Small Ruminant Research 101 (2011) 160172

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Small Ruminant Research

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Review

Genetic related effects on sheep meat quality

D.L. Hopkins a, , N.M. Fogarty b , S.I. Mortimer c
a
b
c

NSW Department of Primary Industries, Centre for Red Meat and Sheep Development, P.O. Box 129, Cowra, NSW 2794, Australia
NSW Department of Primary Industries, Orange Agricultural Institute, Forest Road, Orange, NSW 2800, Australia
NSW Department of Primary Industries, Agricultural Research Centre, Trangie, NSW 2823, Australia

a r t i c l e

i n f o

Article history:
Available online 29 September 2011
Keywords:
Meat quality
Sheep
Eating quality
Nutritive value

a b s t r a c t
This review examines the inuence of sheep genotype on meat quality traits where genotype includes breeds, cross-breeds and major gene effects. Differences in experimental
design have constrained some of the robustness of the conclusions which can be drawn
from the studies reviewed, but there is good evidence that, Merinos do have a propensity
to produce meat with a higher pH and in some cases specic muscles have reduced colour
stability. On the other hand, there is some evidence that increasing levels of Merino genetics improves the level of unsaturated fatty acids. There are clear effects of major genes on
traits such as tenderness and intramuscular fat and their mechanisms of action and effects
on eating quality are worthy of more attention. There is also evidence that sires selected
for extremes in breeding values for muscling or fatness will produce progeny with lower
eating quality. Recent large studies have indicated that many of the meat quality traits
are moderately heritable and, despite some potentially less than desirable genetic correlations between traits, there is scope to incorporate new and novel traits such as shear force
(indicator of tenderness), intramuscular fat and omega-3 fatty acids in breeding programs.
Crown Copyright 2011 Published by Elsevier B.V. All rights reserved.

1. Introduction
Meat quality includes many factors including palatability, water-holding capacity, colour, nutritional value and
safety (Hopkins and Geesink, 2009) and it can be affected
by changing the genetics and the production and processing environment. The relative importance of meat quality
traits varies according to the user of the product and the
type of product. For example, tenderness is more important for beef (Thompson, 2002) than sheep meat (Hopkins
et al., 2005c). As improvements are made in individual
traits their relative importance changes (Thompson, 2004),
which impacts on their emphasis in breeding programs.

This paper is part of the special issue entitled Products from Small
Ruminants, Guest Edited by A. Govaris and G. Moatsou.
Corresponding author.
E-mail address: david.hopkins@industry.nsw.gov.au (D.L. Hopkins).

Genetic change can occur through cross-breeding,

direct selection for quantitative traits or using genes with
major effects. The void of information on genetic variation for meat quality traits in sheep was highlighted
by Safari et al. (2005) who reported only 2 estimates
of heritability for pH and meat colour, both for Merino
rams. In recent years, more information has become available for both genetic variation and major gene effects on
meat quality traits and for the use of molecular markers.
Marker-assisted selection has the potential to signicantly
increase the rate of gain from selection for meat traits
(Meuwissen and Goddard, 1996). In Australia, large data
sets have been generated from the Cooperative Research
Centre for Sheep Industry Innovation (Sheep CRC) Information Nucleus (Fogarty et al., 2007) and the Sheep Genomics
(Oddy et al., 2007) programs. They are providing estimates
of genetic parameters for a large range of traits including
meat quality as well as developing molecular markers and
evaluating whole genome selection using single nucleotide

0921-4488/$ see front matter. Crown Copyright 2011 Published by Elsevier B.V. All rights reserved.
doi:10.1016/j.smallrumres.2011.09.036

D.L. Hopkins et al. / Small Ruminant Research 101 (2011) 160172

polymorphism technology. Development of strategies to

combine the quantitative and molecular information into
effective breeding programs is required (Fogarty, 2009).
This paper reviews studies that have compared the meat
quality traits between different sheep genotypes (breeds,
cross-breeds and major gene effects), estimates of genetic
parameters and approaches for using this information in
industry breeding programs. This review focuses on ndings since those of Bishop and Karamichou (2009) and
Simm et al. (2009), who identied the challenges for inclusion of meat quality traits in sheep breeding programs, and
covers a more extensive range of traits than Warner et al.
(2010).
2. Effect of genotypes on meat quality
2.1. Impact on tenderness
Tenderness can be evaluated by objectively measuring
shear force (Hopkins et al., 2010) and using trained panelists (Safari et al., 2001) or consumers (Hopkins et al.,
2005c). The differences between genotypes may vary with
the method used, as each measures subtle differences in
tenderness. Some studies have shown either no differences
in objectively measured tenderness between breeds and
crossbreds (Dranseld et al., 1979; Hopkins and Fogarty,
1998; Hopkins et al., 2005c) or inconsistent differences
that were not explained by variation in other traits that
inuence tenderness, such as pH, sarcomere length, carcass
weight or fat levels (Purchas et al., 2002). Different strategies have been used to minimize the impact of processing
on tenderness, including conditioning after slaughter and
ageing (Dranseld et al., 1979), electrical stimulation and
ageing (Hopkins et al., 2005c) and ageing for 7 days
(Hopkins and Fogarty, 1998). Such approaches are needed
to estimate genetic variation because of the potential confounding due to processing factors.
No sire breed effects on taste panel assessed tenderness
were reported by Dranseld et al. (1979) or Safari et al.
(2001) in comparisons of Merino lambs and other breeds
including Texel Merino or Poll Dorset Merino (PDM).
Hopkins et al. (2005c) reported minimal differences in
consumer assessed tenderness between genotypes, except
that the Merinos had lower sensory scores than Border
Leicester Merino (BLM) lambs for two different muscles,
which may have reected a slower rate of pH decline in
the Merino lambs. Rambouillet lambs produced tougher
leg steaks than Karakul and crossbred (Suffolk or Hampshire Rambouillet) lambs (Edwards et al., 1982), although
the reason cannot be conrmed as other traits, such as pH,
were not reported. This was also the case in a comprehensive study of lighter weight lambs, in which Merino lambs
were rated more tender by trained panelists than Rasa
Aragonesa and Churra breeds as slaughter weight increased
(Martnez-Cerezo et al., 2005). Merino lambs had the tenderest m. longissimus in the work of Young et al. (1993),
which was attributed to signicantly higher pH, although
this was not found by Hopkins and Fogarty (1998).
There were no effects of sire breed on sensory tenderness of lamb from 3 sire breeds (Charollais, Suffolk
and Texel) over 3 years (Ellis et al., 1997). Similarly

