Meat Science 95 (2013) 828–836

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Meat Science
journal homepage: www.elsevier.com/locate/meatsci

Review

Control of fresh meat quality through manipulation of muscle

fiber characteristics
S.T. Joo a,⁎, 1, G.D. Kim b, 1, Y.H. Hwang a, Y.C. Ryu b
a
Department of Animal Science, Institute of Agriculture & Life Science, Gyeongsang National University, Jinju, Gyeongnam 660-701, South Korea
b
Division of Biotechnology, College of Applied Life Sciences, Jeju National University, 102 Jejudaehakro, Jeju 690-756, South Korea

a r t i c l e i n f o a b s t r a c t

Article history: Variations of fresh meat quality exist because the quality traits are affected by various intrinsic and extrinsic
Received 8 February 2013 factors. Because the meat quality is basically dependent on muscle fiber characteristics, numerous studies
Received in revised form 14 April 2013 have reported the relationship between quality traits and fiber characteristics. Despite intensive research,
Accepted 15 April 2013
the relationship is yet to be fully established, however, the present knowledge suggests several potential
ways to manipulate muscle fiber characteristics to improve meat quality. The present paper reviews the
Keywords:
Meat quality traits
definition of fresh meat quality, meat quality traits and variations of meat quality. Also, this review presents
Muscle fiber characteristics recent knowledge underlying the relationship between fresh meat quality traits and muscle fiber character-
Muscle fiber types istics. Finally, the present work proposes several potential factors including breed, genotype, sex, hormone,
growth performance, diet, muscle location, exercise and ambient temperature that can be used to manipulate
muscle fiber characteristics and subsequently meat quality in animals.
© 2013 The Authors. Published by Elsevier Ltd. Open access under CC BY-NC-SA license.

1. Introduction (CSAF) are major determinant factors of muscle mass as well as

Meat quality has always been important to the consumer, and it is
an especially critical issue for the meat industry in the 21st century.
As consumer demand for high quality meat is increasing in most
countries, the meat industry should consistently produce and supply
quality meat that is tasty, safe and healthy for the consumer to ensure
continued consumption of meat products. In order to produce high
quality meat, it is necessary to understand the characteristics of
meat quality traits and factors to control them.
Fresh meat quality is difficult to define because it is a complex
concept determined by consumer preferences. Because fresh meat is
animal tissue that is suitable for use as food, the quality characteris-
tics are influenced by various factors such as muscle structure, chem-
ical composition, chemical environment, interaction of chemical
constituents, postmortem (p.m.) changes in muscle tissues, stress
and pre-slaughter effects, product handling, processing and storage,
microbiological numbers and populations, etc. In particular, fresh
meat quality is directly related to muscle fiber characteristics because
skeletal muscles mainly consist of muscle fibers. The muscle fibers are
characterized by their morphological traits, and contractile and
metabolic properties (Lee, Joo, & Ryu, 2010). Morphology traits such
as total number of fibers (TNF) and cross-sectional area of fibers
meat quality. Also, contractile and metabolic properties of muscle
are differentiated by muscle fiber types, and thus fresh meat quality
is strongly related to fiber type composition (FTC) in muscle.
In general, there are four different muscle fiber types in adult skeletal
muscle, which are slow-oxidative or type I, fast oxido-glycolytic or type
IIA, and fast glycolytic IIX and IIB (Schiaffino & Reggiani, 1996). All of
these fiber types are observed in most muscles of meat animals, and
their relative composition in the different muscles can determine the
predominance of muscle's metabolic properties (Ozawa et al., 2000;
Ryu & Kim, 2005). Consequently, p.m. muscle metabolism which is a
crucial factor to determine fresh meat quality is affected by TNF, CSAF
and FTC (Kim, Jeong, et al., 2013; Ryu, Lee, Lee, & Kim, 2006). These mus-
cle fiber characteristics vary by various factors including breed (Ryu et
al., 2008), selection (Larzul et al., 1999), gender (Ozawa et al., 2000),
hormone (Rehfeldt, Fiedler, & Stickland, 2004), growth performance
(Gondret, Lefaucheur, Juin, Louveau, & Lebret, 2006; Kim, Kim, et al.,
2013), diet (Jeong et al., 2012) and muscle location (Beermann et al.,
1990; Hwang, Kim, Jeong, Hur, & Joo, 2010). Therefore, understanding
the relationship between muscle fiber characteristics and meat quality
traits will improve the production of quality meat, and manipulation
of muscle fiber characteristics would have profound impacts on the
profitability of the meat industry. The present paper reviews the scien-
tific literature in meat quality traits, muscle fiber characteristics and
potential factors to manipulate muscle fiber characteristics.

2. Fresh meat quality

⁎ Corresponding author. Tel.: +82 55 772 1943; fax: +82 55 772 1949.
E-mail address: stjoo@gnu.ac.kr (S.T. Joo). The term ‘fresh meat quality’ is very ambiguous because its defini-
1
These authors contributed equally to this work. tion varies depending on the background of consumers in different

0309-1740 © 2013 The Authors. Published by Elsevier Ltd. Open access under CC BY-NC-SA license.
http://dx.doi.org/10.1016/j.meatsci.2013.04.044
 S.T. Joo et al. / Meat Science 95 (2013) 828–836
regions of the world. Accordingly, first of all, meat quality should be
defined by most consumer preferences. Consumer preferences are re-
lated directly to the human senses such as appearance, smell, taste
and mouthfeel. Also, fresh meat quality can be defined by scientific
factors including composition, nutrients, colorants, water-holding
capacity (WHC), tenderness, functionality, flavors, spoilage, contami-
nation, etc.
