009-2007


   2550;5(1):81-91.

Review article

ANIMAL BIOTECHNOLOGY: PIG MUSCLE PART I, FACTORS

AFFECTING MUSCLE FIBER TYPES IN PIGS

Nguyen Trong Ngu,1 Korakot Nganvongpanit2

Department of Agricultural Genetics and Breeding, College of Agriculture and

1

Applied Biology, Cantho University, Cantho, Vietnam,

2
Department of Veterinary Preclinical Science,
Faculty of Veterinary Medicine, Chiang Mai University, Chiang Mai, Thailand

Abstract Today, the study of muscle fibers is critical because they are responsible for the
variation of growth performance and meat quality traits in farm animals. There are several
factors that can contribute to alter the muscle fiber composition. Some of them geneti-
cally originate from individual, breed, gender, birth&slaughter weight and genetics,
while others are environmental factors such as prenatal and postnatal nutrition, tempera-
ture and physical activity. Chiang Mai Veterinary Journal 2007;5(1):81-91.
Keywords: pig, muscle fiber type, factor

The mammalian skeletal muscle is com-
posed of a heterogeneous collection of fiber
types with polynucleated and elongated cells
that can be classified according to their
myosin heavy chain (MyHC) isoforms, contrac-
tile elements (microfilaments), energy metabo-
lism, fiber color or cross-sectional area (CSA)
as showed in table 1.(1-3) In farm animals, the
structural and functional diversity of skeletal
muscle is represented by the variety of myosin
isoforms. Understanding factors influencing
muscle fibers will help to optimize the efficiency
of the muscle growth and meat quality, which
are of important concerns in animal production.
In this review, the effects of both genetics and
environments on the distribution of muscle
fibers will be discussed with the emphasis on
pigs.
Genetic factors
Muscle and individual
The composition of fiber types varies be-
tween anatomical muscles (Fig. 1). The distri-
bution of pig muscles is unique and highly