161

Esenbuga et al. (2001) found no difference in shear force

or sensory assessed tenderness between 4 fat-tailed types
(Awassi, Red Karaman, Tushin and Awassi Tushin) when
slaughtered at similar weights. Likewise Hoffman et al.
(2003) only reported tougher meat (shear force of the m.
semimembranosus; SM) for one of the 6 genotypes they
studied, with the effect associated with the dam breed
(Dohne Merino), although it did not affect sensory traits.
In hill breeds, Carson et al. (2001) reported no difference
in shear force of loin meat from 6 genotypes, although the
low absolute shear force values indicate the meat was aged
for an extended period, which may have reduced any differences between the genotypes.
Inconsistent effects were reported for taste panel tenderness of roasted hind leg lamb meat from 3 Greek dairy
breeds (Arsenos et al., 2002), although there was confounding of slaughter days and few animals evaluated. In
a larger study, Navajas et al. (2008) reported a reduction
in taste panel tenderness for both the loin and the SM
from pure Texel compared to Scottish Blackface lambs. The
authors suggested this reected lower intramuscular fat
(IMF) levels in the Texel, although it was not analysed and
surprisingly there was no difference between breeds for
muscularity.
In a large study across 6 countries, lamb meat from the
Icelandic breed was the tenderest, whether determined
by objective or subjective means, whereas the Bergam
asca breed was the toughest (Berge et al., 2003; Sanudo
et al., 2003). However, the data suggested that some of the
effect was due to differences in sarcomere length (Berge
et al., 2003), nal pH, lambs raised under different production systems and slaughtered in different countries over
a wide range in carcass weights (5.430.5 kg). In another
study, genotype was confounded with feeding system and
age (Fisher et al., 2000). These reports are not informative
for understanding any genetic differences in tenderness
between genotypes and overall no large genotype effects
on tenderness are apparent.
2.2. Impact on eating quality
Young et al. (1993) reported no differences in the juiciness, avour and overall acceptability of loin meat from 6
genotypes when tested by trained panelists. A comparison
of roasted legs from Romney, Border Leicester Romney,
Perendale, Corriedale and Merino animals by Kirton et al.
(1974) found that those from Merinos rated the lowest,
although they had very light carcase weights with minimal
fat cover. Safari et al. (2001) reported no difference in
overall acceptability, tenderness or juiciness for roasted
loin meat from rst cross (BLM), Merino or second cross
lambs. In another study of the hindleg (m. biceps femoris),
Merino lambs had lower juiciness, avour liking and
overall liking scores than BLM and second cross lambs, but
were similar to PDM lambs (Hopkins et al., 2005c) when
assessed by consumers.
Dranseld et al. (1979), Edwards et al. (1982), Crouse
et al. (1981), Crouse (1983), Ellis et al. (1997) and Esenbuga
et al. (2001) reported no signicant differences between
genotypes. Hoffman et al. (2003) did nd initial juiciness
of the SM was lower from Suffolk Merino than other

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D.L. Hopkins et al. / Small Ruminant Research 101 (2011) 160172

genotypes, but it was of no practical signicance. In other

work from South Africa, Webb et al. (1994) reported that
roasted loin meat from South African Mutton Merino
(SAMM) lambs had better avour and overall acceptability than from Dorper lambs. This was attributed to the
signicantly higher fat levels in the Dorper, but it is noteworthy that the subcutaneous fat of the Dorpers also had
higher levels of unsaturated fatty acids which the authors
suggested could have depressed the avour scores of the
meat. The results may also have reected the composition
of the taste panel which has been shown to impact on

scores even when they are trained (Sanudo

et al., 1998). A
decrease in overall liking for meat from Texel over Scottish
Blackface lambs (Navajas et al., 2008) seems to have been
due to differences in tenderness, with inconsistent differences for juiciness across the various cuts. There does not
appear to be consistent genotype effects on eating quality
traits.
2.3. Impact on intramuscular fat
Thompson (2004) concluded that as the variation in
tenderness of meat is reduced through uptake of grading schemes such as Meat Standards Australia (MSA),
marbling or IMF will become a more important determinant of palatability because of its contribution to juiciness
and avour. Many carcass grading systems, particularly
for beef, use marbling score as a major factor in predicting eating quality because higher IMF is considered
to improve eating quality traits (Hocquette et al., 2010).
For lamb, a target of 5% IMF is required to help ensure
a good every day (grade 3 out of a 5 level system)
score despite the literature suggesting that marbling is
only one component of eating quality (Hopkins et al.,
2006).
There were no differences in loin IMF between Merino
and various crossbred types with a mean level of 4.8%
(Hopkins et al., 2005c). However Daz et al. (2005) showed
variation in IMF between breeds for loins, but the comparison was confounded by differences in production systems.
There were some differences in IMF between 5 genotypes
(Merino, BLM and rst and second cross types with sires
having different breeding values for growth and muscling)
in a rigorous study in which the animals were run together
and slaughtered at 4 ages from 4 to 22 months (McPhee
et al., 2008). The BLM had the highest IMF across both ewes
and wethers and the highest increases in IMF between 14
and 22 months.
The claim for Merino sired lambs to have 0.42% more
IMF than terminal and maternal sired lambs (Pethick et
al., 2010) was contrary to Hopkins et al. (2007b) who
found that Merino lambs actually had the lowest levels of
IMF. Merinos in the Pethick et al. (2010) study were older
than the comparative types. Lambe et al. (2008) reported
Texel lambs selected for muscling and reduced fatness had
very low IMF levels (1.60%) in loins compared to Scottish
Blackface (2.3%), although the difference was unlikely to
be signicant. Indeed the low values for IMF suggest the
meat was sub-optimal for good eating quality. This report
of Lambe et al. (2008) illustrates that in some specic cases
extreme genotypes can decrease traits like eating quality