The quality of fresh meat indicates its usefulness to the consumer
and its acceptability for cooking. The important quality traits for fresh
meat are color, WHC, texture and amount of fat (intramuscular fat/
intermuscular fat/subcutaneous fat), while the important traits for
eating quality of cooked meat are tenderness, flavor and juiciness. In
general, consumers rate color as the most important quality trait for
fresh meat, while tenderness is rated as the most important palatabil-
ity trait for cooked meat followed by flavor and juiciness (Glitsh,
2000). However, this can vary among consumers depending upon
past experiences and cultural background. Therefore, the order of
importance of meat quality traits can vary by country (Warner,
Greenwood, Pethick, & Ferguson, 2010).
The appearance of meat is determined by meat color, packaged
meat color, amount and distribution of fat, fat color, amount of drip
on the surface of the meat, purge in the tray, and texture of the
meat (Becker, 2000). These appearance quality traits (AQT) strongly
influence the consumer's decision to select good quality meat at the
point of purchase. However, the consumer determines the actual
meat quality at the point of consumption with eating quality traits
(EQT) such as tenderness, flavor, juiciness and succulence (Acebron
& Dopico, 2000). Additionally, consumers assess meat quality by reli-
ance quality traits (RQT) such as safety, nutrition, animal welfare,
ethics, price, product presentation, origin, and brand of meat products
(Troy & Kerry, 2010). Therefore, it is appropriate to define the term
‘fresh meat quality’ by consumer preferences that are determined
by RQT as well as AQT and EQT of meat (Joo & Kim, 2011).
3. Meat quality traits
Quality traits of fresh meat are categorized based on major intrin-
sic and extrinsic factors. Generally, intrinsic factors are the physiolog-
ical characteristics of meat such as AQT and EQT, whereas extrinsic
factors are the RQT of meat products (Joo & Kim, 2011). All these
traits contribute to the consumer's expectation of high quality meat.
Consumers determine quality meat as one with desirable color, firm
texture, less drip, high marbling, and moderate visible fat and fresh
meat odor, while discoloration, soft texture, large amount of drip,
less marbling, excessive visible fat and abnormal meat odor are
considered as poor quality traits for fresh meat. Also, the consumer
expects quality meat that is reliable in relation to safety, nutrition,
sustainability and ethics (Troy & Kerry, 2010).
3.1. Appearance quality traits (AQT)
Meat color is the most important AQT because it is the first factor
seen by the consumer and is used as an indication of freshness and
wholesomeness. Basically, meat color is dependent on species, age
and muscle type, and the color differences are due to the different
content of myoglobin (Mb) in muscle. The higher Mb content in
type I muscle fiber is due to Mb's function of storing and delivering
oxygen in the muscle. The Mb content in muscle is affected by factors
such as exercise and diet of the animal as well as genetic and environ-
mental factors. Many factors contribute to the discoloration of meat
during processing, storage and display. The predominant determinant
of meat color stability is the rate of OxyMb oxidation (Faustman, Sun,
Mancini, & Suman, 2010), and the rate of discoloration in meat is
muscle-specific. Rapid discoloration occurs in muscles that contain
greater relative proportions of type I muscle fibers because of higher
oxygen consumption rate (Jeong et al., 2009).
829
Two other important AQT for fresh meat are the amount of drip on
the surface of meat and purge in the tray. Drip and purge loss depend
on the WHC of meat, and WHC is closely related to the color of meat
due both to its role in the loss of Mb and reflectance at the surface of
the meat (Joo, Kauffman, Kim, & Kim, 1995). Additionally, WHC influ-
ences other physical properties including texture and firmness of raw
meat, and eating properties of cooked meat. Drip loss originates from
the spaces between muscle fiber bundles and the perimysial network,
and the spaces between muscle fibers and the endomysial network
(Offer & Cousins, 1992). These spaces appear during rigor develop-
ment when muscle converts to meat. It is well known that excessive
drip exudation and soft texture result from the combination of rapid
pH decline, and high temperature in p.m. muscle (Joo, Kauffman,
Kim, & Park, 1999; Warner, Kauffman, & Greaser, 1997). This is an
especially prevalent problem for pork which contains greater relative
proportions of type II muscle fibers compared to beef or lamb.
Meat texture is directly related to the size of muscle fiber and the
amount of connective tissue, and is partially affected by the quantity
of intramuscular fat (IMF). Relatively large muscle bundles are
responsible for the coarse, undesirable texture on the transversely
cut surface of meat. The diversity of muscle is attributed to the
heterogeneous characteristics of the individual muscle fibers and
the mosaic composition (Taber, 1998). Muscle fiber diameter varies
with species, chronological age, state of nutrition of the animal,
genetic background and composition of muscle fiber types (Karlsson
et al., 1993). The coarseness of the meat surface is increased with
thickened connective-tissue strands as well as increased size of
muscle bundles. The connective tissue content of meat varies with
species, chronological age, state of nutrition of the animal and muscle
fiber characteristics (Klont, Brocks, & Eikelenboom, 1998). Meat firm-
ness is also influenced by the status and quantity of the subcutaneous
fat surrounding muscles and IMF. Because IMF deposits mainly in the
perimysium between muscle bundles, meat firmness is partially
influenced by the IMF firmness which is affected by composition of
fatty acids and temperature.
It is known that IMF produces effects on flavor, juiciness, tender-
ness and visual characteristics of meat with increased marbling in
meat, although there has been extensive debate about the contribu-
tion of IMF to the tenderness of meat. The quantity of IMF is affected
by many factors including animal breed, slaughter weight (Park et al.,
2002), feeding strategy (Du, Yin, & Zhu, 2010), and growth rate
(Smith et al., 2009). In animals, adipogenesis occurs the earliest in
the visceral fat deposit, closely followed by subcutaneous and
intermuscular deposits, and adipogenesis in intramuscular fat occurs
last (Hausman et al., 2009). This adipogenesis can be affected by
genetic, nutritional and environmental factors that are the key signal-
ing pathways regulating adipogenesis in skeletal muscle (Du &
Dodson, 2011). Although there are variations among species, IMF
tends to increase with advancing age when the major stages of mus-
cle growth have been completed. IMF deposition is highly heritable
and is positively correlated with general body fatness in the animal.
