SF5 only),
and the residual variances. As appropriate for pH24ST
and SF5, the ratios of maternal (0.18 ± 0.09) and sire x
site variances to phenotypic variance, respectively, were
estimated. The ratio of genetic group variance to addi-
tive genetic variance was also calculated as a compari-
son of the relative size of the between genetic group vari-
ance to the within genetic group variance. Phenotypic
and genetic covariances were estimated using a series
of bivariate analyses involving all combinations of traits
at each stage of measurement. Fixed effects and signifi-
cant 2-way interactions were fitted based on the univari-
ate analyses. The mixed bivariate models included all
significant fixed and random effects from the univariate
models for the meat quality and wool traits, where for
the latter traits the maternal effect and sire x site inter-
action were significant for most traits (Mortimer et al.,
2017a). However, in the instances where convergence
did not occur, the mixed models were reduced to exclude
nonestimable (co)variance components and achieve
convergence. The correlations involving EPA, DPA, and
EPA+DPA+DHA were not estimable because of the lack
of any genetic variance for these traits. As no males were
measured for both carcass and adult traits, the residual
covariances for these combinations of traits were fixed at
zero and phenotypic correlations were not estimated. For
all other trait combinations, phenotypic and genetic cor-
relations and their standard errors were estimated from
the appropriate covariances using ASReml.
RESULTS AND DISCUSSION
Heritability
The estimates of heritability and variances for the
meat quality traits are shown in Table 2, whereas those
for the wool traits were reported by Mortimer et al.
(2017a). There were high estimates of heritability for
IMF (0.58 ± 0.11) and retail display meat color light-
ness (retL*, 0.59 ± 0.15). There were several other traits
 4264 Mortimer et al.

Table 2. Estimates of phenotypic variance1, coefficient of variation (CV) and the proportion due to additive genetic
(h2), and genetic group variance ratio (b2)2 (± SE) for meat quality and nutritional value traits3
Trait Phenotypic variance CV (%) Heritability (h2) Genetic group (b2)
Meat quality
IMF (%) 0.827 19.6 0.58 ± 0.11 0.00 ± 0.00
SF5 (N) 69.15 26.8 0.10 ± 0.09 0.00 ± 0.00
pH24LL 0.011 1.8 0.15 ± 0.07 0.00 ± 0.00
pH24ST 0.032 3.1 0.20 ± 0.10 0.33 ± 0.54
ICDH (μmol/min/g tissue) 0.932 18.5 0.06 ± 0.10 2.08 ± 4.51
Myo (mg/g tissue) 1.77 17.9 0.31 ± 0.09 0.14 ± 0.25
cfa* 1.96 7.6 0.07 ± 0.07 0.00 ± 0.00
cfb* 1.11 28.9 0.08 ± 0.08 0.58 ± 1.10
cfL* 4.42 6.2 0.14 ± 0.08 0.21 ± 0.49
reta* 1.83 8.6 0.00 ± 0.00 n.e.4
retb* 1.84 8.0 0.13 ± 0.13 0.80 ± 1.26
retL* 5.18 6.4 0.59 ± 0.15 0.15 ± 0.19
retOxy/Met 0.243 15.5 0.06 ± 0.12 0.83 ± 2.55
reth 4.56 4.5 0.27 ± 0.15 0.53 ± 0.68
retC* 2.95 7.4 0.00 ± 0.00 n.e.
Nutritional value
IRON (mg/kg wet tissue) 9.72 14.1 0.20 ± 0.09 0.78 ± 0.73
ZINC (mg/kg wet tissue) 18.35 16.6 0.11 ± 0.09 1.12 ± 1.37
EPA (mg/100g tissue) 15.83 27.0 0.00 ± 0.00 n.e.
DPA (mg/100g tissue) 30.47 22.5 0.00 ± 0.00 n.e.
DHA (mg/100g tissue) 4.36 28.4 0.01 ± 0.07 7.15 ± 37.33
EPA+DHA (mg/100g tissue) 30.83 25.1 0.00 ± 0.00 n.e.
EPA+DPA+DHA (mg/100g tissue) 108.92 22.4 0.00 ± 0.00 n.e.
LA (mg/100g tissue) 942.50 19.0 0.14 ± 0.07 0.00 ± 0.00
ARA (mg/100g tissue) 108.91 18.9 0.00 ± 0.00 n.e.
LA+ARA (mg/100g tissue) 1419.1 17.4 0.10 ± 0.07 0.14 ± 0.50
1Phenotypic variance = sum of additive genetic variance and residual variance components, plus sire x site (SF5 only) and maternal (pH24ST only)
variance components.
2b2 = ratio of genetic group to additive genetic variance.
3IMF = intramuscular fat; SF5 = shear force after 5 days ageing; pH = pH at 24 h after slaughter for LL and ST muscles; ICDH = isocitrate dehydroge-
24
nase activity; Myo = myoglobin concentration; cfa* = fresh meat redness; cfb* = fresh meat yellowness; cfL* = fresh meat lightness; reta* = retail display
meat redness; retb* = retail display meat yellowness; retL* = retail display meat lightness; retOxy/Met = retail display meat oxy/met value; reth = retail dis-
play meat hue [h = tan-1(b*/a*)]; retC* = retail display meat chroma (C* = [a*2 + b*2]0.5); IRON = iron content; ZINC = zinc content; EPA = eicosapentae-
onic acid content; DPA = docosapentaeonic acid content; DHA = docosahexaeonic acid content; EPA+DHA = sum of EPA and DHA; EPA+DPA+DHA =
sum of EPA, DPA and DHA; LA = linoleic acid content; ARA = arachidonic acid content; LA+ARA = sum of LA and ARA.
4 n.e. = not estimable.

with moderate heritabilities of 0.2 or greater (Myo, reth,
pH24ST, IRON), although most traits had low or negli-
gible estimates of heritability. These Merino data are a
subset, representing 16 to 23% of the records depending
on the trait, from the earlier report by Mortimer et al.
(2014), which included the majority of carcasses sam-
pled from the IN program. The traits that had moderate
to high heritabilities in this Merino study were gener-
ally similar to those in the larger dataset which included
records on terminal sire and maternal breeds. However,
many more traits had moderate heritabilities in the latter
data set. For example, SF5 (0.27), reta* (0.25), retOxy/
Met (0.27), and ZINC (0.27) all had greater heritabili-
ties (Mortimer et al., 2014) than the low or negligible
values in the current Merino dataset. Similarly, greater
heritabilities for more of the meat quality traits were also
reported from data across two multibreed sheep popula-
tions (Bolormaa et al., 2016), as well as for shear force
of loins from terminal sire flocks (Brito et al., 2015).
Quality issues have been reported with Merino meat,
especially associated with high ultimate pH and suscep-
tibility to discoloration during retail display (Warner et
al., 2007; Hopkins and Mortimer, 2014). This is sup-
ported by the mean values found here (Table 1) for SF5
(31 N) which should be less than 27 N for acceptable
eating quality (Hopkins et al., 2006), high pH values in
the LL and ST which should be less than 5.7 (Warner et
al., 2010), and an retOxy/Met value (3.19) for which <
3.3 is considered discolored by consumers on average
(Khliji et al., 2010), whereas the mean values for these
traits were at acceptable levels for the larger meat sheep
data set (Mortimer et al., 2014). High pH and LA and