and IMF and the sheep industry needs screening systems

to prevent the production of such genotypes.
2.4. Impact on pH and colour
High mean pH values for meat affect keeping quality and
can adversely affect avour and aroma (Young et al., 1993).
Shelf life is reduced when pH exceeds 5.8 (Egan and Shay,
1988) and as pH increases meat becomes darker (Fogarty
et al., 2000), affecting consumer purchase decisions.
Higher muscle ultimate pH for Merino and BLM lambs
compared to terminal sire second cross lambs has been
reported (Hopkins and Fogarty, 1998; Gardner et al., 1999;
Fogarty et al., 2000; Hopkins et al., 2007a). Under high
stress commercial slaughter conditions, the Merino loses
a greater amount of muscle glycogen than other types
(Gardner et al., 1999), but under low stress slaughter the
meat from Merinos can have a similar ultimate pH as that
from other genotypes (Hopkins et al., 2005c). The m. semitendinosus muscle has more type 2X fast glycolytic bres
than the m. longissimus (Greenwood et al., 2007) and as the
content of these bres increases so do the nal pH levels
(Gardner et al., 2006). This effect was clearly evident in the
Merino lambs studied by Hopkins et al. (2007a), which had
a higher ultimate pH indicative of a faster rate of glycogen depletion during the pre-slaughter period. However,
it does not appear that these Merinos were more agitated
than the other genotypes as measured by an isolation test
(Warner et al., 2006). Hopkins et al. (2005a) reported signicant differences in muscle pH among Merino wether
bloodlines for both the loin and the m. semitendinosus with
higher muscle pH for the Merino superne wool bloodlines.
In contrast, Fogarty et al. (2003) found that the broad wool
strain of Merino rams had a higher loin pH, than both ne
and medium wool strains.
The higher muscle pH for Merinos compared to other
genotypes was consistent across 3 different muscles for
lambs sourced and slaughtered in New South Wales, but the
effect was not found in lambs sourced and slaughtered in
Victoria (Hopkins et al., 2005c). The Victorian animals had a
much greater weight gain than those in New South Wales
which may have led to higher levels of muscle glycogen
and, together with a lower stress slaughter environment,
may have resulted in low pH meat from the Merinos. The
results of Gardner et al. (1999) support this because Merinos slaughtered at a research abattoir did not exhibit high
pH values. The higher ultimate pH reported by Hopkins
et al. (2005c) cannot be ascribed to an increasing proportion
of Merino genes as the pH levels for the BLM and Merino
animals were similar for the loin at least, which is also
supported by other reports (Hopkins and Fogarty, 1998;
Hopkins et al., 2007a).
However, Hopkins et al. (2007a) reported a higher muscle pH in the m. semitendinosus of Merino lambs compared
with all other crosses including BLM lambs. The samples
were aged for 5 days and measured after homogenisation
to ensure an accurate measure of ultimate pH, overcoming
electrode placement issues which can arise in situ. A New
Zealand experiment with various breeds of terminal sires
joined to Merino ewes, reported that loins from Merino
lambs had a signicantly higher pH than BLM and PDM

D.L. Hopkins et al. / Small Ruminant Research 101 (2011) 160172

lambs (Young et al., 1993). Loin pH values were also greater

for BLM compared to PD sired lambs (Fogarty et al., 2000;
Hopkins et al., 2007a). Higher loin pH values have also been
reported in lambs from Merino dams compared to those
from Dohne Merino or SAMM dams (Hoffman et al., 2003).
There is little evidence of differences in muscle pH
between British breed crossbred lambs (Dranseld et al.,
1979; Carson et al., 2001), although purebred Scottish
Blackface lambs had higher loin pH levels than crossbred
lambs in the latter study with no explanation for the result.
Other studies involving purebreds and crossbreds have not
found any differences in muscle pH levels with Romney and
East Friesian (Purchas et al., 2002) or the Elliottdale carpet
wool breed (Hopkins et al., 1992).
While meat from Merinos is more susceptible to high
pH than from other types, there is little evidence of type
impacting on objectively measured fresh colour (Dranseld
et al., 1979; Fogarty et al., 2000, 2005; Hopkins et al.,
2005c, 2007a). Even when Merino lambs produce meat
with a higher pH than other types, they do not produce
darker fresh meat, which may reect the low correlation
between pH and L* values (Menzies and Hopkins, 1996).
Loin muscle from Merinos browns quicker and to a greater
extent through formation of metmyoglobin than muscle
from the other types (Warner et al., 2007). This could
explain the industry perception that meat from Merinos
has poor colour. Loin meat from Romney lambs was darker
than that from crossbred Texel lambs (Purchas et al., 2002),
which was independent of pH, with time of slaughter
having a greater effect. The loin meat from these Romney lambs was below the acceptable threshold lightness
value of 34 (L*) established by consumer evaluation (Khliji
et al., 2010). There is variation within the Merino breed
for muscle pH, but the physiological basis of higher pH
levels in the Merino remains to be fully established and,
given the importance of this breed to sheep production
in a number of countries, is an area worthy of further
research.
2.5. Impact on iron, zinc and omega-3 fatty acids
The importance of red meat in human diets in achieving
recommended levels of iron (Fe), zinc (Zn) and omega-3
fatty acids has been proposed as a future key marketing tool (Pethick et al., 2006). Iron is a component of
several important proteins such as haemoglobin and myoglobin, zinc is a component of various enzymes that help
maintain structural integrity of proteins and regulates
gene expression and the omega-3 fatty acids, particularly
eicosapentaenoic (EPA) and docosahexaenoic (DHA), have
cardiovascular and anti-inammatory benets (NHMRC,
2006).
There is little information on the impact of genotype
on these traits and the muscle type needs to be considered as levels vary especially for Fe (Lin et al., 1989) and
omega-3 fatty acids (Ponnampalam et al., 2010). Pearce
et al. (2009) reported the BLM had the highest Fe level in
the m. semimembranosus, but not in two other muscles
with smaller differences for Zn. There was an interaction between genotype and muscle, with Merino and BLM
m. semimembranosus and m. semitendinosus having 5%