Moreover, IMF is positively correlated with percentage of red muscle
fiber, but negatively correlated with white muscle fiber in muscle
(Hwang et al., 2010).
3.2. Eating quality traits (EQT)
Tenderness is the most important EQT because it strongly influ-
ences consumer's perceptions of acceptability. Meat tenderness is
mainly affected by the amount and solubility of connective tissue,
the composition and contractile state of muscle fibers, and the extent
of proteolysis in rigor muscle. Also, IMF content indirectly affects
meat tenderness. Tenderness is more important for red meat such
as beef and lamb because of a high composition of red muscle fibers
and connective tissue compared to pork or chicken. The content of
connective tissue is related to muscle fiber characteristics because
830 S.T. Joo et al. / Meat Science 95 (2013) 828–836
muscle fibers occupy 75–90% of the muscle volume, and the morphol-
ogy of the muscle fiber is a major determinant factor of mass (Lee et
al., 2010). The heterogeneity of muscle fiber characteristics in differ-
ent muscles is known to influence tenderness (Maltin, Balcerzak,
Tilley, & Delday, 2003). However, the relationship between muscle
fiber characteristics and meat tenderness is still controversial.
Muscles with diverse muscle fiber characteristics have different
patterns of p.m. change during the conversion of muscle to meat. If
type II fibers are predominant in muscle, p.m. glycolysis is rapid,
resulting in an accelerated pH decline in the muscle. In addition
throughout the p.m. period, sarcomere lengths in muscle vary
because each muscle fiber goes into rigor at different times. Conse-
quently meat tenderness varies with the rate of glycolysis, the rigor
onset post-slaughter and the extent of glycolysis, which are all related
to muscle temperature as well as muscle fiber characteristics (Ali et
al., 2008).
Flavor is also important for the eating quality of meat because
people expect certain attributes such as savoriness. Because meats
consist mainly of the lean portion and the fat portion, the meat flavor
is primarily dependent on the pool of flavor precursors in these two
tissues. Meat flavor is affected by species, sex, age, stress level,
amount of fat, and diet of animal. Beef, pork, lamb, and poultry have
distinctive flavor characteristics due to the variation of the flavor
precursors generally in the fat between and within species. The effect
of animal gender on meat flavor is highly related to testosterone and
skatole that are produced in intact males and females, respectively.
Boar taint in pork from intact males is an unpleasant urine-like
and sweaty odor that is related to the presence of androstenone
(5α-androst-16-en-3-one) and skatole (3-methylindole) (Grindflek
et al., 2011). Androstenone is a metabolite of testosterone, and
skatole is the major contributor to pastoral-flavor (Teixeira, Batista,
Delfa, & Cadavez, 2005). Testosterone increases muscle growth and
decreases intramuscular lipid deposition. In general, intact males
deposit less fat throughout the body and within muscle, and are
more susceptible to long-term pre-slaughter stress than females or
castrated males. Increasing serum-like bloody aromatics and metallic
flavor are due to increased levels of Mb in the meat of older animals.
Juiciness is positively related with the WHC of meat and the IMF
content in meat. The IMF content directly affects juiciness as well as
flavor (Hocquette et al., 2010), and the human perception of juiciness
is increased as the IMF content in meat increases (Jeremiah, Gibson,
Aalhus, & Dugan, 2003). Moreover, the feel of juiciness in the oral
cavity is generally sustained when meat has a large amount of IMF.
In general, juiciness is a more important sensory trait for pork be-
cause consumers of pork place a higher rating on juiciness than flavor
or tenderness (Aaslyng et al., 2007), while consumers of beef rate
tenderness as the most important palatability trait (Cho et al.,
2010). A lack of juiciness is a major quality issue in pork, and pork
muscle that lacks marbling exhibits a lack of juiciness. IMF content
affects juiciness by enhancing the WHC of meat, by lubricating the
muscle fibers during cooking, by increasing the tenderness of meat,
and thus the apparent sensation of juiciness, or by stimulating sali-
vary flow during mastication (Luchak et al., 1998). It is well known
that meat with a high IMF content has improved juiciness after rela-
tively long-heating in a moist environment, whereas meat of lower
IMF content is not deteriorated by severe short-heating under dry
cooking conditions.
3.3. Reliance quality traits (RQT)
Safety is always more important than AQT and EQT, and the mi-
crobial level in meat is the most important RQT for fresh meat. The
categories of meat safety also include physical and chemical residues,
food additives and animal identification of meat products. In general,
consumers evaluate meat safety by visual and odor evaluations which
are the most rapid indications of meat spoilage, although they are
unreliable indicators of safety. The importance of meat as a carrier
of bacterial pathogens is considerable in terms of public health.
Therefore, strict and stringent safety requirements in the processing
of meat have been developed and implemented in many countries.
The Hazard Analysis Critical Control Point (HACCP) system provides
the basis for the meat safety management system within the meat
chain (Troy & Kerry, 2010).
There is no doubt that quality meat is one with high nutritional
value, and meat is one of the most nutritional foods. However, recent-
ly, the concept of nutrition has changed as the nutrition of food has
reached an all-time high. In the past, quality meat was more closely
related to the sensory perceptions, freshness, and safety aspects of
meat products, whereas more recently it is associated with nutrition,
well-being and functionality in relation to human health. Consequently,
consumers may consider the high content of fat and cholesterol in meat
as undesirable and unhealthy, although meat is nutritious because it is a
rich source of protein, essential amino acids, minerals and vitamins.
Meat composition can be manipulated to alter the nutritional profile
in most cases. Dietary supplementation is the key factor which can
most easily be manipulated and has one of the most profound effects
on meat composition. Furthermore, the effect of diet on nutritional pro-
file is more profound in meat derived from monogastric animals. These
kinds of meats are categorized as functional foods which are defined as
foods with nutritional profiles that exceed conventional products
(Decker & Park, 2010; Hur, Park, & Joo, 2007).