Address request for reprints: Korakot Nganvongpanit, Department of Veterinary Preclinical Science, Faculty of
Veterinary medicine, Chiang Mai university, Chiang Mai 50100, Thailand; E-mail:korakot@chiangmai.ac.th
Article received date: August 21st, 2006
82
Figure 1. Percentage of muscle fiber types in different muscle areas of the pig (6,10,11)
organized, in which deep muscles contain more
type I fibers surrounded by an internal rosette
of type IIa fibers and an external ring of type IIb
fibers. (4)
Functionally, type I fibers are more
involved in maintaining posture, while type IIb
fibers are responsible for rapid movement.
commercial breeds, the proportions of type IIb
and IIx fibers of longissimus dorsi and psoas
muscle are highest compared with the other
fibers as pointed out by many research
groups. (6-10)
The same tendency was also
noticed in biceps femoris, quadriceps femoris
and adductor. (11)
On the other hand, examples
Nguyen Trong Ngu, Nganvongpanit K.
of high degree of type I and IIa fibers can be
seen in rhomboideus muscle.(9,10)
Animals of the same breed reared in the
same environment also show large variation in
fiber type composition. An example is the varia-
(5)
In tion within litter of Danish Landrace x Large
White (LW) pigs, as described by Nissen et al.
(2004).(12) In this experiment, they slaughtered
and grouped the animals by litter, at the same
age, according to the body weight: heaviest,
middle and lightest weight. It was found that,
(6)
intra-litter growth performance varied largely as
a result of differences in both the number and
Animal biotechnology: pig muscle
growth rate of muscle fibers followed the trend
that heavy pigs had higher total number of fiber
(TNF) than middle and lightest weight pigs,
which did not differ from each other.
Breed
Breed covers probably the main proportion
of genetic factors affecting the muscle fiber
composition of a certain muscle. The compo-
sition of muscle fiber types in different breeds
is listed in table 2. Generally, the proportion of
type IIb fibers is dominant in most modern and
traditional breeds. However, a dramatic change
in the relative expression profile was found in
our study. (13)
Mongcai is a Vietnamese local
breed and known for its highly preferred meat
quality but unsatisfactory muscularity. Larger
loin eye muscle area offered higher proportion
of type IIb fibers(13) and this helps to explain the
difference of fiber distribution in Mongcai
compared to other breeds. Moreover, in com-
parison between the numbers and types of
muscle fibers in large and small pig breeds,
authors(14) announced that the responsible
reasons for muscle size variation between LW
and miniature pigs were due to the difference in
myofiber number, a factor fixed before postna-
tal growth. (15)
In addition, it appeared that pigs
exhibiting postnatal increases in myofiber size
are more related to age than to live weight. The
mechanism to which fewer muscle development
in genetically small animals is different from
that exhibited by nutritional deprivation animals
in utero (14)
and consequences of these differ-
ences are therefore reflected in chemical prop-
erties of the constituent muscle fibers. In com-
parison between miniature and LW pigs, a
higher content of type I fibers in the latter breed,
83
which may support their increased weight and
accord with the hypothesis that at similar stage
of growth, there are great differences across
breeds and fiber sizes. Finally, muscle of wild
pigs was reported to contain higher area
percentage of type IIa and conversely lower
percentage area of type IIb fibers than those
from the same muscle of domestic pigs.(16)
Gender
There have been contradictory documenta-
tions on the distinction between females and
males in term of proportion of muscle fibers. In
general, females have larger fibers than
castrated males and as a result, sex has been
mentioned to affect on the cross-sectional area
but not on the proportion of each fiber.(19-22)
Moreover, Lefaucheur et al(1998)(23) demon-
strated a significant decrease of relative fiber
area of type I fiber in females compared to
intact LW males, which may indicate that
castration decreases relative area of type I
fibers. Nevertheless, Rehfeldt et al(2000)(24)
evidenced smaller values of type IIa and IIb
fibers in boars in comparison to female pigs
and this implies a higher numbers of muscle
fiber in male pigs because the weight of the
longissimus dorsi muscle was similar.
Birth weight
The effect of birth weight on muscle fiber
performances has been mentioned in many
publications. Indeed, a tendency that lower
total fiber number in piglets is associated with
low birth weight was revealed.(10,25,26) Most of
the variations were due to a difference in the
number of secondary myofibers that formed
around each primary. However, results from
84 Nguyen Trong Ngu, Nganvongpanit K.