163

higher Zn levels than PD progeny (Pearce et al., 2009). In a

study where lambs were given excess levels of dietary Fe
and Zn, Field et al. (1985) found that Suffolk-sired lambs
had a lower level of Zn in their muscle than Southdownsired lambs, but there was no difference in Fe levels. In
a larger study based on 2000 lambs of 3 main genotypes
by 94 sires, Pannier et al. (2010) reported no differences
between genotypes for Fe levels, but elevated levels of Zn
in BLM compared to Merino and Terminal sire cross lambs.
While this was indicative of more oxidative muscle bre
type (Gardner et al., 2007), the absolute levels in this study
were approximately half that previously reported from
retail sampling of lamb in Australia (Williams, 2007) and
elsewhere (Lin et al., 1989) and thus any genotype induced
reduction must be validated. Hoffman et al. (2003) found
much higher Zn levels across 6 different types (with no type
effects) than those reported by either Pearce et al. (2009)
or Pannier et al. (2010). However Hoffman et al. (2003)
did report genotype differences for Fe, with lower absolute levels than reported by others (Pearce et al., 2009;
Pannier et al., 2010). The study by Hoffman et al. (2003)
was only based on 42 lambs, which was considerably less
than those of Pearce et al. (2009) or Pannier et al. (2010).
There is obviously a need to more comprehensively document the importance of genotype as a source of variation
for minerals like Fe and Zn.
Ponnampalam et al. (2009) reported a decrease in the
ratio of polyunsaturated to saturated fats as the level of
Merino genes decreased, even after adjustment for IMF%,
although there was no genotype effect for the levels of individual fatty acids. Similarly, Hoffman et al. (2003) found
no genotype effect for the level of EPA, but Suffolk (S)
cross SAMM m. semimembranosus had lower DHA and
docosapentaenoic acid (DPA) than from S M, which is of
interest given EPA is synthesized into DHA. In contrast,
the S SAMM had the second highest level of DPA and
the S M the lowest (Hoffman et al., 2003). The results
strongly suggest that differences due to genotype are likely
to occur in the wider sheep population and Ponnampalam
et al. (2009) suggested that crossbred lambs may require
a higher level of dietary intervention to attain the same
level of omega-3 fatty acids as that found in Merinos In a
study of two Turkish breeds, Demirel et al. (2006) reported
an interaction between breed type and the type of nutrition for the level of DHA in the m. longissimus, suggesting
an effect on the incorporation of fatty acids into phospholipids. This could reect a difference in fat levels (De Smet
et al., 2004) specically IMF%, but Demirel et al. (2006) did
not adjust for the signicant differences in IMF% between
breeds. This same problem is evident in the data of Fisher
et al. (2000), where the different fat levels of the Welsh
Mountain and Soay were not considered in the comparison of the levels of the various fatty acids, where the Soay
was much leaner and had a signicantly higher level of EPA
in the m. semimembranosus.
Given the claims that a greater ratio of polyunsaturated
to saturated fat and a lower ratio of omega-6 to omega-3 in
meat would be desirable to lower the incidence of human
metabolic diseases such as heart disease, inammation and
mental health (Wood et al., 2003; Scollan et al., 2006)
it is important to develop both genetic and non-genetic

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D.L. Hopkins et al. / Small Ruminant Research 101 (2011) 160172

approaches to manipulating the level of the omega-3 fatty

acids.
3. Major gene effects
3.1. The callipyge effect
The callipyge effect is manifested as a major increase in
hindquarter muscling caused by muscle hypertrophy and
was originally identied in USA ocks (Jackson et al., 1993).
The condition is due to a mutation on chromosome 18 and is
expressed in heterozygous animals with maternal imprinting meaning that the mutation is only expressed when
inherited from the sire (Cockett et al., 1994; Shackelford
et al., 1998). The m. longissimus muscle of callipyge
lambs has much higher levels of shear force (Shackelford
et al., 1997; Taylor and Koohmaraie, 1998; Freking et al.,
1999), with signicantly reduced myobrillar fragmentation index values indicative of a decrease in protein
degradation (Koohmaraie et al., 1995). It appears that postmortem proteolysis is delayed in muscle from callipyge
lambs due to higher levels of calpastatin (Koohmaraie
et al., 1995; Geesink and Koohmaraie, 1999). The extent
of proteolysis is also reduced (Geesink and Koohmaraie,
1999), with much less structural disruption of the sarcomere (Taylor and Koohmaraie, 1998). It appears the effect
is manifested solely through the myobrillar component
of the m. longissimus muscle with no impact via collagen
(Field et al., 1996). The IMF% is reduced, with no effect on
the rate of pH decline (Koohmaraie et al., 1995), although
there is a change in the proportion and size of muscle
bres with an increase in the proportion of glycolytic bres
(Koohmaraie et al., 1995). While there is increased toughness (Abdulkhaliq et al., 2007) and reduced juiciness of
the m. longissimus from the callipyge lambs there is no
reduction in tenderness in other muscles (Shackelford et al.,
1997; Abdulkhaliq et al., 2002). Increased cooking loss in
meat from callipyge lambs has been reported (Abdulkhaliq
et al., 2007), although not always (Shackelford et al., 1997).
3.2. Carwell effect
An industry progeny testing program indicated that
some rams from an Australian Poll Dorset stud (Carwell)
produced progeny with a large positive deviation for eye
muscle area (Banks et al., 1995). Subsequently one of the
4 markers for the callipyge gene, TGLA 122 on chromosome 18, was found to be present in Australian animals
exhibiting increased muscling (Barendse, 1995; Nicoll et al.,
1998). The Carwell mutation is thought to be maternally
imprinted and a marker test is now available under the
trademark LoinMax (Dodds, 2007).
The Carwell effect increases the weight of the m.
longissimus by 810% (Nicoll et al., 1998; Hopkins and
Fogarty, 1998). It increased shear force in the m. longissimus by 35% in male lambs that had high pre-slaughter
growth rate (Hopkins and Fogarty, 1998), but there was
no effect in female lambs under low growth rate (Hopkins
et al., 1997). In a more recent study including progeny
by the same Carwell sire, Hopkins et al. (2007a) found
no increase in toughness of the m. longissimus or m.