In recent years there has been a considerable increase in consumer
concern with regard to how meat is produced. Concern about animal
welfare has greatly increased around the world, and there has been
an enormous development of the ‘organic’ rearing of animals. Con-
sumers demand that animals are reared, transported and slaughtered
under humane conditions. Also, consumers want to be confident that
the meat they purchase is derived from ethically robust production
systems. Consequently, farmers, veterinarians, packers and scientists
need to become more knowledgeable on how to assess and audit
animal welfare at the farm and slaughter plant (Grandin, 2010). It
should be emphasized that the importance of traceability has
increased in relation to RQT. Regulatory agencies in many countries
have insisted on the implementation and application of traceability
systems (Troy & Kerry, 2010).
4. Variation of meat quality traits
Attempts to identify common standards of fresh meat quality have
been done for international trade across a number of countries. How-
ever consumers still have difficulty in accurately predicting quality by
perception at the point of purchase (Glitsh, 2000). The eating quality
and the assessment of beef from the same animals in eight countries
in the EU have been investigated, and the results showed that
consumer's preferences differ among countries (Dransfield et al.,
1984). Irish and English panelists preferred flavor more than tender-
ness and juiciness, but Italian panelists tended to value tenderness
more highly than flavor. Pethick, Warner, and Banks (2006) reported
that consumers of lamb in Australia usually place the greatest weight
on flavor/odor, followed by tenderness and finally juiciness. This is in
contrast to consumers of beef who generally rate tenderness as the
most important palatability trait (Moon, Yang, Park, & Joo, 2006).
According to Warner et al. (2010), flavor has increased in importance
for beef consumers as tenderness variation in meat has been reduced.
It is well known that inherent differences in eating quality
between muscles exist. Jeremiah et al. (2003) found that 30 anatom-
ically defined bovine muscles differed significantly in EQT. It should
be emphasized that meat is composed of numerous tissues such as
adipose, epithelial, connective and nervous tissues, although the
major component is muscle. Consequently, regardless of muscle qual-
ity, variation of meat quality could be influenced by these intrinsic
factors. For example, some muscles contain relatively large quantities
 S.T. Joo et al. / Meat Science 95 (2013) 828–836
of connective tissue, which are associated with meat toughness. If
meat contains either an unexpected excess or low amount of fat, it
would be considered a low quality meat. It is obvious that fat content,
connective tissue and muscle fiber characteristics have a significant
influence on meat quality. Also, storage conditions and temperature
are very important because meat quality can deteriorate via spoilage
due to adverse storage conditions or temperature abuse. Moreover,
AQT and EQT are significantly influenced by the method of cooking
and preparation of meat cuts.
5. Relationship between meat quality traits and muscle
fiber characteristics
It is commonly reported that many aspects of meat quality are re-
lated with muscle fiber characteristics that are represented by TNF,
CSAF and FTC in muscle. In particular, these muscle fiber characteris-
tics are closely related with muscle pH that is commonly considered
as an indicator of pork quality. Especially, FTC in muscle is related to
the rate of p.m. pH decline. Increasing the proportion of fast-twitch
glycolytic fibers in porcine longissimus muscle has been shown to
increase the rate and extent of p.m. pH decline (Choi, Ryu, & Kim,
2007; Kim, Jeong, et al., 2013; Ryu & Kim, 2006). Moreover, oxidative
fibers are susceptible to cold temperature and thus rapid temperature
decline p.m. could increase muscle shortening especially if the
muscles contain a high amount of oxidative fibers (Lonergan, Zhang,
& Lonergan, 2010).
Muscle fiber characteristics influence AQT including meat color,
WHC, texture and marbling in meat. The Mb content and the rate of
Mb oxidation are muscle-specific, and increasing the proportion of
red muscle fibers is known to increase the redness and Mb content
of meat (Kim et al., 2010). It is well documented that increasing the
proportion of type I fibers decreases color stability with a possible
shift to a brownish MetMb color (Renerre, 1990). The composition
of fast-twitch glycolytic (IIB) fibers in pork muscle is related to higher
lightness and lower WHC (Kim, Jeong, et al., 2013). Hypertrophy of
fast-twitch oxido-glycolytic fibers (IIA) is more specifically detrimen-
tal to WHC (Larzul et al., 1997; Maltin et al., 1998). The size of muscle
fibers affects muscle growth potential and the size of the fiber bundle,
resulting in the visible coarseness of transverse sections of meats
(Kim, Kim, et al., 2013; Rehfeldt & Kuhn, 2006; Ryu & Kim, 2005).
The content of connective tissue including IMF also varies with
muscle fiber characteristics (Klont et al., 1998). There is a strong pos-
itive genetic correlation between CSAF and IMF content in porcine
longissimus muscle (Larzul et al., 1997). Kim, Jeong, et al. (2013),
and Kim, Kim, et al. (2013) also reported that the proportion and
size of type IIB fibers are positively related with IMF content in por-
cine longissimus muscle. In beef muscle, IMF is positively correlated
with the percentage of red muscle fiber, but negatively correlated
with white muscle fiber (Hwang et al., 2010). It is commonly stated
that red oxidative muscles contain more IMF than white glycolytic
muscle. However, Lefaucheur (2010) reported no universal relation-
ship between IMF and FTC, and suggested that both characteristics
are rather independent and can be manipulated separately.
EQT are also closely associated with muscle fiber characteristics. The
heterogeneity of muscle fiber type in different muscles is known to in-
fluence meat tenderness (Maltin et al., 2003). Slow-twitch muscles
have been reported to contain more collagen, which plays an important
role in binding muscle fibers and decreasing tenderness of meat
(Kovanen, Suominen, & Heikkinen, 1984). However, the relationship
between FTC and meat tenderness is still controversial, and no clear re-
lationship between collagen content and FTC has been reported in live-
stock species (Lefaucheur, 2010). Muscles with a larger fiber size,
especially type IIB fiber, exhibit tougher meat than muscles of smaller
fiber size in cattle (Renand, Picard, Touraille, Berge, & Lepetit, 2001)
and in pig (Karlsson et al., 1993). Conversely, Berri et al. (2007) reported
831
that CSAF showed a positive relationship with tenderness in broiler
breast muscle.