other studies did not observe any association
between birth weight and TNF and thereby
rejected this suggestion. (27,28)
Although birth
weight was indicated to have an association
with enlarged muscle fiber area,(29) its influence
on muscle fiber composition of longissimus
dorsi and rhomboideus muscle was not estab-
lished.(10,22) Similarly, Rehfeldt et al(2006)(30)
Animal biotechnology: pig muscle
stated that the ranking of fiber number at slaugh-
ter was almost the same as at birth, with low
fiber numbers in low birth weight and high
numbers in high birth weight piglets. Because
no differences were observed in the frequencies
of different fiber types, Rehfeldt and Kuhn (2006)
(30)
concluded that postnatal fiber differentiation
is independent of birth weight.
Age and slaughter weight
At birth, muscle fibers are oxidative and the
relative number of slow-twitch numbers contin-
ues to increase until 8 weeks after. (4)
The
proportions of white fibers were observed to
intensively increase up to 4 months of age,
together with the rapidly increased size and
continue at a slower rate afterward. (31)
Surpris-
ingly, although type IIa fibers are a minor in
porcine skeletal muscle but their relative fiber
type-restricted expression was highest among
4 isoforms at both stages 6 weeks and 22 weeks
postnatal. (32)
A relation of age and muscle
fibers was additionally recorded in such a way
that increasing weight and age at slaughter did
change the cross-sectional area but not the
numerical percentage of any fibers. (33,34)
Candidate genes
Muscle fiber formation occurs during embry-
onic development including two stages of
primary and secondary generation. (35)
During
this period, the meiogenesis is under control of
the MyoD gene family consisting of four struc-
tural related genes MYOD1 (MYF3), myogenin
(MYOG or MYF4), MYF5 and MRF4 (MYF6). (36,37)
The MYOD1 and MYF5 genes are involved in
myoblasts proliferation and they were found to
directly affect the proportion of fast-twitch
85
oxidative fibers and the fast-twitch low-oxida-
tive fibers of pigs being crosses of Pietrain x
(Polish LW x Polish Landrace) and thereby also
influence the metabolic properties of muscle.(38)
The other gene, myogenin, is expressed in all
myoblasts starts from differentiation to cell
fusion and also marks the end of the myoblast
proliferation. In deed, a significant difference of
two homozygous genotypes for birth weight,
growth rate and lean weight was reported by te
Pas et al. (1999).(39) The expression of the last
gene in the MyoD family, MYF6, also involves
in the differentiation and maturation of myotubes
and highly expressed postnatally(40) but experi-
mental outcome demonstrating its effects on
muscle fibers are still scare.
It is well known that pigs homozygous for
the halothane (HAL) sensitivity allele (nn) are
highly stress susceptible and often induces
accelerated pH-ultimate (pHu) decline post-
mortem (p.m.) leading to a pale, soft and
exudative (PSE) meat. Different halothane
genotypes have been described to associate
with the content of muscle fibers and thereby
affect on meat quality. For instance, Depreux
et al. (2002)(41) showed a greater amounts of
type IIb but less slow fibers in pigs carrying the
“n” gene (Nn or nn), whereas the NN pigs
exhibited higher proportion of type IIax fibers.
Moreover, a positive correlation between the
relative abundance of type IIb fibers and pH
was observed in pigs free of “n” gene, but across
all genotypes, the relationship between type IIb
fibers and pH45 was negative. This was curious
because, it is opposite with the whole assump-
tion that type IIb isoform hydrolyzes ATP
rapidly and increases the glycolysis rate.(41) In
the other hand, it was suggested that the rela-
86
tive amounts of individual fibers are not attrib-
uted by the effects of HAL, or in other words,
the process of maturation from one fiber to an-
other is free from the HAL accelerated effect.
Conversely but similar detrimental effect on
meat quality, the RN gene mainly increase the
CSA of red fibers leading to a decrease in rela-
tive area of white muscle (IIx and IIb) and thus
the fibers are more to oxidative and less to
glycolytic metabolism. The RN effect, namely,
“acid meat” phenotype indicated a positive
correlation between glycolytic potential and lac-
tate content and pHu. (43)
In fact, results from
Marinova et al(1992) (44)
proved that pigs
carrying RN gene have higher glycogen con-
tent in white muscles especially in glycolytic
fibers. This was later confirmed by Lebret et
al(1999), (45)
who concluded that white muscles
are more affected than red muscles (I and IIa)
and that the glycogen content increase in long-
issimus muscle. Further findings from these
authors also emphasized a higher enzyme
activities and relative area of type II red fibers in
the dominant RN carriers. A causative muta-
-
tion (R200Q) for the RN gene in the PRKAG3
-
gene encoding for a muscle-specific isoform of
the regulatory(³) subunit of adenosine mono-
phosphat-acitvated protein kinase additionally
reported. (46)
Calpastatin (CAST) is a specific inhibitor of
calpain, a Ca -activated protease family,
2+
considered to be involved in the initiation of myo-
fibrillar protein degradation in living muscle.
In a preliminary study, Klosowska et al(2005)
