68

JacksonAnimal
and Blanton,
Jr. 17:6874
The Professional
Scientist

: The Callipyge Gene

Rin Sheep
EVIEW

S. P. JACKSON1 and J. R. BLANTON, JR.

Department of Animal Science and Food Technology, Texas Tech University,
Lubbock, TX 79409

Introduction

pattern was subsequently used to

identify this variant as callipyge
In recent years, there has been
(CLPG), which comes from the Greek
considerable economic pressure to
language meaning beautiful butproduce meat animals as efficiently as tocks (8). Since the discovery of the
possible while still meeting consumer CLPG phenotype, extensive research
needs. As a result of these pressures,
has been conducted to determine the
many scientists and producers have
effects of this mutation on the
been looking more closely at utilizing biological systems of sheep and what
the genetic variation available in
effect it may have on commercial
current meat animal populations to
lamb production. Therefore, this
meet these goals. Although some of
review will attempt to summarize the
these genetic mutations can improve current research on CLPG animals to
animal performance, many are
provide an unbiased source of inforassociated with characteristics that
mation for researchers and producers.
make daily livestock management
more difficult. For example, dystocia
problems are associated with double
muscle cattle and susceptibility to
stress in lean pigs with the ryanodine
In 1983, a Dorset lamb was born
receptor mutation (4, 6, 38, 39).
that
transmitted postnatal developHowever, in 1983, a unique muscling
ment
of extreme heavy muscling to
phenotype was discovered in sheep
his
progeny
in a non-Mendalian
that improved overall performance,
inheritance
pattern
(17). Since this
but lacked many of the production
time,
researchers
have
determined
problems found in the cattle and pig
that
the
CLPG
phenotype
is a single
mutations. This mutation is identifigene
that
is
dominant
when
inherable by the extensive postnatal
ited
from
the
sire.
The
occurrence
of
muscle accretion in the hindquarters
this
phenotype
is
50%
when
a
and loin of affected animals (Figure
heterozygous male is mated to a
1) (7, 8). This unique muscling
population of normal, noncarrier
females. However, mating heterozygous carrier females to normal
1To whom correspondence should be ad- noncarrier males results in 100%
dressed: sajackso@ttacs.ttu.edu
normal phenotype. When lambs

Chromosomal Location
and Transmission

from the latter matings are

phenotyped, 50% carry the CLPG
gene, which results in 50% CLPG
expression in lambs produced from
these males when mated to normal
noncarrier females. This unique
inheritance pattern has been termed
polar overdominance (10, 14).
Preliminary mapping studies using
bovine DNA markers determined that
the CLPG locus is on ovine chromosome 18 (9). Subsequently, a linkage
group of 25 marker loci spanning
87.2 cM has been used to determine
that the CLPG gene is located within
the telomeric region of ovine chromosome 18 (14, 28). Currently,
researchers are attempting to determine the precise CLPG location and
sequence using positional candidate
cloning techniques (10).
There has been some discussion in
the literature as to the relationship of
the CLPG gene to other muscling
mutations. However, the inheritance
of the double muscle phenotype in
cattle is due to a partially recessive
base pair deletion or mutation (6).
The ryanodine receptor gene that is
responsible for extreme muscling and
leanness in pigs is located on porcine
chromosome 6 (35, 43) and bovine
chromosome 18 (6). Because of the
location of both of these mutations,
it seems very unlikely that they have
any role in the ovine muscle hyper-

69

REVIEW: Callipyge Gene in Sheep

TABLE 1. Performance traits of normal and callipyge rams, wethers, and

ewes.
Phenotype
Trait

Callipyge

Normal

Average daily gain, kg/d

Rams (n = 42)
Ewes (n = 54)
Wethers (n = 24)

0.36
0.26
0.29

0.35
0.29
0.31

0.378
0.802
0.184

Average daily feed intake, kg/d

Rams
Ewes
Wethers

1.75
1.48
1.67

1.90
1.68
2.11

0.016
0.004
0.001

Feed conversion, gain:feed

Rams
Ewes
Wethers

0.203
0.173
0.175

0.185
0.165
0.148

0.007
0.050
0.036

trophy model. Finally, the recently

described Carwell condition in
Australian Polled Dorsets (1), which
results in smaller increases in muscle
accretion when compared with CLPG
animals, has been localized to a
region near the CLPG locus on
chromosome 18. Although the
polar overdominance inheritance
model has been found to occur with
the Carwell condition, the extreme
muscling has not been seen. Therefore, this condition may be a different allelic form of the same gene,
which causes the CLPG phenotype
(37).