FTC of muscle is related to p.m. proteolytic degradation as well as
glycolysis and the rate of pH decline. Fast-twitch IIB fibers are known
to be highly glycolytic fibers and their metabolism contributes to a
fast metabolic rate early p.m. period (Ryu & Kim, 2005, 2006). If
fast-twitch glycolytic fibers are predominant in muscle, rapid glycol-
ysis is induced, resulting in a rapid pH decline in the muscle (Choe
et al., 2008). Thus, the composition of type IIB fibers are negatively re-
lated to muscle pH but positively related to R-value (adenine/inosine
ratio), allowing for the determination of ATP depletion in the early
p.m. period (Ryu et al., 2008). Contrarily, increasing the proportion
of type I fibers in muscle decreases the rate and extent of p.m. pH
decline (Choi et al., 2007). These differences in p.m. muscle properties
are due to different FTCs and influence meat tenderness. Moreover,
type II fast fibers are more susceptible to early p.m. proteolytic degra-
dation than type I slow fibers (Xiong et al., 2007). Hwang et al. (2010)
also reported improvement of tenderness via increasing the percent-
age of type I fibers and decreasing the percentage of type IIB in cattle
muscle.
The influence of muscle fiber characteristics on p.m. aging is another
important aspect of meat quality. The increase of fast-twitch glycolytic
fibers has beneficial effects on p.m. aging and tenderness in cattle
(Seideman, Crouse, & Cross, 1986). In p.m. aging of meat, the rate of
aging is faster in fast-twitch muscles than in slow-twitch oxidative
muscles (Totland, Kryvi, & Slinde, 1988). The calpain/calpastatin ratio
is higher in fast-twitch glycolytic muscles than in slow-switch oxidative
muscles, which could partly explain the faster rate of aging in glycolytic
muscles (Ouali & Talmant, 1990). Fast-twitch fibers have a more exten-
sively developed sarcoplasmic reticulum, transverse-tubule system,
and thinner Z-band than slow-twitch fibers, and proteins that comprise
the Z-band in fast-twitch fibers are more susceptible to early p.m.
proteolytic degradation than those in slow-twitch fibers (Xiong, 2004).
Meat flavor and juiciness are strongly affected by IMF content in
muscle that is positively correlated with proportion of type I fibers
in muscle (Maltin et al., 1998). A high proportion of type I fibers is
associated with a high level of phospholipids which are important
determinant of cooked meat flavor (Hwang et al., 2010). Also, the
content of type I fiber is positively related with juiciness (Calkins,
Duston, Smith, Carpenter, & Davis, 1981). However, Lefaucheur
(2010) suggested that intramuscular adipocytes within a muscle
were not related to its FTC, although research has shown that red
oxidative muscles contain more total IMF than white glycolytic
muscles. Therefore, the relationship among FTC, IMF and EQT still
remains a challenge.
6. Potential factors to manipulate muscle fiber characteristics
6.1. Breed and genotype
One of the important factors that influence TNF, CSAF and FTC of a
given muscle within a species is breed. In general, wild animals
contain more oxidative fibers, less glycolytic fibers and smaller fibers
compared to domesticated animals (Lefaucheur, 2010). Animal selec-
tion for increased growth rate and lean meat content shifts muscle
metabolism towards a more white glycolytic and less red oxidative
type (Rahelic & Puac, 1981). For example, Meishan pigs have a higher
oxidative and a lower glycolytic metabolism with a decrease in
TNF and CSAF compared to LW pigs (Bonneau, Mourot, Noblet,
Lefaucheur, & Bidanel, 1990), and LW pigs contain more type I fibers
than miniature pigs (Stickland & Handel, 1986). Also, there is a posi-
tive relationship between high muscularity and a high proportion of
myosin heavy chain (MHC) IIb transcript in different pig breeds
(Wimmers et al., 2008). The longissimus and gluteus medius muscles
from Hampshire pigs have a greater oxidative capacity, a lower glyco-
lytic capacity, and a higher concentration of glycogen than those of
832 S.T. Joo et al. / Meat Science 95 (2013) 828–836
the Swedish Yorkshire (Essén-Gustavsson & Fjelkner-Modig, 1985).
The longissimus dorsi muscle of Berkshire pigs has a larger percentage
of type I fiber compared to Landrace and Yorkshire pigs (Ryu et al.,
2008). Therefore, it should be emphasized that an increase in carcass
lean percentage is positively related to glycolytic and oxidative
metabolism in longissimus muscle as well as lean tissue growth rate
(Karlsson et al., 1993), although no opposite or significant correla-
tions between lean tissue growth rate and FTC have been reported
(Candek-Potokar, Lefaucheur, Zlender, & Bonneau, 1999; Ryu, Rhee,
& Kim, 2004). Consequently, there is a strong relationship between
FTC and growth performance when comparing extremely different
genetic breeds, but this is often controversial within conventional
domesticated animals (Lefaucheur, 2010). All these findings imply
that FTC in muscle could be manipulated by breeding of animals.
On the other hand, the progressive improvement in meat quality
can be achieved through selection of animals that have a ‘quality
gene’. According to Warner, Greenwood, and Ferguson (2011), inter-
action between genotype and environment contributes to the pheno-
typic variation in AQT and EQT, and increasing muscle mass is
achievable by targeted selection for specific mutations, for example,
the myostatin gene. The major genes that induce an increase in
carcass lean content are the double-muscling myostatin gene in cat-
tle, the Callipyge gene in sheep and Ryr1, RN (Rendement Napole)
or IGF-II genes in pig. The effects of these genes on TNF, CSAF and
FTC are well documented, and only the mutated myostatin gene is
associated with an additional increase in TNF (Lefaucheur, 2010).