semimembranosus or any other effects on pH or colour

traits. In contrast, Jopson et al. (2001) found that loins,
frozen at 24 h post-mortem, from carrier progeny were
tougher and unacceptable, based on values of 2730 Newtons shear force (Hopkins et al., 2006), although the
differences disappeared when the loins were aged for 6
weeks. As the lambs were processed under an accelerated conditioning procedure this indicates the differences
in shear force could be overcome by myobrillar protein
degradation. While the Carwell mutation does not have
such an impact on carcase and meat traits as callipyge, its
effects can be modied by the production and processing
environments. Unless appropriate production and processing strategies are in place, there is a need for caution when
selecting sires to avoid detrimental effects on meat quality
of progeny.
3.3. Texel muscling
A quantitative trait locus (QTL) that increases m. longissimus weight by less than 10% (Macfarlane et al., 2009) has
been identied on chromosome 18 (TM-QTL) in Texel sheep
(Walling et al., 2004). In male progeny of sires carrying the
mutation, there was increased shear force and reduced IMF
of the m. longissimus. There was also a signicant interaction with sex as there was no effect in female lambs (Lambe
et al., 2010a). In this study there was no effect on shear
force or IMF of the m. vastus lateralis a muscle from the
hind leg or for the quadriceps femoris group of muscles.
The levels of IMF reported for the m. longissimus in the
Lambe et al. (2010a) study were very low compared with
others (Hopkins et al., 2007a,b) and were below the level
for optimal eating quality (Hopkins et al., 2006). While variable times between death and electrical stimulation and
variable ageing periods may have impacted on the results,
the greater variation for shear force in lambs carrying the
mutation highlights the need for caution in the dispersal of
carrier genetics.
In follow up work, Lambe et al. (2010b) reported that
there was a higher proportion of high shear force longissimus muscles among carriers after 3 and 5 days of ageing,
but not after 9 days. The absolute levels of shear force for
all the genotypes studied suggest unacceptable eating quality (Hopkins et al., 2006), although the exact relationship
between shear force values and sensory assessments will
vary according to the instrument used to measure shear
force (Hopkins et al., 2011). Their relatively high shear
force values (Lambe et al., 2010b) may indicate ineffective
electrical stimulation, which could have been veried by
collection of pH decline data.
3.4. Myostatin
A variant of the myostatin gene (growth differentiation
factor 8; GDF8) on chromosome 2 has been reported to be
segregating in Texel sheep in New Zealand (Broad et al.,
2000) and a marker test (MyoMAX ) is available (Dodds,
2007). The mutation affects muscle and fat traits in some
Texel sired lambs (Johnson et al., 2005a) with no effect on
shear force, colour and pH of the m. longissimus and m.
semimembranosus (Johnson et al., 2005b). The sarcomere
length and shear force data for the m. semimembranosus

D.L. Hopkins et al. / Small Ruminant Research 101 (2011) 160172

muscle in the latter study suggested that cold induced

shortening had occurred in the carcases. Laville et al. (2004)
reported a shift towards more glycolytic muscle bres in
lambs carrying the mutation, but no other meat quality
traits were studied. Kijas et al. (2007) reported a reduction
in IMF% and sensory scores, but no effect on shear force,
although there was evidence of another QTL that affected
meat quality traits.
There is a need for robust sensory assessment of all these
mutations that affect muscling and meat quality. There has
been little investigation of the basis for any increase in
toughness caused by some of these mutations that impact
on protein turnover and this should be pursued.

165

weight, whereas there was a positive unfavourable correlation between shear force and weaning weight (0.45 0.15).
While Safari et al. (2008) reported that the genetic correlations between meat colour measures and reproductive
traits were small and generally positive. Afolayan et al.
(2008) reported by contrast large negative genetic correlations between meat colour measures and reproductive
traits. More precise estimates of genetic correlations are
required among a wider range of meat quality traits and
between the meat quality traits and other traits currently
included in breeding programs.
5. Relationships with breeding values for live
animal traits