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 Correlations between wool and meat quality
the Merino breed have all been found to reduce meat
color stability (Jacob et al., 2014). The mean values
for iron and zinc contents observed in the present study
were slightly greater than those reported by Pannier et
al. (2014b) from a multibreed population. Nonetheless,
based on calculations using recommended dietary guide-
lines described by Pannier et al. (2010, 2014b), our mean
values fit with their conclusion that Australian lamb can
be claimed only as “good sources” of iron for women
over 50 yr old and men, and of zinc for women of all
ages. As levels of these minerals should be increased
if Australian lamb can be claimed as “good sources” of
iron for women under 50 yr old and of zinc for men, se-
lection offers one option of achieving this improvement.
The low heritability estimates for several meat qual-
ity traits (for example, SF5, reta*, retOxy/Met, ZINC,
and long-chain fatty acids) in the present study indi-
cate that selection to improve these traits in the Merino
would be relatively slow, although improvement could
be more rapid in a subset of traits such as IMF, meat pH,
meat color, and IRON. However, additional studies are
needed to obtain more accurate heritability estimates in
order to verify these expected responses for meat qual-
ity traits of Merino sheep. Such studies would require
pedigreed data based on greater numbers of records on
progeny than those available from our study, as well
as dams having records from several progeny. In their
review of the genetics of meat quality, Hopkins et al.
(2011) concluded that despite the relatively few esti-
mates there appeared to be moderate heritability for
traits such as shear force, pH, color, iron and zinc, and
long-chain fatty acids. In particular, our estimates of
heritability for pH and color are quite close to those
reported by Greeff et al. (2008) for Merino rams and
are generally within the range of estimates reviewed by
Hopkins and Mortimer (2014). Lorentzen and Vangen
(2012) reported greater heritabilities, with high SE, for
fresh meat color (L*, 0.53 ± 0.18; a*, 0.17 ± 0.14; b*,
0.29 ± 0.17) and muscle pH at 48h (LL, 0.20 ± 0.12;
M. semimembranosus, 0.30 ± 0.15) in lambs of a
Norwegian terminal sire breed. While there is consider-
able phenotypic variation for the health claimable ome-
ga-3 polyunsaturated fatty acids (EPA, DPA, DHA),
there is little or no genetic variation evident from these
Merino data and any improvement would need to be
through green pasture nutrition (Ponnampalam et al.,
2014). Nonetheless, further phenotyping of the long-
chain fatty acid content of lamb may be warranted in
order to confirm this conclusion, as earlier heritability
estimates were generally moderate from a Merino data
set of similar size (Greeff et al., 2007). There was a sig-
nificant sire x site variance ratio for SF5 (0.10 ± 0.05),
but there was no evidence of genotype x environment
interactions for any other traits.
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4265
Though standard errors were large, moderate to
large genetic group variances relative to the additive
genetic variances (b2) were estimated for several meat
color traits (cfb*, retb*, reth, and retOxy/Met), IRON,
ZINC, and ICDH (Table 2). From the few studies that
have compared meat quality traits of different types of
Merinos, the Merino strains have been reported to dif-
fer significantly in fresh meat b* (Fogarty et al., 2003;
Hopkins et al., 2007a), as well as L*, pH for the LL
(Fogarty et al., 2003; Hopkins et al., 2005a; Hopkins
et al., 2007a) and ST (Hopkins et al., 2005a; Hopkins
et al., 2007a), and ICDH activity, with higher levels in-
dicating greater aerobic capacity of muscle (Hopkins
et al., 2005a). An understanding of the size of genetic
group variation influencing the meat quality traits, as
well as for the range of wool traits (Swan et al., 2016;
Mortimer et al., 2017a), live weights (Swan et al., 2016;
Mortimer et al., 2017a), and carcass traits (Swan et al.,
2016; Mortimer et al., 2017b), will assist us to iden-
tify where modifications to genetic evaluation models
should be made to account for this source of variation in
data from breeds that consist of distinct subpopulations.
In addition, genetic group covariances also are needed,
though were not estimated during the present study due
to the relatively small numbers of available records.
More data on meat quality traits recorded on appropri-
ately structured and pedigreed Merino populations are
required to more accurately estimate genetic group (co)
variances. This is needed for the Merino breed as ap-
propriate modeling of genetic group effects in genetic
evaluation models has been shown to substantially re-
duce biases in estimated breeding values (Atkins et al.,
1999). In response, the process of revising the genetic
group covariances used in the MERINOSELECT ge-
netic evaluation model implemented by Sheep Genetics
has been initiated (Swan et al., 2016).
Genetic Correlations
Wool and Meat Quality Traits. All estimates of
genetic correlations between the yearling wool produc-
tion traits and meat quality traits were low or negligible,
with SE generally between 0.10 and 0.25 (Table 3).
Greeff et al. (2008) also reported low or negligible ge-
netic correlations between wool production traits and
meat pH and color in Merino rams. In an earlier study
of Merino rams, Fogarty et al. (2003) reported a signifi-
cant genetic correlation (-0.66 ± 0.27) between yCFW
and cfL*. The corresponding phenotypic correlations in
our data were generally smaller in magnitude than 0.1.
The corresponding genetic correlations for adult wool
production traits were very consistent with the yearling
traits and are available in Table S1 (see the article on-
line at http://journalofanimalscience.org).
 4266 Mortimer et al.