The improvement of tenderness in double-muscled cattle is due to a
lower total and higher soluble collagen content (Ngapo et al., 2002).
Increased muscle content of Callipyge sheep is related to increased pro-
portion of white muscle fibers (Koohmaraie, Shackelford, Wheeler,
Lonergan, & Doumit, 1995). The specific increased CSAF of fast muscle
fiber in Callipyge sheep reduces protein degradation, including a lower
p.m. proteolysis, maturation and tenderization of meat (Lorenzen et
al., 2000). Increasing muscle content of Ryr1 pigs is due to increased
CSAF with similar or reduced TNF (Depreux, Grant, & Gerrard, 2002).
The Ryr1 gene pigs show decreased IIA mRNA and increased MHC IIX
mRNA in muscle, while the RN mutation decreases MHC IIB mRNA and
increases IIA and IIX mRNA expression (Park, Gunawan, Scheffler,
Grant, & Gerrard, 2009). The Ryr1 gene induces a punctual mutation in
the SR Ca2+ channel, leading to an abnormally high release of Ca 2+ in
the sarcoplasm and rapid pH decline early p.m., resulting in PSE pork
(Lefaucheur, 2010). All these changes in FTC due to mutated genes in
pigs are strongly related to poor pork quality. Therefore, efforts to elim-
inate these genes from pig herds are necessary to improve the AQT and
EQT of pork.
6.2. Sex and hormone
There are sex differences relative to muscle fiber characteristics,
although Staun (1963), Rowe and Goldspink (1969) and Miller,
Garwood, and Judge (1975) did not find any sex differences in TNF,
in CSAF and in FTC, respectively. The smaller type IIA and type IIB fi-
bers in the longissimus dorsi muscles from boars indicate that TNF is
higher in male pigs compared to female pigs (Petersen, Henckel,
Oksbjerg, & Sørensen, 1998). The CSAF of fiber types I, IIA and IIB is
smaller in the longissimus dorsi muscle of entire male pigs compared
to gilts (Karlsson et al., 1993). Essentially, females exhibit larger fibers
with no difference in fiber type percentages and relative areas (Larzul
et al., 1999; Miller et al., 1975; Solomon, Campbell, & Steele, 1990).
Some hormones have a profound influence on the muscle fiber
characteristics of specific muscles. Differences in TNF and CSAF are
primarily controlled by sex hormones. Differences in TNF between
males and females can arise by hormonal action if differences in
androgen hormones are sufficiently high during periods of prenatal
fiber formation (Rehfeldt et al., 2004). Yoshioka, Boivin, Bolduc,
and St-Amand (2007) reported that testosterone treatment in later
postnatal periods can stimulate muscle hypertrophy in a direct or
indirect manner by satellite cell proliferation and muscle protein
synthesis, without increasing TNF. Hormonal differences, especially
testosterone, may also contribute to the gender differences in specific
fiber type sizes that ultimately affect the relative concentrations of
MHC isoforms (Staron et al., 2000).
Muscle fiber phenotypes are dramatically changed by thyroid hor-
mones. Hypothyroidism causes fast-to-slow transitions, while hyper-
thyroidism induces transitions in the reverse direction (Fitts, Winder,
Brooke, Kaiser, & Holloszy, 1980; Ianuzzo, Patel, Chen, O'Brien, &
Williams, 1977). Many studies show that low levels of thyroid hor-
mones cause fast-to-slow shifts in MHC isoform expression: MHC
IIB → MHC IIX(D) → MHC IIA → MHC I/slow, whereas high levels
of thyroid hormones induce slow-to-fast shifts in MHC isoform
expression: MHC I/slow → MHC IIA → MHC IIX(D) → MHC IIB
(Caiozzo, Herrick, & Baldwin, 1992; Fitzsimons, Herrick, & Baldwin,
1990; Izumo, Nadal-Ginard, & Mahdavi, 1986). In addition, low
thyroid hormone levels inhibit or delay the appearance of adult
fast-twitch muscle fibers, whereas high levels of thyroid hormone ac-
celerates the transition from developmental fibers to adult fast-twitch
fibers (Adams, McCue, Zeng, & Baldwin, 1999).
According to Ryu, Choi, Ko, and Kim (2007), serum insulin-like
growth factor-I (IGF-I) concentrations are negatively related to the
composition of type I fibers, whereas serum epidermal growth factor
is positively correlated to the composition of type I fiber. The propor-
tion of type IIB fibers has a positive correlation with IGF-I expression
(Owens, Campbell, Francis, & Quinn, 1994). In β-agonist-fed lambs,
muscle hypertrophy is associated with a selective increase in CSAF
of glycolytic fibers, whereas effects on FTC are unclear (Beermann et
al., 1987). In pigs, treatment with β-agonist increases the frequency
of type IIB fibers, mainly at the expense of type IIA fibers, resulting
in lower activities of oxidative enzymes (Oksbjerg, Henckel, &
Rolph, 1994).
6.3. Growth performance and diet
The muscle fiber characteristics are significant for growth perfor-
mance. In most farm animals, myogenesis is a biphasic phenomenon
involving the successive differentiation of a primary and secondary
generation of myotubes during the fetal period. The number of
secondary generation fibers varies by fetus weight or gestation period
of animals. In general, TNF is considered to be established by 90 dg of
fetus (i.e. after 80% of gestation) in pig and 180 dg of fetus (i.e. after
66% of gestation) in cattle (Picard, Lefaucheur, Berri, & Duclos, 2002;
Wigmore & Stickland, 1983). The fetal stage of animals is character-
ized by the differentiation of fibers strongly expressing the slow
MHC isoform derived from primary fibers, and of fibers strongly
expressing the neonatal MHC isoform derived from both primary
and secondary fibers (Condon, Silberstein, Blau, & Thompson, 1990;
Lyons, Ontell, Cox, Sassoon, & Buckingham, 1990). CSAF remains con-
stant during gestation, but FTC shifts dramatically to a greater extent
in future glycolytic than oxidative fibers after birth (Lefaucheur,
2010). At birth, muscle is composed of oxidative fibers and mRNAs
of the fast MHC isoform can be detected a few days after birth
(Moody, Enser, Wood, Restall, & Lister, 1978). The proportion of
oxidative fibers decreases while the proportion of glycolytic fibers
increases during growth (Lefaucheur & Vigneron, 1986; Solomon et
al., 1990).