4. Heritability and genetic correlations

There are few estimates of heritability for meat quality traits in sheep (Safari et al., 2005) and these and more
recent reports are summarised in Table 1. Shear force
appears to be moderately to highly heritable (Table 1) with
higher estimates for aged meat (Mortimer et al., 2010),
which is of interest as the effects of processing should
be diminished with ageing. The heritability estimates for
IMF are also moderate to high (Karamichou et al., 2006b;
Mortimer et al., 2010). Estimates of the genetic correlation
between IMF and shear force of 0.54 0.24 (Karamichou
et al., 2006b) and 0.69 0.11 (Mortimer et al., 2010) suggest that selection for increasing IMF will have a favourable
affect on shear force.
The heritability estimates for eating quality traits were
low (Karamichou et al., 2006b, 2007), albeit from a very
small data set. Positive genetic correlations were found
between IMF and the sensory traits juiciness and avour
(0.12 0.06 and 0.20 0.06, respectively) (Karamichou
et al., 2006b).
The estimates of heritability for meat pH and measures
of meat colour were generally low to moderate (Table 1)
with low phenotypic variation which suggests selection
response for these traits would be slow. There appears to be
a favourable genetic correlation (0.6) between pH and L*
colour values in Merino hogget rams (Fogarty et al., 2003;
Greeff et al., 2008), whereas others have reported correlations not signicantly different from zero in meat sheep
breeds (Ingham et al., 2007; Payne et al., 2009).
The preliminary report on the minerals iron and zinc
in meat suggest low to moderate heritabilities (Table 1),
although there is one very high estimate for zinc from limited data (Bennett and Field, 1985). More precise estimates
will be available as more data are obtained (Mortimer
et al., 2010). The estimates of heritability for the omega3 fatty acids EPA and DHA range from 0.16 to 0.29, with
relatively high phenotypic variation, supporting the conclusions of Karamichou et al. (2006a) and Greeff et al.
(2007) that it may be possible to improve omega-3 fatty
acids through selection. Karamichou et al. (2007) reported
negative residual correlations between both EPA and DHA
with overall eating quality liking. There are few reports
of genetic correlations between meat quality traits, production traits (reviewed by Mortimer et al., 2010) and
reproduction traits. Mortimer et al. (2010) reported a negative correlation (0.19 0.11) between IMF and weaning

Although there have been few reported estimates of

genetic correlations between meat quality and production
traits, several recent studies have evaluated the associations between estimated breeding values (EBVs) for various
production traits and the meat quality and muscle biochemistry in their progeny (Hopkins et al., 2005b; Gardner
et al., 2006). These associations assist industry to understand the implications of selection for production traits on
meat quality. Early work by Woodhams et al. (1966) found
no effect of sire on tenderness of loins among crossbred
lambs sired by 6 Southdown rams, although individual sires
can affect tenderness (Hopkins and Fogarty, 1998). Table 2
summarises the impact of sire EBVs for growth, muscling
and fat on tenderness, IMF and other aspects of eating quality from several reports.
In the rst study involving 9 Poll Dorset sires with
ranges in EBVs for growth, fatness and muscling, there was
no impact of any of these EBVs on shear force of either the
m. longissimus or m. semimembranosus muscles in electrically stimulated carcasses, although there was a decrease
in consumer assessed tenderness as sire EBV for eye muscle
depth increased (Hopkins et al., 2005b). In a larger follow up study there was no effect of growth, fatness or
muscling EBVs on shear force of the m. longissimus or
the m. semimembranosus across 16 sires of 3 genotypes,
although there was an 8.3% higher shear force (P < 0.05)
for the m. semimembranosus from high muscle EBV Poll
Dorset sires compared with other types irrespective of
whether loin pH was included as a covariate (Hopkins
et al., 2007a). In another study involving 637 crossbred
lambs by 20 sires (Hopkins et al., 2007b); the semimembranosus from early-weaned, restricted growth lambs had
a signicantly lower shear force. In the same study, sensory
tenderness increased as sire EBV for fat increased with the
range of 4 points for sensory tenderness across the range
in the EBVs. The only estimates of genetic correlations with
shear force are small for muscling (0.15 0.17) and fatness (0.10 0.16), but larger positive for body weight
(0.30 0.15) (Mortimer et al., 2010).
Other eating quality traits declined as the sire EBV for
muscling increased (Hopkins et al., 2005b). This negative
relationship could be partly explained by the reduced IMF
as marbling is positively related to eating quality traits
(Karamichou et al., 2006b). There is a positive relationship between IMF and sire EBV for fat, which is consistent
with a small positive genetic correlation (Mortimer et al.,

166

Table 1
Estimates of heritability (h2 ), standard error (s.e.) and phenotypic variance ( 2 P ) for meat quality traits.
2P

h2

s.e.

Shear force
(kg) 2 day aged
Initial (kg) 10 day aged
(N)
Peak (N) 1 day aged
Peak (N) 5 day aged

0.28
0.39
0.44
0.27
0.38

0.10
0.16

3.84

0.07
0.08

59.30
39.20

0.15
0.21
0.11
0.22

0.13
0.12
0.11
0.13

99.7
35.1
23.8
51.7

0.32
0.39

0.09
0.05

0.27
0.21
0.18
0.22
0.09
0.12
0.10
0.09

0.09
0.14
0.07
0.03
0.09

0.14
0.15
0.23
0.45
0.18
0.29
0.21

0.07
0.12
0.07
0.19
0.03
0.04

3.21

0.02
0.45
0.10
0.04

0.06
0.19
0.06
0.10

2.17
2.01

Eating quality (0100)

Tenderness
Juiciness
Flavour
Overall liking
Intramuscular fat
Mg/100 g muscle
%
pH
Measured 24 h post-mortem
Measured 24 h post-mortem
Measured 24 h post-mortem
Measured 24 h and 48 hr post-mortem
Measured 48 h post-mortem
Measured 24 h post-mortem
Measured 1924 h post-mortem
Colourc (Lightness, L*)
Colour meat calibrated with a white tile
Calibration details not given
Calibration details not given
Calibration details not given
Colour meat calibrated with a white tile
Colour meat calibrated with a white tile
Colour meat calibrated with a white tile
Redness (a*)

0.03
0.04

0.63

0.01
0.108

0.006
0.025

3.43
2.05

Number
of records

Breed base

Reference

802a
349
586a , b
1637
1759

Rambouillet, Columbia and Corriedale crosses

Scottish Blackface
Merino and meat crosses
Merino, BLM, Terminal Merino and Terminal BLM

Botkin et al. (1969)

Karamichou et al. (2006b)
Cloete et al. (2008)
Mortimer et al. (2010)

Scottish Blackface - trained panels

Karamichou et al. (2006b)