the effects of the CLPG mutation on

commercially important performance
parameters of 311 sheep. There were
no significant differences detected for
dystocia between CLPG and normal
ewes (P<0.10), most likely from the
delayed phenotypic expression of
CLPG phenotype. Furthermore, these
data indicate that normal and CLPG
animals are indistinguishable at birth
(18).
To evaluate growth parameters
thoroughly, 120 heterozygous normal
and CLPG lambs were fed identical
commercial NRC-balanced grower/
finishing diets for 120 d. Growth
rate was independent of the CLPG
phenotype (P>0.10), and sex did not
significantly interact with the CLPG
First, it appears that the CLPG
phenotype (P>0.10; Table 1). Howgene results in a phenotype that is
ever, CLPG animals were found to
very different from other muscle
have superior muscling (P<0 .05);
hypertrophy mutations as evidenced
lower average daily intakes (P<0.02);
by the lack of dystocia or reduced
and, consequently, improved feed to
growth rate that is associated with
gain ratios (P<0.05) (17). The authors
other muscling mutations (3, 33, 44,
suggest that improved feed efficiency
46). Second, the CLPG phenotype is
of CLPG lambs might be a result of
not expressed until 4 to 6 wk postna- increased nitrogen retention as
tally. Current research indicates that measured in a 14-d feeding trial. In
sheep expressing the CLPG phenotype this study, lambs were fed diets
do not have the negative characteris- containing 12 and 18% CP. Callipyge
tics associated with double muscling
lambs were found to be more effiin other agricultural species (19). In
cient than controls on the 18% CP
fact, Jackson et al. (18) conducted a
diet and utilized a higher percentage
series of investigations to determine
of their dietary nitrogen than the

Animal Performance

normal lambs (16). Fleece weights

and staple lengths of the CLPG lambs
have been found to be 12.7% lower
(P<0.03) and 8.7% shorter (P<0.05),
respectively, than normal lambs (18).
Overall, it seems that the CLPG
mutation improves growth efficiency
without causing dystocia. Unfortunately, the increased feed efficiency
and growth rate translate into a
decrease in fleece weight.

Muscle Characteristics
Early researchers suggested that
the CLPG phenotype is a hypertrophy condition as evidenced by the
lack of dystocia and alterations in
muscle fiber type (18, 25). Second,
hyperplastic growth would have most
likely resulted in larger birth weights
when compared with normal lambs.
However, many researchers have
shown that CLPG and normal lambs
are indistinguishable at birth (9, 10,
18). Histological examination of
CLPG phenotypes revealed a higher
percentage of fast twitch glycolytic
muscle fibers (P<0.001) and a lower
percentage of slow twitch oxidative
and slow twitch glycolytic muscle
fiber (5, 25). In fact, the overall
average cross-sectional muscle fiber
area increased as much as 62% in
CLPG animals (25).
Further support for the hypertrophy model arises from studies revealing that there is an increase in RNA
concentration, DNA content, and
DNA:RNA ratios of CLPG animals
(P<0.05) (25). Percentage of protein
degradation was lowered in CLPG
animals as indicated by an increase
in the activity of the protease inhibitor calpastatin (P<0.05) (29). In fact,
the fractional protein degradation
rate for CLPG phenotypes is significantly decreased when compared
with normal controls. Normal lambs
had 2.3 g/d less protein accretion
than CLPG animals and 6.9 g/d more
fat accretion (14). Total body fat and
lean for normal carcasses were 31.5
and 64.0%, respectively, whereas the
CLPG animals had 24.3% fat and
71.3% lean (14). Overall, the large
increase in muscling in CLPG lambs

70

Jackson and Blanton, Jr.

may result from an increase in

satellite cell proliferation and incorporation combined with a decrease in
muscle protein degradation (41).
Therefore, the expression of the CLPG
phenotype at 4 wk of age results in
larger muscles (hypertrophy) and not
an increase in muscle fiber numbers
(hyperplasia).