TNF, CSAF and FTC are factors that affect muscle growth perfor-
mance and meat quality (Rehfeldt & Kuhn, 2006). The size of various
fiber types in longissimus dorsi muscle is increased at different rates,
while the CSAF of all fibers is increased during growth (Hegarty &
Allen, 1978). The diameter of type II fibers is increased faster than
that of type I (Oksbjerg et al., 1994). Carter et al. (2010) reported
that fast MHC expression increased during aging in the rat soleus
muscle due to a marked increase in fibers that co-express both fast
 S.T. Joo et al. / Meat Science 95 (2013) 828–836
and slow MHC isoforms. Recently, Kim, Kim, et al. (2013) reported a
decrease in type I and IIA fibers and an increase in type IIB fibers
with increasing pig carcass weight. In the carcass weight range of
70 kg to 109 kg, the diameter of type IIB is positively correlated
with loin-eye area and carcass weight, and an excessively high weight
of carcass has an influence on pale and exudative properties in pork
because of muscle fiber characteristics, especially increased type IIB
fibers (Kim, Kim, et al., 2013).
TNF and CSAF are negatively correlated with each other, and mus-
cle mass is closely related to CSAF at a constant TNF (Kim, Jeong, et al.,
2013; Rehfeldt, Fiedler, Dietl, & Ender, 2000). However, when low
birth weight pigs are compared to normal weight pigs, CSAF shows
a negative correlation with lean meat content in carcass, and low
birth weight pigs have a low TNF, larger CSAF, higher IMF content in
muscle and a decreased lean meat content in carcasses compared to
those of normal birth weight pigs (Gondret et al., 2006; Rehfeldt &
Kuhn, 2006). These differences imply that low birth weight pigs
induce a higher physiological maturity, and muscle fiber hypertrophy
is achieved earlier because of a lower TNF (Lefaucheur, 2010). There-
fore, a high TNF combined with normal CSAF is suggested as a way to
increase muscle growth potential without decreasing oxidative ca-
pacity which could be induced by excessive muscle fiber hypertrophy
(Lefaucheur, 2010).
On the other hand, the adequate level of energy and the balance of
nutrients play an important role in determining growth rate and feed
efficiency in the fetal as well as in the postnatal period. During the
fetal period, malnutrition specifically decreases the number of
secondary fibers, leading to a permanent decrease in postnatal mus-
cle growth potential (Hegarty & Allen, 1978; Wigmore & Stickland,
1983). In contrast, over-nutrition of the sow between 25 and
50 days of gestation can increase the total number of fibers in the
developing pigs (Rehfeldt, Fiedler, Weikard, Kanitz, & Ender, 1993).
In lactating sows, a selective decrease in CSAF of glycolytic fibers,
and an increase in relative CSAF of type I fibers are found in
longissimus muscle (Lefaucheur, 1990).
In general, muscle fiber hypertrophy is impaired by restricting
feed intake regardless of the pig's growth stage, whereas the effect
of restricted feeding on FTC appears to vary depending on the growth
stage. According to Harrison, Rowlerson, and Dauncey (1996), feed
restriction at an early stage (between 3 and 7 weeks of age) does
not change FTC in longissimus muscle, but does lead to a dramatic in-
crease in the proportion of type I fibers in the red rhomboideus muscle
and lower CSAF of all fibers. Lefaucheur (1990) also reported that feed
restriction between 7 and 100 kg of body weight did not change fiber
type percentages in pig longissimus and tibialis cranialis muscles, but
increased the CSAF. The FTC in longissimus muscle is not changed
due to feed restriction in post-weaning and growing-finishing pigs
(Bee et al., 2007). However, restricted feeding induces a lower
proportion of type IIB fibers and more type IIA fibers in porcine
longissimus muscle during growth (Solomon & Lynch, 1988). These
changes in muscle fiber characteristics due to restricted feeding
would be related to deterioration of meat quality. Candek-Potokar
et al. (1999) reported that a 30% restriction in feed intake during
the growing-finishing period caused an increase in drip loss and light-
ness with a decrease in IMF content in porcine longissimus dorsi mus-
cle. In lambs, feed restriction induced atrophy of muscle fibers and
increased the percentage of type I fibers at the expense of type IIB
fibers (Solomon, Caperna, Mroz, & Steele, 1994).
The energetic density of the diet (2.79 vs 1.87 Mcal ME/kg diet)
was shown to influence FTC in lambs given ad libitum access to feed
and slaughtered at 45 kg of body weight (Solomon & Lynch, 1988).
The longissimus muscles from lambs fed the lower-energy diet
contained more type I and fewer type IIB fibers (Solomon & Lynch,
1988). Jeong et al. (2012) reported that FTC was not affected by
dietary energy level (3.0 vs 3.2 Mcal DE/kg) in porcine longissimus
muscle, while type IIB fiber size was greater in the low energy group
833
(3.0 Mcal DE/kg) than in the medium energy group (3.2 Mcal DE/kg).
Consequently, control of diet is another potential tool which can be
combined with growth performance in different rearing systems to
manipulate muscle fiber characteristics in relation to meat quality.