349
3811

Scottish Blackface
Merino, BLM, Terminal Merino and Terminal BLM

Karamichou et al. (2006b)

Mortimer et al. (2010)

957
349
1330
5700
672
6565
3709
3766

Merino
Scottish Blackface
Merino cross.
Merino
Merino and meat crosses
Terminal cross
Merino, BLM, Terminal Merino and Terminal BLM
Measured on the m. semitendinosus

Fogarty et al. (2003)

Karamichou et al. (2006b)
Ingham et al. (2007)
Greeff et al. (2008)
Cloete et al. (2008)
Payne et al. (2009)
Mortimer et al. (2010)

1035
349
1913
580
5107
6565
3432

Merino
Scottish Blackface
Merino cross
Merino and meat crosses
Merino
Terminal cross
Merino, BLM, Terminal Merino and Terminal BLM

Fogarty et al. (2003)

Karamichou et al. (2006b)
Ingham et al. (2007)
Cloete et al. (2008)
Greeff et al. (2008)
Payne et al. (2009)
Mortimer et al. (2010)

1011
349
1331
580

Merino
Scottish Blackface

Fogarty et al. (2003)

Karamichou et al. (2006b)
Ingham et al. (2007)
Cloete et al. (2008)

349
349
349
349

Merino and meat crosses

D.L. Hopkins et al. / Small Ruminant Research 101 (2011) 160172

Trait

Minerals
Iron (mg/kg dried muscle tissue)

0.21

0.38

Iron (mg/kg wet muscle tissue)

Zinc (mg/kg dried muscle tissue)
Zinc (mg/kg wet muscle tissue)

0.12
0.92
0.21

0.05
0.48
0.06

Long chain fatty acids

Eicosapentaenoic acid (EPA, mg/100 g wet muscle tissue)
EPA
EPA
Docosahexaenoic acid (DHA, mg/100 g wet muscle tissue)
DHA
DHA

0.21
0.18
0.29
0.16
0.19
0.25

0.13
0.07
0.07
0.10
0.08
0.06

Derived for the m. longissimus unless otherwise indicated.

a
No information given to indicate if peak or initial yield.
b
Ageing period not given.
c
Measured under the same conditions as the pH.

0.03
0.10

1.43
0.28

12.77
12.83

0.129
0.039
0.032
0.051

5080
6565
3431
1156

Merino
Terminal cross
Merino, BLM, Terminal Merino and Terminal BLM

Greeff et al. (2008)

Payne et al. (2009)
Mortimer et al. (2010)

100

Suffolk or Southdown sires Perendale, Romney,

Coopworth, Merino-Perendale and Merino-Romney
ewes
Merino, BLM, Terminal Merino and Terminal BLM

Bennett and Field (1985)

1915
100
1915
350
1109
1919
350
1069
1915

Scottish Blackface
Merino
Merino, BLM, Terminal Merino and Terminal BLM
Scottish Blackface
Merino
Merino, BLM, Terminal Merino and Terminal BLM

Mortimer et al. (2010)

Bennett and Field (1985)
Mortimer et al. (2010)
Karamichou et al. (2006a)
Greeff et al. (2007)
Mortimer et al. (2010)
Karamichou et al. (2006a)
Greeff et al., 2007
Mortimer et al. (2010)

D.L. Hopkins et al. / Small Ruminant Research 101 (2011) 160172

0.03

Metmyoglobin/Oxymyoglobin (indicator of browness)

0.10
0.19
0.06
0.40

167

For m. longissimus
For m. longissimus
For m. longissimus
For m. longissimus
For m. semitendinosus
0.05
0.05
0.05
0.05
0.05
0.05
0.05
0.05
0.05
0.05
0.05
0.14 0.06

0.027
0.012
0.041 0.015
Not given

Lightness (L*)
Redness (a*)
Zinc (mg/100 g)

pH

0.15 0.06

0.05

0.11 0.06

Eating quality (score 0100)

Juiciness
Overall liking
Flavour liking
Flavour liking
Intramuscular fat

0.019 0.004a

1.59 0.57
1.32 0.58
1.37 0.49

1.30 0.60

1.66 0.76
1.91 0.59
Tenderness (score)

PWWT, postweaning weight; PEMD, postweaning muscle depth; PFAT, postweaning fat depth; YEMD, yearling muscle depth; YFAT, yearling fat depth.
a
Log scale.

Hopkins et al. (2005b)

Hopkins et al. (2005b)
Hopkins et al. (2005b)
Hopkins et al. (2007b)
Hopkins et al., 2005b
Hopkins et al. (2007b)
Pethick et al. (2010)
Hopkins et al. (2007a)
Hopkins et al. (2007a)
Hopkins et al. (2005b)
Hopkins et al. (2007a)
Pannier et al. (2010)

Consumer panel (m. longissimus).

Consumer panel (m. longissimus).
Shear force (m.
semimembranosus low growth)
0.05
0.05
0.01

Background information
Signicance
level
YFAT (mm)
YEMD (mm)
PFAT (mm)
PEMD (mm)
PWWT (kg)
Trait

Table 2
Summary of the signicant regression coefcients for meat quality traits on sire estimated breeding values.