Blood Profiles
A 1998 study was designed to
evaluate typical growth-promoting
serum hormone profiles of both
CLPG and normal lambs. Differences
did not exist in serum growth hormone concentration, growth hormone pulse amplitude, or pulse
frequency (P>0.3). Furthermore,
there were no differences in IGF-1,
insulin, or thyroxine concentrations
(45). In the investigation of stress
hormones, CLPG animals placed
under restraint stress had the same
serum cortisol levels as normal lambs
placed under similar restraint (45).
Therefore, these researchers concluded that the excessive muscling of
the CLPG animals is not a result of or
caused by any differences in systemic
hormone concentrations (45).
In an attempt to identify CLPG
lambs prior to phenotypic expression,
serum creatine and lipid concentrations were selected as marker compounds (30). Serum lipid concentrations were indistinguishable between
CLPG and normal lambs at 1 mo of
age. However, by 2-mo-old normal
lambs had lower concentrations of
triacylglycerol, cholesterol, high
density lipoproteins, and very low
density lipoproteins. These results are
understandable when considering
that the CLPG phenotype is not
expressed until 4 to 6 wk of age.
Surprisingly, by slaughter age, animals did not have any differences in
serum lipid profiles. Although these
data suggest that serum lipids could
be used to identify CLPG animals,
these differences are not evident until
phenotypic expression. Therefore,
visible identification of CLPG animals is probably more reliable,

Figure 1. Rambouillet (A,B) and Hampshire (C,D) ewes expressing callipyge (A,C) and normal
(B,D) phenotypes.

controls of the same live weight.

Lambs expressing the CLPG gene had
a higher (40.2 vs 32.9) percentage of
boneless retail yield; higher protein
(16.6 vs 15.2), moisture (63.6 vs 58.6),
It is important to examine the
and ash (0.85 vs .077) percentages,
effects of the CLPG phenotype on
and a lower fat (18.9 vs 25.4) percentcarcass quality to ensure that these
age than normal controls (18).
lamb products meet consumer accep- Following complete dissection, the
tance parameters. To determine
CLPG carcasses yielded 5% less total
carcass characteristics of the CLPG
body fat, 2% less fat trim, 1% less
phenotype, carcass dissections were
seam fat, 3% less bone (18), and
performed (18) on normal and CLPG significantly lighter pelts (15) than
lambs at 54.5 kg live weight. There
normal controls.
were no significant differences in any
These results were confirmed by
internal organ weights, with the
Koohmaraie et al. (25), who deterexception of the large intestine,
mined that the CLPG phenotype did
which was larger in the normal
not affect slaughter weight. However,
animals (P<0.05) (15, 18). In addiCLPG lambs had higher dressing
tion, no differences were seen in any percentages and reduced fat thickness
carcass linear measurements includand marbling scores (P<0.05).
ing height, hindsaddle, or carcass
Muscles of CLPG lamb hindquarters,
length. Following the dissection and including the adductor (30%), biceps
weighing of 19 economically imporfemoris (42%), gluteus group (31%),
tant muscle groups, it was determined longissimus (18%), semimembranosus
that the CLPG lambs had 30% more
(38%), and semitendinosus (26%),
overall muscle weight than normal
were significantly heavier in the
controls (Figure 2). In fact, CLPG
CLPG phenotypes (P<0.01). Most of
lambs had a 5% increase in dressing
the increase in muscle mass of CLPG
percentage, 70% larger loin eye area,
lambs appears to be concentrated in
and higher leg and conformation
the area of the hind leg and loin
scores when compared with normal
(Table 2) (19). Muscles excised from

accurate, and expedient than serum

lipid profiles (Figure 1).

Carcass Characteristics

71

REVIEW: Callipyge Gene in Sheep

TABLE 2. Mean muscle weights from Rambouillet rams of both

phenotypesa.
Phenotype
Muscle

Callipyge

Superficial gluteal
Tensor facia latae
Gluteus medius
Gracilis
Semitendinosus
Adductor
Semimembranosus
Rectus femoris
Vastus group
Peronius tertius
All excised pelvic muscles

575.1
94.1
396.5
85.7
169.3
233.3
526.7
214.1
385.0
56.9
2,736.7

Normal

360.2
76.9
265.4
66.5
133.3
158.2
323.1
173.6
321.7
50.8
1,929.7

0.001
0.002
0.001
0.001
0.001
0.001
0.001
0.01
0.002
0.06
0.001

(g)

an

= 9 carcasses of each phenotype.

the hind legs of the CLPG lambs were

nearly 40% heavier than the same
muscles from normal lambs. Longissimus muscle from CLPG lambs were
45% larger; however, there was only a
13% increase in total muscle weight
when compared with normal lambs.
The importance of this unique
muscle accretion pattern becomes
very important when considering
that these animals had similar live
weights (19). Together these data
indicate that CLPG animals are
considerably more efficient at producing muscle.

