Research Article

Received: 17 March 2014 Revised: 23 July 2014 Accepted article published: 23 August 2014 Published online in Wiley Online Library: 23 September 2014

(wileyonlinelibrary.com) DOI 10.1002/jsfa.6883

Meat quality and the histological structure

of breast and leg muscles in Ayam Cemani
chickens, Ayam Cemani × Sussex hybrids
and slow-growing Hubbard JA 957 chickens
Monika Łukasiewicz,* Jan Niemiec, Agnieszka Wnuk
and Natalia Mroczek-Sosnowska

Abstract
BACKGROUND: The purpose of this study was to determine the quality of meat and the histological structure of muscles of Ayam
Cemani chickens, Ayam Cemani × Sussex hybrids and slow-growing Hubbard JA 957 chickens and to examine whether crossing
generally available Sussex chickens with little available Ayam Cemani gives a good quality product of interest to the poultry
industry and in food technology.

RESULTS: The size of breast and leg muscle fibers varied among genotypes. The breast and leg muscles of slow-growing Hubbard
JA 957 chickens had the largest fiber diameter. The histological and biochemical properties of muscles, including the type,
number, proportions, diameter and metabolic profile of fibers, had a significant effect on the pH and water-binding capacity
of meat, thus affecting its quality. The muscle fibers of Ayam Cemani chickens were approximately half the size of the muscle
fibers of Hubbard JA 957 chickens. Ayam Cemani and Ayam Cemani × Sussex gave a product of as good quality as Hubbard JA
957 chickens.

CONCLUSION: Meat from Ayam Cemani chickens is a rich source of protein and could be highly valued by gourmet consumers,
connoisseurs and dieticians for its rarity and originality. The results of this study show that genotype (Ayam Cemani, Ayam
Cemani × Sussex, Hubbard JA 957) affected the quality and color of meat and the histological profile of chicken breast and leg
muscles.
© 2014 Society of Chemical Industry

Keywords: chickens; histological structure of muscles; meat quality

INTRODUCTION
The quality and texture characteristics of meat and meat products
are considered essential to commercial success. Meat quality
is determined, among others, by its sensory properties, chem-
ical composition and physical characteristics.1,2 However, it
should be noted that fast-growing broiler chickens selected for
increased muscle mass are also characterized by thicker muscle
fibers.3 The metabolic profile of muscle fibers is closely corre-
lated with the color, taste, tenderness and juiciness of meat. The
biochemical and microstructural properties of muscle fibers4
are also associated with a drop in meat pH, reduced cooking
loss and meat color. Quality is an important determinant of
the market share of meat and meat products, the processing
suitability of meat and its preparation for human consump-
tion, Meat quality is largely dependent on muscle structure,
including the content of connective tissue and myofibrillar
proteins.5
Ayam Cemani is a breed of chickens with characteristic black
plumage, comb and wattles, beak, eyes, skin and legs and even
1730
country of origin, Ayam Cemani are raised for eggs and meat to
meet domestic market needs. In Java, Ayam Cemani are also used
in ritual ceremonies and in folk medicine to treat cardiovascular
and respiratory diseases.6,7 In Bali, Ayam Cemani are used as
gamecocks in cockfighting, which is considered an age-long cul-
tural tradition and an ancient religious ritual rather than a sport or
an entertainment. In Europe, the Ayam Cemani is an ornamental
chicken breed.
The purpose of this study was to determine the quality of meat
and the histological structure of muscles of Ayam Cemani chick-
ens, Ayam Cemani × Sussex hybrids and slow-growing Hubbard
JA 957 chickens and to investigate whether crossing generally
available Sussex chickens with little available Ayam Cemani gives
∗ Correspondence to: Monika Łukasiewicz, Department of Poultry Breeding, Fac-
ulty of Animal Science, Warsaw University of Life Sciences – SGGW, Warsaw,
Poland. E-mail: monika_lukasiewicz@sggw.pl
Department of Poultry Breeding, Faculty of Animal Science, Warsaw University

black internal organs, bones and muscles. In Indonesia, their of Life Sciences – SGGW, Warsaw, Poland

J Sci Food Agric 2015; 95: 1730–1735 www.soci.org © 2014 Society of Chemical Industry
Meat quality of Ayam Cemani chickens www.soci.org