Hopkins et al. (2005b)

Hopkins et al. (2007b)
Hopkins et al. (2007b)

D.L. Hopkins et al. / Small Ruminant Research 101 (2011) 160172

Reference

168

2010) and responses in IMF reported in divergent lines

selected for carcass lean content (Karamichou et al., 2006b).
Sire group had no effect on myobrillar fragmentation
index (Hopkins and Hegarty, 2004), which indicates that
the poorer consumer scored tenderness was not due to a
decrease in proteolytic degradation in the meat that was
aged for 5 days. Despite the high error around the regression coefcients for the relationships between sire EBVs
and sensory traits, the results suggest that selection for
increased muscling may impact adversely on sensory traits.
Flavour score increased as the sire EBV for fat increased.
There was also an increase in avour score for meat that
was at a higher temperature when the pH had reached
6.0, with the score increasing by 6.6 units over the range
from 10 to 25 C (Hopkins et al., 2007b). For overall liking
score there was a signicant interaction between feeding
level and the sire EBV for fat, with overall liking increasing
as the EBV increased (regression 2.80 0.89) for restricted
growth lambs but not for high growth lambs (regression
0.01 0.89) (Hopkins et al., 2007b). This emphasises the
importance of IMF in eating quality. The positive effect of
fat EBV on sensory traits must be balanced by the negative effect on lean meat production (Hopkins et al., 2007c),
emphasising the need for EBVs to be used in an index. It
should also be stressed that the range in EBVs for a trait will
determine the absolute signicance of an effect. For example, a reduction of 4 sensory points was predicted for the
range in sire EBVs for fat in Hopkins et al. (2007b), which
was less than half the effect for the range of sire EBVs for
muscle in Hopkins et al. (2005b).
Gardner et al. (2006) found no association between EBVs
and pH among a sub-sample of lambs from the Hopkins
et al. (2005b) study, which is consistent with the generally small genetic correlations reported (Ingham et al.,
2007; Greeff et al., 2008; Payne et al., 2009; Mortimer et
al., 2010). There was however a signicant reduction in
myoglobin as the EBV for muscle increased in progeny on
a high plane of nutrition (Gardner et al., 2006). This suggested a potential reduction in oxidative capacity in the
muscle, but this did not impact on pH or colour measures
in the larger group (Hopkins et al., 2005b). Hopkins et al.
(2007a) found a decrease in redness (a* values) as the EBV
for muscle increased, suggesting a reduction in oxidative
capacity. Gardner et al. (2006) also found a rise in lactate
dehydrogenase activity as sire EBV for muscle increased
which indicates an increase in glycolytic bres. Pannier
et al. (2010) reported a reduced level of zinc in the m.
longissimus as sire EBV for muscle increased (Table 2),
although there was no effect on iron levels. No relationship between omega-3 fatty acids and sire EBVs has been
reported.
6. Breeding programs
Substantial annual rates of genetic improvement are
occurring in the Australian (Swan et al., 2009), New Zealand
(Young and Amer, 2009) and UK (Amer et al., 2007) sheep
industries. The use of performance recording and genetic
evaluation technologies in Australia over the last 2 decades
has resulted in an annual increase of $2.00 per ewe in terminal sire breeds through improvements in growth, leanness

D.L. Hopkins et al. / Small Ruminant Research 101 (2011) 160172

and muscling (Swan et al., 2009). There has been little

opportunity to include meat quality traits (Simm et al.,
2009), although market research indicates they are becoming more important to consumers (Pethick et al., 2006;
Bermingham et al., 2008).
It is critical to understand the impact of current industry
breeding programs on meat quality and avoid any detrimental changes (Fogarty, 2009), although at present there
is no economic incentive to justify inclusion of meat quality traits (Simm et al., 2009). It is clear from this review
that genetic variation exists for many meat quality traits
and genetic parameter estimates are becoming available.
While there may be some unfavourable genetic correlations between meat quality and production traits, they
appear to be sufciently small to be countered by appropriate breeding objectives and selection indexes. For New
Zealand sheep, Payne et al. (2009) have predicted that index
selection for growth rate and meat yield would result in little change in meat quality traits, except for small increases
in meat lightness and pH and a decrease in fat yellowness.
Incorporation of meat quality traits into industry breeding programs, requires development of an appropriate
breeding objective including meat quality and production
traits (Bishop and Karamichou, 2009), as well as identifying the selection criteria. In Australia, estimates of genetic
variance and correlations among a wide range of meat
quality traits that are difcult and expensive to measure,
are being obtained in the Sheep CRC Information Nucleus
(van der Werf et al., 2010; Mortimer et al., 2010). Direct
measurements of meat quality traits are expensive and
require slaughter of the animals. Estimated breeding values for some meat quality traits are now being developed to
allow breeders to screen potential sires. Two-stage selection could reduce costs by live assessment of indirect
predictors of the meat quality trait and direct measurement
of a subset of animals (Simm et al., 2009). Use of genomic
information has the potential to increase the accuracy of
selection and allow direct selection of breeding animals at
a young age (Daetwyler et al., 2010). van der Werf (2009)
estimated a 1632% increase in value from genomic selection by including carcass traits in a terminal sire objective,
depending on the level of accuracy. Genomic information
for meat quality traits is currently being generated by the
Information Nucleus.
7. Conclusions
There appear to be few differences between genotypes
of sheep for meat quality traits, although the Merino has a
tendency for a high pH. Given the importance of this breed
to sheep production in a number of countries research to
establish the physiological basis of the higher pH levels in
the Merino is worthy of further investigation. The literature
also shows that there may be adverse effects on meat quality for extreme genotypes, such as extreme muscling and
major gene effects. Given the importance of eating quality
and IMF to consumers and the evidence that some extreme
genotypes can change these traits unfavourably, there is
room for more investigation of the basis of any increase in
toughness caused by some of the more recently identied
mutations which impact on protein turnover. Estimates of

169

genetic parameters are only just becoming available and

indicate there is genetic variation for most meat quality
traits. More accurate estimates of the genetic correlations
between meat quality and production traits are required
to develop appropriate objectives and selection criteria for
industry breeding programs. Increasingly, it will be important to develop both genetic and management approaches
to manipulate the level of traits like omega-3 fatty acids, if
consumer demand follows expectations. Incorporation of
molecular information into breeding programs also offers
potential for improvement of meat quality. Both these areas
of research are currently being addressed in the Sheep CRC
Information Nucleus.
Conict of interest
The authors Drs Hopkins, Fogarty and Mortimer have
no nancial or personal relationship with other people or
organisations that could inappropriately inuence or bias
the paper entitled Genetic related effects on sheep meat
quality.
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