Table 3. Estimates of genetic and phenotypic correlations (± SE) between yearling wool production traits and
meat quality traits1
yGFW yYLD yCFW yFD yFDSD yFDCV ySS ySL yCUR
Genetic correlations
IMF -0.18 ± 0.11 0.03 ± 0.11 -0.17 ± 0.11 -0.03 ± 0.09 0.10 ± 0.11 0.12 ± 0.11 0.04 ± 0.12 -0.08 ± 0.11 0.09 ± 0.11
SF5 -0.04 ± 0.23 0.32 ± 0.24 0.11 ± 0.24 -0.01 ± 0.19 0.18 ± 0.24 0.23 ± 0.24 -0.25 ± 0.26 -0.22 ± 0.23 0.08 ± 0.22
pH24LL -0.05 ± 0.18 -0.09 ± 0.18 -0.08 ± 0.18 -0.26 ± 0.16 -0.02 ± 0.19 0.26 ± 0.18 -0.26 ± 0.20 0.11 ± 0.17 -0.17 ± 0.18
pH24ST -0.04 ± 0.16 -0.01 ± 0.15 -0.02 ± 0.16 -0.06 ± 0.13 0.10 ± 0.15 0.18 ± 0.14 -0.28 ± 0.16 0.10 ± 0.14 -0.02 ± 0.15
ICDH -0.23 ± 0.22 0.34 ± 0.24 -0.11 ± 0.22 -0.08 ± 0.20 -0.18 ± 0.23 -0.11 ± 0.23 0.24 ± 0.26 -0.38 ± 0.24 0.21 ± 0.22
Myo 0.03 ± 0.13 -0.12 ± 0.13 -0.01 ± 0.13 0.06 ± 0.11 -0.03 ± 0.13 -0.08 ± 0.13 -0.13 ± 0.14 0.03 ± 0.12 -0.02 ± 0.13
cfa* -0.07 ± 0.23 -0.17 ± 0.23 -0.14 ± 0.23 0.07 ± 0.20 0.19 ± 0.22 0.13 ± 0.22 -0.40 ± 0.27 -0.09 ± 0.21 0.36 ± 0.24
cfb* -0.02 ± 0.21 -0.11 ± 0.21 -0.10 ± 0.21 0.00 ± 0.18 0.08 ± 0.20 0.09 ± 0.20 -0.01 ± 0.22 -0.15 ± 0.20 0.17 ± 0.21
cfL* 0.09 ± 0.16 -0.12 ± 0.16 0.05 ± 0.17 0.14 ± 0.14 0.22 ± 0.16 0.14 ± 0.16 0.19 ± 0.18 -0.16 ± 0.16 0.12 ± 0.16
Phenotypic correlations
IMF 0.05 ± 0.03 -0.03 ± 0.04 0.02 ± 0.04 0.06 ± 0.04 0.02 ± 0.04 -0.02 ± 0.04 -0.01 ± 0.05 0.04 ± 0.05 0.00 ± 0.04
SF5 0.00 ± 0.04 -0.01 ± 0.05 0.00 ± 0.04 0.01 ± 0.04 0.05 ± 0.05 0.05 ± 0.05 -0.01 ± 0.05 -0.04 ± 0.05 0.01 ± 0.05
pH24LL -0.03 ± 0.04 -0.01 ± 0.05 -0.02 ± 0.04 -0.07 ± 0.04 -0.03 ± 0.05 0.00 ± 0.05 0.08 ± 0.05 0.05 ± 0.05 -0.07 ± 0.05
pH24ST -0.01 ± 0.04 0.03 ± 0.04 0.00 ± 0.04 -0.04 ± 0.04 -0.07 ± 0.04 -0.06 ± 0.04 0.00 ± 0.05 0.10 ± 0.05 -0.07 ± 0.04
ICDH 0.01 ± 0.04 0.03 ± 0.04 0.02 ± 0.04 -0.01 ± 0.04 0.03 ± 0.04 0.04 ± 0.05 0.01 ± 0.05 -0.04 ± 0.05 -0.02 ± 0.05
Myo 0.05 ± 0.03 0.02 ± 0.04 0.06 ± 0.04 0.09 ± 0.04 -0.01 ± 0.04 -0.07 ± 0.04 0.10 ± 0.05 0.01 ± 0.05 0.01 ± 0.04
cfa* 0.02 ± 0.04 -0.01 ± 0.04 0.02 ± 0.04 0.00 ± 0.04 -0.10 ± 0.04 -0.12 ± 0.04 0.04 ± 0.05 -0.01 ± 0.05 0.04 ± 0.05
cfb* 0.03 ± 0.04 0.04 ± 0.04 0.04 ± 0.04 0.04 ± 0.04 -0.04 ± 0.05 -0.09 ± 0.05 0.07 ± 0.05 -0.01 ± 0.05 0.04 ± 0.05
cfL* -0.04 ± 0.04 -0.01 ± 0.04 -0.05 ± 0.04 -0.01 ± 0.04 0.02 ± 0.05 0.04 ± 0.05 -0.04 ± 0.05 -0.12 ± 0.05 -0.01 ± 0.05
1IMF = intramuscular fat; SF5 = shear force after 5 days ageing; pH = pH at 24 h after slaughter for LL and ST muscles; ICDH = isocitrate dehydrogenase
24
activity; Myo = myoglobin concentration; cfa* = fresh meat redness; cfb* = fresh meat yellowness; cfL* = fresh meat lightness; yGFW = yearling greasy fleece
weight; yYLD = yearling clean yield; yCFW = yearling clean fleece weight; yFD = yearling fiber diameter; yFDSD = yearling fiber diameter SD; yFDCV =
yearling fiber diameter CV; ySS = yearling staple strength; ySL = yearling staple length; yCUR = yearling fiber curvature.