Meat Characteristics
Overall palatability of muscle is
determined by the tenderness, juiciness, and flavor of the product.
These values are measured using
subjective evaluations of trained
sensory panels or objective evaluations with mechanical equipment
such as the Warner-Bratlzer shear
(WBS) force machine, which applies
a measured amount of force to a
cooked core of muscle sample. Utilizing these techniques, CLPG muscle
has been examined to determine
whether the product is acceptable to
todays lamb consumer.

collagen percentages (P<0.05) were

lower in CLPG lambs (13), which
indicates that collagen cross-linking
is not the cause for CLPG longissimus
toughness.
Higher levels of calpastatin, a
protease inhibitor, have been found
in CLPG lambs muscle products (20,
22, 25). As a consequence of the
increased calpastatin activity, postmortem protein degradation is
slowed, which may increase toughness. In fact, calpastatin activity is
higher in CLPG animals until 56 d
post-mortem when compared with
normal controls (P<0.05) (25, 29).
Interestingly, -calpain activity did
not differ between normal and CLPG
animals at 0 and 56 d post-mortem
but was higher at day 14. However,
m-calpain levels were higher at all
time points for CLPG carcasses.
Consistent with calpastatin activity
studies, Western blot analysis of
muscle structural proteins (e.g.,
dystrophin, titin, desmin) revealed
that postmortem proteolysis was
greater in normal lambs when
compared with CLPG animals. In a
study utilizing a transmission electron microscope, sarcolemma detach-

Researchers have determined that

some muscles from CLPG lambs are
less tender than those from normal
control lambs (13, 18, 23, 24, 31). In
fact, the longissimus muscle of CLPG
lambs has always been found to be
less tender when compared with
normal lambs, with less tenderness
variation being detected in other
hindquarter muscles (12, 31, 40).
Warner-Bratzler shear force values for
longissimus muscle at 1, 7, and 21 d
post-mortem in CLPG lambs were
found to be significantly higher
(P<0.01) than normal lambs (13, 18,
24). Utilization of tenderization
methods such as electrical stimulation did not eliminate the tenderness
problems of the longissimus muscle
when compared with normal controls (21). One hypothesis for the
increased toughness of the longissimus muscle was an increase in
connective tissue, or more specifically,
collagen cross-linking in the CLPG
muscle. A representative component
of connective tissue is hydroxylysylpyridinoline, a trivalent collagen
cross-linker that has been found to
increase with age and toughness (32).
However, researchers revealed that the
levels of hydroxylysylpyrodinoline
Figure 2. Normal (a) and callipyge (B)
cross-link percentages (P<0.04) and
Rambouillet carcasses.

72

ment was delayed in CLPG animals

when compared with normal lambs.
This study also revealed that 15% of
I-bands were fractured by 14 d postmortem in normal lambs, but only
3% fracture was detected for CLPG
animals at the same post-mortem
storage (42). Therefore, a possible
reason for the decreased tenderness of
the longissimus muscle from CLPG
lambs is the stability of muscle
myofibrils. Overall, the increased
calpastatin activity of CLPG animals
may play a significant role in postmortem tenderization of CLPG meat.
However, it is important to remember
that as calpastatin increased in the
CLPG lambs, some increase in calpain and m-calpain was seen.
These increases may offset or negate
the increased calpastatin activity
post-mortem.
Because of tenderness problems
associated with CLPG longissimus
muscle, many researchers have
studied technologies to improve the
tenderness problems associated with
the CLPG phenotype. One of the
earliest approaches explored was to
extend post-mortem aging. Aging
CLPG longissimuss muscle for 14 to
21 d resulted in decreased WBS force
values by 12 to 25% (2, 25). Unfortunately, these aging periods were
unable to create muscle tenderness
similar to the normal lambs.
Muscle tenderness should be
improved if sarcomere shortening can
be prevented during the early postmortem period. Therefore, methods
of rapid freezing to reduce sarcomere
shortening were investigated as
methods to prevent CLPG tenderness
problems. Submersion of muscle
samples in liquid nitrogen resulted in
a decrease in longissimuss sarcomere
length shortening from 1.33 to 1.99
m. Although the reduction in
sarcomere shortening decreased WBS
force values when compared with
controls, the CLPG longissimus
muscle WBS force was still 63%
higher than normal lambs at 14 d
post-mortem (24). In a similar study,
researchers determined that freezing
CLPG longissimus muscle in liquid
nitrogen reduced calpastatin activity

