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290 Part 3: Meat, Poultry and Seafoods
metabolic enzymes (mitochondrial, lysosomal, microsomal, nu-
clear, or free in the cytosol) and myoglobin. Some of these en-
zymes play a very important role in postmortem meat and during
further processing, as described in Chapter 16. Minor amounts
of hemoglobin may be found in the muscle if blood has not been
drained properly. Myoglobin is the main sarcoplasmic protein,
responsible for the red color of meat as well as the typical pink
color of drippings. The amount of myoglobin depends on many
factors. Red fibers contain higher amounts of myoglobin than
white fibers. The species is very important, and thus beef and
lamb contain more myoglobin than pork and poultry. For a given
species, the myoglobin content in the muscle increases with the
age of the animal.
Connective Tissue Proteins
Collagen, reticulin, and elastin constitute the main stromal pro-
teins in connective tissue. There are several types (I–V) of colla-
gen containing different polypeptide chains (up to tenα-chains).
Type I collagen is the major component of the epimysium and
perimysium that surround the muscles. Types III, IV, and V col-
lagen are found in the endomysium, which provides support to
the muscle fiber (Eskin 1990). There are a high number of cross-
linkages in the collagen fibers that increase with age, and this
is why meat tougher in older animals. Elastin is found in lower
amounts, usually in capillaries, tendons, nerves, and ligaments.
Enzymes
Muscle contains a large number of enzymes, mainly peptidases
like calpains, cathepsins, proteasome, tri- and dipeptidylpep-
tidases, aminopeptidases, carboxypeptidases, and dipeptidases,
but also lipases, glycohydrolase, nucleotidases, and so on (Toldra ́
2007). All these enzymes have a special relevance for meat pro-
cessing and they are covered in-depth in Chapter 16.
Non-protein Compounds
Free Amino Acids
The action of muscle aminopeptidases contributes to the gener-
ation of free amino acids in living muscle. An example of the
typical content of free amino acids, at less than 45 minutes post-
mortem, in glycolytic and oxidative porcine muscles is shown
in Table 15.3. It can be observed that most of the amino acids
are present in significantly higher amounts in the oxidative mus-
cle (Aristoy and Toldra 1998). The free amino acid content is ́
relatively low just postmortem, but it is substantially increased
during postmortem storage due to the action of the proteolytic
chain, which is very active and stable during meat aging.
Dipeptides
Muscle contains three natural dipeptides: carnosine (β-alanyl-l-
histidine), anserine (β-alanyl-l-1-methylhistidine), and balenine
(β-alanyl-l-3-methylhistidine). These dipeptides perform some
physiological functions in muscle, for example, as buffers, an-
Table 15.3.Example of the Composition in Free Amino
Acids (Expressed as mg/100g of Muscle) of the
Glycolytic MuscleLongissimus Dorsiand Oxidative
MuscleTrapezius.
Amino Acids
Muscle
Longissimus
dorsi
Muscle
Trapezius
Essential
Histidine 2.90 4.12
Threonine 2.86 4.30
Valine 2.78 2.09
Methionine 0.90 1.01
Isoleucine 1.52 1.11
Leucine 2.43 1.82
Phenylalanine 1.51 1.25
Lysine 2.57 0.22
Nonessential
Aspartic acid 0.39 0.74
Glutamic acid 2.03 5.97
Serine 2.02 4.43
Asparagine 0.91 1.63
Glycine 6.01 12.48
Glutamine 38.88 161.81
Alanine 11.29 26.17
Arginine 5.19 5.51
Proline 2.83 4.45
Tyrosine 2.11 1.63
Ornithine 0.83 0.83
Source: Aristoy and Toldra 1998. ́
tioxidants, neurotransmitters, and modulators of enzyme action
(Chan and Decker 1994, Gianelli et al. 2000). Dipeptide content
is especially higher in muscles with glycolytic metabolism (see
Table 15.4), but it varies with the animal species, age, and diet
(Aristoy and Toldr ́a 1998, Aristoy et al. 2004). Beef and pork
have a higher content of carnosine and are lower in anserine,
lamb has similar amounts of carnosine and anserine, and poultry
is very rich in anserine (see Table 15.3). Balenine is present in
minor amounts in pork muscle but at very low concentrations
in other animal muscle, except in marine mammalians such as
dolphins and whales (Aristoy and Toldr ́a 2004).
Muscle and Adipose Tissue Lipids
Skeletal muscle contains a variable amount of lipids, between
1% and 13%. Lipid content mainly depends on the degree of
fattening and the amount of adipose tissue. Lipids can be found
within the muscle (intramuscular), between muscles (intermus-
cular), and in adipose tissue. Intramuscular lipids are mainly
composed of triacylglycerols, which are stored in fat cells, and
phospholipids, which are located in cell membranes. The amount
of cholesterol in lean meat is around 50–70 mg/100 g. Intermus-
cular and adipose tissue lipids are mainly composed of tria-
cylglycerols and small amounts of cholesterol, around 40–60
mg/100g (Toldr ́a and Flores 2004).