The estimates of genetic and phenotypic correla-
tions between the yearling wool production traits and
the meat nutritional value traits are shown in Table 4,
with the genetic correlations for adult wool produc-
tion traits available in Table S2 (see the article online
at http://journalofanimalscience.org). Most genetic cor-
relations were low or negligible, with some exceptions.
The long-chain omega-3 polyunsaturated fatty acids
(EPA + DHA) have favorable genetic correlations with
fleece weight and yield (moderate positive) and yFD
(moderate negative), although the correlations were
generally no larger than their very high SE. Similarly,
the long-chain omega-6 polyunsaturated fatty acids
(LA, ARA, LA + ARA) had low to moderate positive
genetic correlations with fleece weight and yield and
low negative correlations with yFD, ySL, and yCUR.
All the corresponding phenotypic correlations were
generally smaller in magnitude than 0.1. The genetic
correlations of EPA+DHA with adult fleece weight and
yield were approximately half the yearling values and
those for aSL with the long chain omega-6 polyunsatu-
rated fatty acids were reduced to negligible (Table S2).
The estimates of genetic and phenotypic correlations
between the yearling wool color and visual traits and the
meat quality traits are shown in Table 5, with genetic
correlations for adult wool color and visual traits avail-
able in Table S3. Most genetic correlations were neg-
ligible, with some exceptions. Meat shear force (SF5)
was lowly and positively genetically correlated with
both wool scoured brightness (yY) and yellowness (y(Y-
Z)), indicating that shear force values would be slightly
increased after selection for improved clean wool color.
However, these correlations were reduced to about half
for adult wool (Table S3). The genetic correlations for yY
with indicators of meat redness (Myo, cfa*) were low to
moderate and negative (unfavorable relationships), but
low positive with cfL* (favorable relationship). These
correlations were also reduced in magnitude with adult
wool, with a higher correlation between a(Y-Z) and cfL*
(0.32 ± 0.18; Table S3). While there were some low ge-
netic correlations between the meat quality traits and the
yearling visual traits, there were no consistent patterns.
The corresponding phenotypic correlations were gener-
ally smaller in magnitude than 0.1.
The estimates of genetic and phenotypic correlations
between the yearling wool color and visual traits and the
meat nutritional value traits are shown in Table 6, with
the genetic correlations for adult wool color and visual
traits available in Table S4. The genetic correlations be-
tween wool brightness (yY) and meat IRON and ZINC
were low and negative, although low and positive with
wool yellowness (y(Y-Z)), indicating that selecting for
improved clean wool color would be associated with
lower mineral content of lamb. The genetic correlation

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 Correlations between wool and meat quality 4267

Table 4. Estimates of genetic and phenotypic correlations (± SE) between yearling wool production traits and
meat nutritional value traits1
yGFW yYLD yCFW yFD yFDSD yFDCV ySS ySL yCUR
Genetic correlations
IRON -0.05 ± 0.14 -0.04 ± 0.13 -0.08 ± 0.14 -0.15 ± 0.11 -0.22 ± 0.13 -0.10 ± 0.13 -0.07 ± 0.14 -0.24 ± 0.13 0.12 ± 0.13
ZINC -0.06 ± 0.15 0.00 ± 0.15 -0.06 ± 0.15 -0.02 ± 0.13 -0.09 ± 0.15 -0.07 ± 0.15 0.14 ± 0.16 -0.09 ± 0.15 -0.13 ± 0.15
DHA -0.04 ± 0.19 0.21 ± 0.19 0.00 ± 0.19 -0.22 ± 0.17 -0.50 ± 0.22 -0.36 ± 0.22 -0.05 ± 0.20 0.04 ± 0.19 -0.23 ± 0.20
EPA+DHA 0.36 ± 0.54 0.52 ± 0.53 0.47 ± 0.57 -0.50 ± 0.40 n.e.2 n.e. 0.24 ± 0.51 0.23 ± 0.46 -0.74 ± 0.81
LA 0.26 ± 0.19 0.34 ± 0.20 0.34 ± 0.20 -0.17 ± 0.16 -0.16 ± 0.19 -0.05 ± 0.19 0.16 ± 0.21 -0.25 ± 0.19 -0.26 ± 0.19
ARA 0.41 ± 0.48 0.57 ± 0.50 0.55 ± 0.59 -0.12 ± 0.28 -0.14 ± 0.36 -0.09 ± 0.34 0.15 ± 0.38 -0.29 ± 0.34 -0.50 ± 0.44
LA+ARA 0.31 ± 0.22 0.42 ± 0.23 0.40 ± 0.24 -0.18 ± 0.18 -0.17 ± 0.21 -0.06 ± 0.21 0.17 ± 0.23 -0.28 ± 0.21 -0.32 ± 0.22
Phenotypic correlations
IRON 0.05 ± 0.03 -0.04 ± 0.04 0.05 ± 0.04 -0.01 ± 0.04 -0.03 ± 0.04 -0.02 ± 0.04 0.01 ± 0.05 -0.04 ± 0.05 -0.02 ± 0.04
ZINC 0.03 ± 0.04 -0.04 ± 0.04 0.02 ± 0.04 0.04 ± 0.04 0.02 ± 0.04 -0.00 ± 0.05 0.01 ± 0.05 0.01 ± 0.05 0.01 ± 0.04
DHA -0.04 ± 0.04 0.01 ± 0.04 -0.05 ± 0.04 0.05 ± 0.04 0.06 ± 0.04 0.03 ± 0.04 0.01 ± 0.05 0.02 ± 0.05 -0.08 ± 0.04
EPA+DHA -0.06 ± 0.03 0.02 ± 0.04 -0.06 ± 0.04 0.01 ± 0.04 n.e. n.e. 0.01 ± 0.04 -0.02 ± 0.04 -0.09 ± 0.04
LA 0.08 ± 0.04 -0.02 ± 0.04 0.06 ± 0.04 0.03 ± 0.04 0.06 ± 0.04 0.03 ± 0.04 0.04 ± 0.05 0.07 ± 0.05 -0.10 ± 0.04
ARA -0.01 ± 0.04 0.03 ± 0.04 0.01 ± 0.04 0.05 ± 0.04 0.11 ± 0.04 0.10 ± 0.05 0.03 ± 0.05 0.04 ± 0.05 -0.05 ± 0.05
LA+ARA 0.07 ± 0.04 -0.01 ± 0.04 0.05 ± 0.04 0.04 ± 0.04 0.07 ± 0.04 0.05 ± 0.04 0.04 ± 0.05 0.07 ± 0.05 -0.10 ± 0.04
1IRON = iron content; ZINC = zinc content; DHA = docosahexaeonic acid content; EPA = eicosapentaeonic acid content; EPA+DHA = sum of EPA
and DHA; LA = linoleic acid content; ARA = arachidonic acid content; LA+ARA = sum of LA and ARA; yGFW = yearling greasy fleece weight; yYLD =
yearling clean yield; yCFW = yearling clean fleece weight; yFD = yearling fiber diameter; yFDSD = yearling fiber diameter SD; yFDCV = yearling fiber
diameter CV; ySS = yearling staple strength; ySL = yearling staple length; yCUR = yearling fiber curvature.
2n.e. = not estimable.