Jackson and Blanton, Jr.

to the levels of normal control

animals, which should increase the
postmortem proteolysis of the CLPG
longissimus muscle. Unfortunately,
trained sensory panel tenderness
scores were still higher for CLPG
muscle (P<0.05) aged after freezing
(12).
Electrical stimulation was found to
increase tenderness of CLPG longissimus muscle by 30 to 34% (P<0.05),
but had no effect on consumer
acceptance (21). Calcium chloride
(CaCl2) injections or calcium-activated tenderization has been utilized
to increase the rate of post-mortem
proteolysis and to improve muscle
tenderness in several animal species
(26). Researchers found that 5% (wt/
wt) injection of 200 mM CaCl2 will
improve overall tenderness of CLPG
meat (25, 27). Calcium chloride
injections resulted in WBS force
values that are up to 100% lower
than control CLPG, with final WBS
force values being similar to control
animals (20, 24, 27). Interestingly, by
combining liquid nitrogen freezing,
calcium injection, and 14-d aging,
CLPG lamb carcasses have tenderness
values virtually indistinguishable
from normal control animals.
However, a very rapid and often
overlooked method for tenderization
of CLPG longissimus muscle is the
hydrodyne process (40). This process
generates a sonic shock wave that
reflects off particles in the muscle
that are an acoustic match. This
reflection of the sonic wave acts as a
mechanical disruption of muscle
proteins (40). When the hydrodyne
process was applied to CLPG muscle,
it instantaneously produced meat
with WBS shear force values similar
to or better than normal controls.

Flavor and Consumer

Acceptance
Palatability of CLPG muscle is
critical to consumer acceptance and
appears to be influenced by many
factors including age, diet, genetics,
and cooking method. Robinson et
al. (36) reported that consumers

prefer the larger, leaner loin associated with the CLPG phenotype.
However, when Moore et al. (31)
compared CLPG and normal lamb
chops, they determined that consumers preferred normal lambs because of
the decreased tenderness associated
with the CLPG lambs. In this study,
CLPG meat was rated as too lean by
18% of consumers; 23% of consumers
felt that the normal lambs were too
fat. Freezing CLPG chops prior to the
aging period seems to have alleviated
the tenderness problem for some
consumers (11). Interestingly, sensory evaluations of leg roasts and
bone-in square-cut shoulder roasts
did not differ between CLPG and
normal lambs (20, 34). Overall,
sensory studies have found that
CLPG lamb flavor is not significantly
different for normal lambs (18).
Juiciness, tenderness, and overall
acceptability are lower for CLPG loin
chops and shoulders when compared
with normal lambs; however, leg
chops or roasts were not significantly
different for these same sensory
characteristics (20).

Implications
The CLPG gene in sheep offers
tremendous potential for improving
the growth efficiency and composition of lamb carcasses. It has been
well documented that CLPG lambs
have more muscle mass, improved
muscle:fat ratios, and higher dressing
percentages. Productivity of CLPG
ewes does not seem to be any different than the productivity of normal
ewes. Reproductive parameters of the
males are unaffected by the CLPG
phenotype (18). Tenderness of CLPG
longissimus muscle is the only trait
that seems to be affected negatively
by the CLPG gene, and palatability
and juiciness are relatively unaffected. Techniques such as the
hydrodyne system or extended aging
and freezing may alleviate the tenderness problems of CLPG longissimus
muscle. Therefore, through proper
meat processing, the CLPG gene
could have a profound impact on the
profitability of lamb production.

REVIEW: Callipyge Gene in Sheep

13. Field, R. A., R. J. McCormick, D. R. Brown,

F. C. Hinds, and G. D. Snowder. 1996.
Collagen crosslinks in longissimus muscle
from lambs expressing the callipyge gene. J.
Anim. Sci. 74:2943.

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