6.4. Muscle location, exercise, and ambient temperature
It is well known that muscle fiber characteristics are affected by mus-
cle type, location and function within an animal. Basically, the histochem-
ical characteristics of muscle fiber depend on muscle location and
function in animals. In general, a high degree of type IIB fibers is found
in porcine longissimus dorsi (Kiessling & Hansson, 1983; Kim, Kim, et al.,
2013), gluteus medius (Essén-Gustavsson & Fjelkner-Modig, 1985), biceps
femoris (Barton-Gade, 1981), quadriceps femoris (Barton-Gade, 1981),
vastus lateralis (Kiessling & Hansson, 1983), and semimembranosus
(Barton-Gade, 1981; Monin, Mejenes-Quijano, Talmant, & Sellier, 1987)
muscles. Examples of porcine muscles containing a high degree of I and
IIA fibers are masseter, trapezius (Monin et al., 1987) and triceps brachii
(Kiessling & Hansson, 1983) muscles. In bovine muscles, longissimus
and semimembranosus muscles have a high proportion of IIB fibers
(Hwang et al., 2010; Ozawa et al., 2000), whereas a high degree of type
I fibers is found in psoas major, biceps brachii and brachialis muscles
(Hwang et al., 2010; Kirchofer, Calkins, & Gwartney, 2002). Generally,
deep muscles involved in maintaining posture are more oxidative and
contain more type I fibers than more superficial muscles involved in
rapid movements (Rosser, Norris, & Nemeth, 1992). The proportion of
type I fiber in porcine semitendinosus muscle ranges from approximately
4% in the superficial white portion to 45% in the deep red portion
(Beermann et al., 1990). In cattle, the increase in fiber diameter stops
later in the light than in the dark area of semitendinosus muscle, coincid-
ing with an increase in the percentage of type IIB fibers in the light area
(Dreyer, Naudé, Henning, & Rossouw, 1977). This information regarding
muscle fiber characteristics in relation to individual muscles is very useful
in producing quality meat cuts in meat markets, as the meat industry in
many countries has shown trends towards marketing individual muscles
for consumption (Hwang et al., 2010; Jeremiah et al., 2003; Kirchofer et
al., 2002).
Although muscle fibers are dynamic structures that exhibit high
plasticity, fibers undergo type shift following an obligatory pathway:
I ↔ IIA ↔ IIX ↔ IIB (Pette & Staron, 2000; Schiaffino & Reggiani,
1996). The FTC in muscle can be changed by physical exercise,
depending on the type and duration of the activity. Prolonged endur-
ance induces a IIB → IIX → IIA → I transition in muscles involved in
the exercise (Lefaucheur, 2010). This transition of fiber types in mus-
cle has been found in miniature pigs (Mcallister, Reiter, Amann, &
Laughlin, 1997), and confirmed in more commercial pigs (Petersen
et al., 1998). Recently, understanding the transition mechanisms of
fiber types and manipulating FTC in individual muscles via exercise
of animals have become important to the meat industry, because of
the development of ‘organic’ rearing of the animal and consumer
concern about animal welfare. Moreover, the FTC can be changed
through ambient temperature at animal rearing systems. For exam-
ple, long term cold exposure shows a shift in FTC to a slower type in
oxidative muscles involved in posture in LW pigs (Herpin &
Lefaucheur, 1992). Contrarily, if pigs are reared in warm environ-
ments, both oxidative and glycolytic metabolism decreased (Rinaldo
& Le Dividich, 1991). Consequently, the FTC in individual muscles
can be manipulated by physical exercise combined with ambient
temperature in conventional rearing systems.
7. Conclusion
The term ‘fresh meat quality’ should be defined by consumer's
preferences that are determined by AQT and EQT as well as RQT of
meat. The AQT include meat color, drip and purge, texture and
firmness, and marbling, while EQT consist of tenderness, flavor and
834 S.T. Joo et al. / Meat Science 95 (2013) 828–836
juiciness. The components of RQT are safety, nutritional value, animal
welfare and ethics. These quality traits are evaluated by the consumer
at the point of purchase, and contribute to the consumer's expecta-
tion of high quality meat. However, consumer's preferences are dif-
ferent according to country because their significance is determined
by regional preference and by the experiences of the individual con-
sumer. Moreover, variations of meat quality traits exist according to
individual muscles, and the inherent differences in eating quality
among muscles also exist. Therefore, better control of meat quality
requires a thorough understanding of meat quality as a complex
global concept submitted to the influence of various meat quality
traits.
AQT and EQT are dependent on muscle fiber characteristics such
as TNF, CSAF and FTC in meat. The rate of p.m. pH decline that influ-
ences AQT and EQT is closely related to muscle fiber characteristics,
especially FTC in muscle. Numerous studies have reported that a
high proportion of fast-twitch glycolytic fibers (type II fibers) in mus-
cle increases the rate and extent of p.m. pH decline, while a high pro-
portion of oxidative fibers (type I fiber) increases muscle shortening
and level of phospholipids in muscle influencing tenderness and fla-
vor. Increasing the proportion of type IIB fibers in porcine muscles de-
creases WHC and tenderness of pork due to increased CSAF, whereas
increasing the proportion of type IIB fibers in bovine muscle increases
tenderness of beef due to improved p.m. aging. Although these are
general findings in the present paper, controversial perspectives still
remain, and the relationship between meat quality traits and muscle
fiber characteristics is yet to be fully established. Because both quality
traits and fiber characteristics are influenced by various factors
including species, breeds, gender, age, birth weight, exercise, use of
hormones, slaughter weight, muscle type and location, sampling
sites within muscle, etc., further strictly controlled studies are needed
to understand this uncertain relationship.
Nevertheless, the present knowledge and technologies suggest
that the progressive improvement in meat quality could be achieved
by manipulation of muscle fiber characteristics in animals. This
manipulation may be possible through breeding of animals and
using specific genes or gene markers. Also, muscle fiber characteris-
tics could be changed via control of growth performance and gender
of animals. Finally, dramatic changes in FTC could be achieved by
treatment with specific hormones, and energy level and nutrient
balance during animal growth. In addition, the FTC in individual
muscles can be manipulated by physical exercise combined with
ambient temperature at conventional rearing systems. Of course, an
approach combined with all potential factors may be the best way
to achieve the progressive improvement in meat quality. This would
ensure continued production of high quality meat through manipula-
tion of muscle fiber characteristics in animals.
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