Table 5. Estimates of genetic and phenotypic correlations (± SE) between yearling wool color and visual traits
and meat quality traits1
yY y(Y-Z) yBCOV mBRWR yBRWR yBDWR
Genetic correlations
IMF 0.12 ± 0.13 -0.14 ± 0.11 0.01 ± 0.15 -0.03 ± 0.12 0.04 ± 0.12 0.06 ± 0.12
SF5 0.26 ± 0.27 0.37 ± 0.25 -0.21 ± 0.30 0.15 ± 0.25 -0.20 ± 0.25 0.00 ± 0.24
pH24LL 0.01 ± 0.20 -0.15 ± 0.18 0.08 ± 0.24 -0.03 ± 0.20 -0.17 ± 0.20 -0.03 ± 0.19
pH24ST 0.03 ± 0.19 0.02 ± 0.15 -0.03 ± 0.22 -0.01 ± 0.18 0.00 ± 0.17 -0.05 ± 0.16
ICDH -0.15 ± 0.26 0.17 ± 0.22 0.37 ± 0.31 0.14 ± 0.25 0.35 ± 0.26 0.35 ± 0.25
Myo -0.26 ± 0.15 -0.03 ± 0.13 -0.10 ± 0.17 -0.14 ± 0.14 -0.05 ± 0.14 0.09 ± 0.14
cfa* -0.51 ± 0.29 0.13 ± 0.22 -0.20 ± 0.33 -0.24 ± 0.26 0.06 ± 0.25 -0.15 ± 0.24
cfb* -0.13 ± 0.24 0.05 ± 0.21 0.13 ± 0.28 0.05 ± 0.23 0.40 ± 0.24 0.15 ± 0.22
cfL* 0.32 ± 0.19 0.05 ± 0.16 0.36 ± 0.21 0.19 ± 0.17 0.32 ± 0.18 0.21 ± 0.18
Phenotypic correlations
IMF 0.04 ± 0.04 0.00 ± 0.04 -0.05 ± 0.04 -0.08 ± 0.03 -0.07 ± 0.04 -0.06 ± 0.04
SF5 0.01 ± 0.05 0.02 ± 0.05 0.11 ± 0.05 0.06 ± 0.04 0.06 ± 0.05 0.05 ± 0.05
pH24LL -0.10 ± 0.05 0.01 ± 0.05 0.09 ± 0.05 0.04 ± 0.04 -0.02 ± 0.05 -0.06 ± 0.05
pH24ST 0.01 ± 0.05 -0.05 ± 0.04 0.08 ± 0.04 0.03 ± 0.04 -0.02 ± 0.05 -0.05 ± 0.05
ICDH 0.04 ± 0.05 0.01 ± 0.04 -0.04 ± 0.04 0.03 ± 0.04 0.01 ± 0.05 0.11 ± 0.05
Myo 0.00 ± 0.04 0.01 ± 0.04 -0.08 ± 0.04 -0.07 ± 0.03 -0.09 ± 0.04 -0.02 ± 0.05
cfa* 0.14 ± 0.05 -0.02 ± 0.04 -0.10 ± 0.04 -0.02 ± 0.04 -0.10 ± 0.05 -0.14 ± 0.05
cfb* 0.07 ± 0.05 0.02 ± 0.05 0.00 ± 0.05 -0.02 ± 0.04 -0.04 ± 0.05 -0.07 ± 0.05
cfL* 0.01 ± 0.05 0.00 ± 0.05 -0.01 ± 0.04 -0.01 ± 0.04 0.00 ± 0.05 0.00 ± 0.05
1IMF = intramuscular fat; SF5 = shear force after 5 days ageing; pH = pH at 24 h after slaughter for LL and ST muscles; ICDH = isocitrate dehydrogenase
24
activity; Myo = myoglobin concentration; cfa* = fresh meat redness; cfb* = fresh meat yellowness; cfL* = fresh meat lightness; yY = yearling brightness, y(Y-Z) =
yearling yellowness; yBCOV = yearling breech cover; mBRWR = marking breech wrinkle; yBRWR = yearling breech wrinkle; yBDWR = yearling body wrinkle.

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Table 6. Estimates of genetic and phenotypic correlations (± SE) between yearling wool color and visual traits
and meat nutritional value traits1
yY y(Y-Z) yBCOV mBRWR yBRWR yBDWR
Genetic correlations
IRON -0.30 ± 0.15 0.14 ± 0.13 -0.04 ± 0.18 0.14 ± 0.15 0.16 ± 0.15 0.25 ± 0.14
ZINC -0.24 ± 0.17 0.26 ± 0.14 -0.10 ± 0.21 -0.04 ± 0.16 -0.01 ± 0.17 -0.06 ± 0.16
DHA 0.26 ± 0.21 -0.12 ± 0.19 0.05 ± 0.25 -0.10 ± 0.21 -0.10 ± 0.21 -0.18 ± 0.21
EPA+DHA 0.17 ± 0.41 -0.17 ± 0.38 0.11 ± 0.50 0.15 ± 0.43 0.01 ± 0.41 -0.19 ± 0.44
LA 0.28 ± 0.22 -0.02 ± 0.18 0.22 ± 0.26 0.43 ± 0.19 0.47 ± 0.21 0.42 ± 0.20
ARA -0.43 ± 0.39 0.52 ± 0.44 -0.13 ± 0.42 0.21 ± 0.36 0.64 ± 0.54 0.50 ± 0.50
LA+ARA 0.17 ± 0.25 0.07 ± 0.20 0.18 ± 0.29 0.42 ± 0.22 0.53 ± 0.25 0.47 ± 0.24
Phenotypic correlations
IRON -0.01 ± 0.04 0.01 ± 0.04 -0.03 ± 0.04 -0.04 ± 0.04 -0.04 ± 0.04 0.07 ± 0.04
ZINC -0.09 ± 0.05 0.10 ± 0.04 -0.03 ± 0.04 -0.04 ± 0.04 -0.05 ± 0.05 -0.03 ± 0.05
DHA -0.06 ± 0.04 0.08 ± 0.04 0.00 ± 0.04 -0.02 ± 0.04 -0.08 ± 0.04 -0.04 ± 0.04
EPA+DHA -0.05 ± 0.04 0.04 ± 0.04 0.01 ± 0.04 0.00 ± 0.04 -0.03 ± 0.04 -0.01 ± 0.04
LA 0.00 ± 0.05 0.06 ± 0.04 -0.04 ± 0.04 -0.01 ± 0.03 0.03 ± 0.05 0.08 ± 0.05
ARA -0.04 ± 0.05 0.08 ± 0.04 -0.01 ± 0.04 0.01 ± 0.04 0.08 ± 0.05 0.14 ± 0.05
LA+ARA -0.01 ± 0.05 0.07 ± 0.04 -0.04 ± 0.04 -0.01 ± 0.04 0.05 ± 0.05 0.10 ± 0.05
1IRON = iron content; ZINC = zinc content; DHA = docosahexaeonic acid content; EPA = eicosapentaeonic acid content; EPA+DHA = sum of EPA
and DHA; LA = linoleic acid content; ARA = arachidonic acid content; LA+ARA = sum of LA and ARA; yY = yearling brightness, y(Y-Z) = yearling
yellowness; yBCOV = yearling breech cover; mBRWR = marking breech wrinkle; yBRWR = yearling breech wrinkle; yBDWR = yearling body wrinkle.