showed that longissimus lumborum diameters
of all types of muscle fibers are significantly
related to the Stamboek pigs’ genotype at
locus CAST. The percentage of fast-twitch
Nguyen Trong Ngu, Nganvongpanit K.
glycolytic fibers in a bundle was also concluded
to change the metabolic properties of muscles
and thereby meat quality. Although this locus
(42)
is located in the 6th intron of the gene, this
intronic mutation should be considered both as
a marker for muscle microstructure character-
istics and as the causal mutation itself.(48)
Environmental factors
Prenatal feeding
During gestation, prenatal muscle develop-
ment includes two successive generations and
thus produces primary fibers (up to 50-55 days)
and secondary fibers (up to 90 days). In most
cases, the nutritional manipulation has little
effect on the early period of myogensis, a stage
involves in differentiation of primary muscle
fibers. However, nutrition may change the
number of primary fiber differentiation possibly
because of indirect influence on the placenta
development.(30,49) In pigs, between 25 to 90 days
of gestation, the differentiation and hyperplasia
of secondary fibers have been demonstrated
as a cause for under-nutrition, which can lead
to runting, a decrease in muscle fiber numbers,
especially secondary fibers.(25,27,50) Likewise,
findings on the relationship between over-nutri-
tion and muscle fiber characteristics are still
unclear. Over-nutrition of the sow between 25
to 50 days of gestation might increase the TNF
in developing pigs(51) whereas increased mater-
nal nutrition of sows from day 25 to 50, or 25 to
(47)
70 of gestation did not give any benefits on
(48)
muscle fiber number and area in the offspring.(52)
Postnatal feeding
There has been much attention on the role
of nutrition on muscle development. Under-
Animal biotechnology: pig muscle
nutrition was demonstrated to account for a
reduction of cross-sectional area of future fast-
twitch glycolytic fibers in longissimus dorsi
muscle. (53)
Chilibeck et al(2005) (54)
carried out
an experiment on rapidly growing gilts, in which
limited overfeeding at 75% more energy than
needed for weight maintenance were offered at
two stages, from days 1-7 (early luteal phase)
and from day 8-15 (late luteal phase) of the
estrous cycle. Results showed that muscle
fiber area and fiber type composition were
independent on restricted overfeeding at any
time exclusive of a significant decrease in type
IIa fiber percentage over time. Partly opposite
results were presented by Chilibeck et al
(2005),(55) who failed to evaluate any effect of
restricted feeding (approximatly 30%, started
from 30 to 100 kg) on fiber number, CSA diam-
eter and relative area of fast-twitch oxidative,
fast-twitch glycolytic fibers. However, a signifi-
cant difference was noted in case of slow-twitch
oxidative fibers, in which restricted pigs had
larger cross-sectional area, diameter and rela-
tive area than those from ad libitum fed pigs.
Supportably, an association between postna-
tal nutritional status and type I fiber expression
both at mRNA and protein levels was unrav-
eled. (56)
Also, restricted feeding (50% of ad
libitum) at an early stage (between 3 and 7
weeks of age) had no influence in myofiber type
composition in longissimus dorsi but led to a
remarked increase in type I fibers proportion in
the red rhomboideus muscle.(57) An assump-
tion for these findings is, because the energy
usage per unit tension is lower in type I fibers,
as a result, a selective increase in type I pro-
portion in muscle during the period of reduced
available energy would be physiologically
87
relevant to spare energy.(23,57)
Other factors
Other factors that may have an impact on
muscle fiber characteristics include physical
activity, ambient temperature and growth-pro-
moting agents. In fact, climate conditions and
physical exercise can influence on pig perfor-
mance raised in an outdoor production system.
Animals born outdoor at low temperature had a
higher percentage of type I, but lower percent-
age of type IIa fibers in the longissimus muscle.
Interestingly, this difference changes in pigs
finished outdoor environment as Harrison et al.
(1996)(58) indicated a higher proportion of type
IIa fibers in both longissimus and semimem-
branosus muscle, and vice versa for type IIb/IIx
fibers. An increase of type I fiber percentage in
pigs long-term exposing to cold temperature is
generally accompanied by an increase in
oxidative metabolism.(59) Moreover, outdoor
pigs have more spontaneous activities leading
to a shift of muscle fibers from type IIb to IIx to
IIa and to I, respectively, which can explain the
more type IIa and less type IIb/IIx in muscle
compared with indoor pigs.(23) In addition to
some environmental factors, growth promoters
such as growth hormone, b-agonists and
steroids can influence muscle fiber composi-
tion of farm animals. An excellent review
regarding to these promoting agents is avail-
able.(60)
Conclusion
It is clear that the distribution of skeletal
muscle fibers is heterogeneous among individu-
als. However, conclusions whether these
differences are genetically determined or
88 Nguyen Trong Ngu, Nganvongpanit K.

consequence of environmental influences are 8. Chang KC, da Costa N, Blackley R, South-

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