for ARA with yY was moderate negative and moderate
positive with y(Y-Z), although with high SE. There were
low to moderate positive genetic correlations between all
the wrinkle scores and the long-chain omega-6 polyun-
saturated fatty acids (LA, ARA, LA + ARA). The corre-
sponding phenotypic correlations were generally smaller
in magnitude than 0.1. The genetic correlations with the
adult wool and visual traits (Table S4) were consistent
with the yearling traits, except that the correlations be-
tween aBCOV and the long-chain omega-6 polyunsatu-
rated fatty acids were all increased (0.33 to 0.76).
The low to negligible genetic correlations between
the wool production and meat quality traits indicate that
wool breeding programs will have little or no effect on
meat quality. Additionally, it is likely that the appropri-
ate definition of dual-purpose breeding objectives and
design of selection indexes should allow improvement
of both wool production and meat quality from breed-
ing programs even where the genetic relationships are
unfavorable. As noted earlier with respect to the herita-
bility estimates, validation of these expected outcomes
will require more accurate estimates of the genetic cor-
relations between the wool production and meat qual-
ity traits derived from well-designed studies (including
more accurate estimates among the meat quality traits
as will become evident from estimates presented later
in this study). The low negative genetic correlation
of FD with pH24LL reported here supports an earlier
suggestion by Greeff et al. (2008) that Merino breed-
ing programs should also aim to produce meat that is
less prone to having high pH, avoiding meat that has
a reduced shelf life and is dark cutting (Hopkins and
Mortimer, 2014). Similarly, the low negative genetic
correlation of fleece weight with IMF suggests that
Merino breeding programs, particularly those with an
emphasis on fleece weight, should also consider IMF.
While the mean value of IMF in the present study
was within the range considered acceptable for lamb
by Australian consumers (4-5%; Warner et al., 2010;
Pannier et al., 2014c), wool and dual purpose breeding
programs should ensure that this level is maintained.
Correlations among Meat Quality Traits.
Estimates of genetic and phenotypic correlations
among the meat quality traits are shown in Table 7.
There was a high negative genetic correlation between
IMF and SF5 (-0.76 ± 0.24), which is consistent with
estimates reported by Mortimer et al. (2014, 2015)
from the larger meat sheep data set sampled for this
study (-0.62 ± 0.07, -0.64 ± 0.06, respectively), and
Karamichou et al. (2006) in Scottish Blackface sheep
(-0.54 ± 0.24). In contrast, Brito et al. (2015) have re-
ported a weak genetic correlation between marbling
score and shear force. These results strongly support
the conclusion of Hopkins and Mortimer (2014) that
selection for increased IMF will have a favorable ef-
fect on shear force. IMF was also positively geneti-
cally correlated with meat color for relative yellow-
ness (cfb* 0.67 ± 0.23), relative lightness (cfL* 0.50 ±
0.23), and relative redness (cfa* 0.17 ± 0.29), but
Lorentzen and Vangen (2012) reported low negative
correlations, albeit with high SE. There were also
highly negative genetic correlations for cfb* (-0.97 ±
0.54) and cfL* (-0.70 ± 0.41) with SF5. These genetic
correlations of the meat color traits with both SF5 and

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 Correlations between wool and meat quality 4269

Table 7. Estimates of genetic (below diagonal) and phenotypic (above diagonal) correlations (± SE) among meat
quality traits1
IMF SF5 pH24LL pH24ST ICDH Myo cfa* cfb* cfL*
IMF - -0.30 ± 0.03 -0.09 ± 0.03 -0.07 ± 0.03 0.04 ± 0.03 0.07 ± 0.03 0.12 ± 0.03 0.23 ± 0.03 0.26 ± 0.03
SF5 -0.76 ± 0.24 - 0.07 ± 0.03 0.13 ± 0.03 0.01 ± 0.04 -0.12 ± 0.03 -0.14 ± 0.03 -0.17 ± 0.03 -0.15 ± 0.03
pH24LL -0.01 ± 0.23 -0.45 ± 0.51 - 0.36 ± 0.03 -0.07 ± 0.04 -0.02 ± 0.03 -0.21 ± 0.03 -0.18 ± 0.03 -0.02 ± 0.03
pH24ST -0.18 ± 0.19 0.38 ± 0.35 0.68 ± 0.21 - -0.12 ± 0.04 -0.01 ± 0.03 -0.12 ± 0.03 -0.10 ± 0.03 -0.07 ± 0.03
ICDH 0.20 ± 0.31 0.37 ± 0.58 -0.53 ± 0.52 0.57 ± 0.45 - 0.17 ± 0.04 0.05 ± 0.04 n.e. -0.16 ± 0.04
Myo 0.09 ± 0.17 -0.16 ± 0.32 -0.19 ± 0.27 -0.10 ± 0.22 0.42 ± 0.32 - 0.07 ± 0.03 0.00 ± 0.03 -0.18 ± 0.03
cfa* 0.17 ± 0.29 0.02 ± 0.58 -0.25 ± 0.45 -0.19 ± 0.37 -0.10 ± 0.63 0.76 ± 0.43 - 0.58 ± 0.02 -0.06 ± 0.03
cfb* 0.67 ± 0.23 -0.97 ± 0.54 -0.63 ± 0.35 -0.59 ± 0.31 n.e.2 0.60 ± 0.29 0.74 ± 0.29 - 0.33 ± 0.03
cfL* 0.50 ± 0.18 -0.70 ± 0.41 -0.23 ± 0.33 -0.37 ± 0.28 0.03 ± 0.47 -0.71 ± 0.22 0.14 ± 0.41 0.72 ± 0.27 -
1IMF = intramuscular fat; SF5 = shear force after 5 days ageing; pH24 = pH at 24 h after slaughter for LL and ST muscles; ICDH = isocitrate dehydro-
genase activity; Myo = myoglobin concentration; cfa* = fresh meat redness; cfb* = fresh meat yellowness; cfL* = fresh meat lightness.
2n.e. = not estimable.

IMF were consistent with Mortimer et al. (2014) and
the high negative correlation between IMF and SF5.
All the meat color measures were negatively ge-
netically correlated (favorable) with meat pH (-0.19
to -0.63), which was generally consistent with other
reports (Fogarty et al., 2003; Safari et al., 2005; Greeff
et al., 2008; Lorentzen and Vangen, 2012; Mortimer
et al., 2014; Brito et al., 2015). There were high posi-
tive genetic correlations for meat relative redness (cfa*
0.76 ± 0.43) and relative yellowness (cfb* 0.60 ± 0.29)
with Myo, while that for meat relative brightness (cfL*
-0.71 ± 0.22) was highly negative, which would be ex-
pected and is consistent with Mortimer et al. (2014) for
cfa* and cfL*, but not cfb*, which they reported as close
to zero. There were high positive genetic correlations for
cfb* with cfa* (0.74 ± 0.29) and cfL* (0.72 ± 0.27; and
0.58 ± 0.02 and 0.33 ± 0.03, respectively, for the phe-
notypic correlations), which were generally consistent
with other studies (Greeff et al., 2008; Lorentzen and
Vangen, 2012; Mortimer et al., 2014). There were also
moderate phenotypic correlations for cfb* with cfa* and
cfL*, but negligible between cfa* and cfL*. Despite a
high positive genetic correlation for pH in the LL and
ST muscles (0.68 ± 0.21; and a moderate phenotypic
correlation of 0.36 ± 0.03), the genetic correlations for
SF5 and ICDH were negative with pH24LL and positive
with pH24ST, albeit with high SE. Lorentzen and Vangen
(2012) also reported a high genetic correlation (0.64 ±
0.32; and moderate phenotypic correlation 0.39) for pH
at 48h in LL and M. semimembranosus, while Brito et al.
(2015) estimated a moderate positive genetic correlation
of pH at 24 h with shear force. The genetic correlation
estimates suggest that pH measures in these 2 muscles
are genetically different traits. Based on simple correla-
tion coefficients, McPhail et al. (2014) had concluded
that ultimate pH in the LL muscle was not a reliable pre-
dictor of ultimate pH in other muscles of lamb carcasses.
The ST muscle in lamb has been found to have more fast
glycolytic fibers than the LL muscle (Greenwood et al.,
2007), and ultimate pH levels increase as the content of
these fibers increases (Gardner et al., 2006).
There were moderate to high genetic correlations
for IRON with both the long-chain omega-3 polyunsat-
urated fatty acids (DHA, 0.50 ± 0.28 and EPA+DHA,
0.67 ± 0.39) and long-chain omega-6 polyunsaturated
fatty acids (LA, 0.38 ± 0.28, ARA, 0.79 ± 0.63 and
LA + ARA, 0.49 ± 0.31), but the corresponding cor-
relations for ZINC were negligible (Table 8). These

Table 8. Estimates of genetic (below diagonal) and phenotypic (above diagonal) correlations (± SE) among meat
nutritional value traits1
IRON ZINC DHA EPA+DHA LA ARA LA+ARA
IRON - 0.26 ± 0.03 0.10 ± 0.03 0.10 ± 0.03 0.14 ± 0.03 0.15 ± 0.03 0.16 ± 0.03
ZINC 0.10 ± 0.23 - 0.04 ± 0.03 -0.01 ± 0.03 0.08 ± 0.03 0.11 ± 0.03 0.09 ± 0.03
DHA 0.50 ± 0.28 0.23 ± 0.33 - 0.82 ± 0.01 0.26 ± 0.03 0.40 ± 0.02 0.33 ± 0.03
EPA+DHA 0.67 ± 0.39 -0.14 ± 0.68 0.80 ± 0.45 - 0.28 ± 0.03 0.43 ± 0.02 0.35 ± 0.03
LA 0.38 ± 0.28 -0.06 ± 0.33 0.28 ± 0.37 0.11 ± 0.76 - 0.58 ± 0.02 n.e.
ARA 0.79 ± 0.63 -0.13 ± 0.57 0.86 ± 0.54 0.23 ± 1.20 0.50 ± 0.47 - 0.75 ± 0.01
LA+ARA 0.49 ± 0.31 -0.08 ± 0.37 0.42 ± 0.38 0.14 ± 0.83 n.e.2 0.61 ± 0.39 -
1IRON = iron content; ZINC = zinc content; DHA = docosahexaeonic acid content; EPA = eicosapentaeonic acid content; EPA+DHA = sum of EPA and
DHA; LA = linoleic acid content; ARA = arachidonic acid content; LA+ARA = sum of LA and ARA.
2n.e. = not estimable.

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 4270 Mortimer et al.

Table 9. Estimates of genetic and phenotypic correlations (± SE) between meat quality and meat nutritional value traits1
IRON ZINC DHA EPA+DHA LA ARA LA+ARA
Genetic correlations
IMF 0.16 ± 0.17 0.41 ± 0.19 0.09 ± 0.24 -0.29 ± 0.55 0.74 ± 0.16 0.11 ± 0.42 0.69 ± 0.20
SF5 -0.09 ± 0.34 -0.25 ± 0.41 -0.20 ± 0.51 -0.31 ± 1.14 -0.52 ± 0.41 0.28 ± 0.84 -0.42 ± 0.47
pH24LL -0.21 ± 0.27 0.14 ± 0.31 0.17 ± 0.38 0.33 ± 0.80 -0.35 ± 0.37 -0.10 ± 0.65 -0.33 ± 0.41
pH24ST -0.13 ± 0.23 0.05 ± 0.27 -0.03 ± 0.32 0.25 ± 0.70 -0.53 ± 0.32 -0.71 ± 0.66 -0.61 ± 0.36
ICDH 0.49 ± 0.34 0.09 ± 0.42 0.52 ± 0.47 n.e.2 0.81 ± 0.44 n.e. 0.94 ± 0.48
Myo 0.89 ± 0.10 0.47 ± 0.21 0.60 ± 0.29 n.e. 0.10 ± 0.27 0.43 ± 0.51 0.17 ± 0.30
cfa* 0.77 ± 0.35 0.15 ± 0.40 0.45 ± 0.50 0.54 ± 1.13 0.77 ± 0.49 0.92 ± 0.96 0.85 ± 0.55
cfb* 0.56 ± 0.31 -0.24 ± 0.35 0.25 ± 0.42 0.20 ± 0.90 n.e. n.e. n.e.
cfL* -0.33 ± 0.23 0.07 ± 0.29 0.01 ± 0.35 -0.44 ± 0.78 n.e. n.e. n.e.
Phenotypic correlations
IMF 0.04 ± 0.03 0.03 ± 0.03 0.02 ± 0.03 -0.06 ± 0.03 0.37 ± 0.03 -0.05 ± 0.03 0.29 ± 0.03
SF5 -0.05 ± 0.03 0.00 ± 0.03 0.03 ± 0.03 0.07 ± 0.03 -0.16 ± 0.03 0.03 ± 0.03 -0.12 ± 0.03
pH24LL -0.09 ± 0.03 0.03 ± 0.03 -0.03 ± 0.03 -0.02 ± 0.03 -0.10 ± .03 -0.04 ± 0.03 -0.10 ± 0.03
pH24ST -0.08 ± 0.03 0.05 ± 0.03 -0.01 ± 0.03 0.01 ± 0.03 -0.03 ± 0.03 0.00 ± 0.03 -0.02 ± 0.03
ICDH 0.25 ± 0.03 0.11 ± 0.04 0.15 ± 0.04 n.e. 0.22 ± 0.04 n.e. 0.24 ± 0.04
Myo 0.40 ± 0.02 0.16 ± 0.03 0.05 ± 0.03 n.e. 0.07 ± 0.03 0.10 ± 0.03 0.08 ± 0.03
cfa* 0.12 ± 0.03 0.04 ± 0.03 0.00 ± 0.03 -0.02 ± 0.03 0.06 ± 0.03 -0.01 ± 0.03 0.05 ± 0.03
cfb* -0.03 ± 0.03 0.04 ± 0.03 0.00 ± 0.03 -0.03 ± 0.03 n.e n.e. n.e.
cfL* -0.19 ± 0.03 0.00 ± 0.03 -0.04 ± 0.03 -0.08 ± 0.03 n.e. n.e. n.e.
1IMF = intramuscular fat; SF5 = shear force after 5 days ageing; pH = pH at 24 h after slaughter for LL and ST muscles; ICDH = isocitrate dehydroge-
24
nase activity; Myo = myoglobin concentration; cfa* = fresh meat redness; cfb* = fresh meat yellowness; cfL* = fresh meat lightness; IRON = iron content;
ZINC = zinc content; DHA = docosahexaeonic acid content; EPA = eicosapentaeonic acid content; EPA+DHA = sum of EPA and DHA; LA = linoleic acid
content; ARA = arachidonic acid content; LA+ARA = sum of LA and ARA.
2 n.e. = not estimable.

correlations with IRON in this Merino data set are
somewhat greater than those reported by Mortimer et
al. (2014) in the larger meat sheep data set.
There was a moderate positive genetic correlation
between IMF and ZINC (Table 9; 0.41 ± 0.19), al-
though Mortimer et al. (2014) found a negligible rela-
tionship (0.09 ± 0.09) in the larger meat sheep data set.
There were moderate to high genetic correlations for
IRON with ICDH, Myo, and meat color and high cor-
relations for IMF with LA and LA+ARA, which were
all generally consistent with Mortimer et al. (2014).
While there were some other high genetic correlations
in Table 9, they generally had very high SE.
Conclusions
This study provides the first report of genetic cor-
relations between a comprehensive range of wool
production and quality traits and meat quality and
nutritive value traits in sheep. In addition, the study
provides the first report of the genetic correlations
among the meat quality traits in Merino sheep, which
is an important breed contributing to sheep meat pro-
duction, but which may have some issues with meat
quality. There is significant genetic variation for some
meat quality traits, and therefore scope for improve-
ment in the Merino, especially for traits such as IMF,
retail meat color, pH, Myo, and IRON. The low to
negligible genetic correlations between the wool
production and meat quality traits indicate that wool
breeding programs will have little or no effect on meat
quality. There were moderately favorable genetic cor-
relations between important yearling wool production
traits and the long-chain omega-3 polyunsaturated fat-
ty acids, but they are unlikely to be important in wool/
meat breeding programs because the correlations have
high SE, were reduced for adult wool production, and
the omega-3 traits have little or no genetic variance.
The genetic correlations among the meat quality traits
in this Merino data set support the earlier report of
the larger meat sheep data set (Mortimer et al., 2014)
and point to ways of improving some of the issues
with Merino meat quality. In particular, selection to
increase IMF, which can be achieved relatively rap-
idly, will improve meat tenderness, meat color, and in-
crease meat ZINC levels. As well as EBV for IMF and
SF5 that are now available for meat-producing sheep
in Australia (Sheep Genetics, 2016), genetic evalua-
tion and reporting of EBV for pH, retail meat color,
and meat iron content should be considered.
Previous studies in this series (Mortimer et al.,
2017a; Mortimer et al., 2017b) and the present study
provide a unique set of estimates of genetic correlations
for a comprehensive range of wool and meat production

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 Correlations between wool and meat quality
and quality traits that provide the basis for development
of effective breeding programs that combine wool and
meat production and quality objectives. Our results show
that many of the meat production and quality traits have
genetic variation and would be expected to respond to
selection. In addition, there is no evidence of important
genotype x environment interactions for the meat traits
so that high genetic merit animals can be sourced from
a wide range of environments. The genetic correlations
between the major wool and meat traits are generally low
to moderate, if not negligible, so there are no major an-
tagonisms in combining wool and meat traits in breeding
programs. However, the large standard errors associated
with some genetic parameter estimates, particularly the
genetic correlations, do mean that further studies should
be conducted to obtain more accurate estimates and
verify the findings from the present series of studies.
Nonetheless, selection to increase weight (including car-
cass weight and dressing percentage) will result in moder-
ate improvement in wool weight, less wrinkled and barer
breech sheep, with some unfavorable increase in FD and
deterioration in wool color. While meat breeding pro-
grams generally emphasize reducing fat, as the Merino
has low carcass fat levels it may be desirable to increase
fat for better reproduction, welfare, and meat quality out-
comes. Carcass fat and lean are highly positively corre-
lated and will respond to selection, with a slight increase
in FD but with little effect on other wool traits. Selection
for increased lean meat yield will reduce carcass fat and
dressing percentage and will result in a moderate increase
in carcass muscle with a slight increase in wool weight.
All of these correlated changes will be small and any that
are unfavorable can easily be addressed by using an ap-
propriate selection index to combine the wool and meat
traits. While these estimated parameters provide a basis
for calculation of more accurate estimated breeding val-
ues, such as Australian Sheep Breeding Values (Brown et
al., 2007), knowledge of the relative size between genetic
group genetic variation influencing wool and meat pro-
duction traits of Merino sheep is also needed. This will
improve genetic evaluation models and allow the possi-
bility of predictions of genetic gains from index selection
based on genetic variation within and between genetic
groups (Swan et al., 2016).
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