a good quality product of interest to the poultry industry and in Statistical analysis
food technology. The data obtained were analyzed statistically by a multifactor anal-
ysis of variance (least squares) using SPSS 21.0 software (SPSS,
Chicago, IL, USA). Only significant interactions between factors
EXPERIMENTAL (P ≤ 0.01 or P ≤ 0.05) were considered in the study. The level of sig-
Animals nificance was determined after performing preliminary statistical
The experimental materials comprised Ayam Cemani chickens analyses.
(group A, 120 birds), Ayam Cemani × Sussex hybrids (group M, 120
birds) and slow-growing Hubbard JA 957 chickens (group H, 120
birds), individually marked. The chickens were raised in confine- RESULTS AND DISCUSSION
ment until 9 weeks of age under typical husbandry conditions on The nutritional value and processing suitability of meat are
cut straw. Day-old chicks were weighed and assigned to one of the affected by its chemical composition. In this study the effect of
above three groups (A, M, H). Each group was further divided into sex on the proximate chemical composition of chicken muscles
four subgroups (replicates) of 30 birds per replicate. Stock density was not significant (P > 0.05), which is why results regarding meat
in a poultry house was 11 birds m−2 . quality are not shown separately for males and females.
A three-stage feeding program was used during the rearing Tables 1–3 present the chemical composition and physico-
period. Experimental diets had identical nutrient levels. Until 21 chemical properties of the breast and leg muscles of chickens. The
days of age, the chickens were fed a standard starter diet con- water content of breast muscles ranged from 73.53 g per 100 g in
taining 209.8 g kg−1 total protein, 2845 kcal kg−1 metabolizable Ayam Cemani to 74.24 g per 100 g in Hubbard JA 957, while the fat
energy, 11.7 g kg−1 lysine and 9.8 g kg−1 methionine + cystine. content ranged from 0.97 g per 100 g in hybrids to 1.03 g per 100 g
From 22 to 56 days of age, the birds received a grower diet con- in Ayam Cemani. Group A and M chickens were characterized
taining 200 g kg−1 total protein, 2900 kcal kg−1 metabolizable by a higher protein content of breast muscles, which reached
energy, 10.4 g kg−1 lysine and 8.8 g kg−1 methionine + cystine. 24.03 and 24.10 g per 100 g respectively compared with 23.13 g
The finisher diet (administered until the end of the rear- per 100 g in group H. Significant (P ≤ 0.01) differences in the ash
ing period) contained 180 g kg−1 total protein, 2965 kcal kg−1 content of breast muscles were observed between groups. Ayam
metabolizable energy, 9.0 g kg−1 lysine and 7.8% g kg−1 methio- Cemani chickens had the lowest ash content of breast muscles,
nine + cystine. The chickens had free access to feed and while higher values were noted in hybrid and Hubbard chickens
water. (Table 1).
At 63 days of age, 12 males and 12 females representing an
average body weight of each group were fasted for 12 h and
slaughtered in a poultry slaughterhouse. The chickens were elec- Table 1. Chemical composition (g per 100 g) of chicken meat (♀ + ♂)
trically stunned in a water bath (120 mA, 50 Hz) for 2 s, slaugh-
Breast muscles Leg muscles
tered by cutting the cervical blood vessels and bled out for ca
7–10 min. After scalding in water at a temperature of 56–58 ∘ C Group Water Protein Fat Ash Water Protein Fat Ash
for ca 60 s, the birds were manually plucked and eviscerated and
their carcasses were placed in a cold store at a temperature of 4 ∘ C H 74.27 23.13B 1.00 1.20A 71.87 19.47b 7.23 1.10
for 24 h. A 73.53 24.03A 1.03 1.10B 72.67 20.63a 5.23 1.07
Sections of M. pectoralis superficialis and M. biceps femoris M 73.57 24.10A 0.97 1.17A 72.80 20.70a 4.97 1.07
were collected for histological analyses. Specimens measuring SEM 0.22 0.14 0.15 0.02 0.65 0.27 0.83 0.04
0.5 cm × 0.5 cm × 1 cm were taken within 15 min post mortem,
after carcass bleeding. After 24 h of fixation, the specimens were Means in a column with different capital or lowercase letters are
significantly different at P ≤ 0.01 or P ≤ 0.05 respectively. H, Hubbard
rinsed in ethanol to remove residual fixative and dehydrated in JA 957 chickens; A, Ayam Cemani chickens; M, Ayam Cemani × Sussex
a graded ethanol series. Dehydrated specimens were infiltrated hybrids; SEM, standard error of mean.
with liquid paraffin for several hours, depending on the muscle
section characteristics, and then embedded in paraffin. A Leica RM
2265 rotary microtome (Leica Microsystems, Nussloch, Germany)
was used to cut paraffin blocks into 6 μm thick cross-sections. Table 2. Physicochemical properties of chicken meat (♀ + ♂)
The sections were stained with hematoxylin and eosin.8 In each
Breast muscles Leg muscles
specimen the diameters of 300 muscle fibers were measured
under a Nikon Eclipse E200 MV RS light microscope (Tokyo, Japan) Cooking WBC Cooking WBC
equipped with a Nikon DS-Fi2 camera and COOL View 2.7 image loss (g per (g per loss (g per (g per
analysis software. Group pH24 100 g) 100 g) pH24 100 g) 100 g)
To determine meat quality, chicken carcasses were air chilled
at 4 ∘ C for 24 h. Nine samples of breast and leg muscles were col- H 5.80a 18.35A 13.15b 6.27 24.36A 10.25b
lected from male and female chickens in each group. At 24 h post A 5.67b 8.23B 7.30a 6.15 12.64B 7.32a
mortem, muscle samples were analyzed to determine pH (accord- M 5.71 13.48 10.73 6.06 20.16 9.67b
ing to PN-ISO 29179 ), water-binding capacity (by centrifugation, SEM 0.03 0.06 1.82 0.06 1.37 0.43
as described by Wierbicki et al.10 ), cooking loss (30 g samples of
ground meat were placed in beakers, covered with self-adhesive Means in a column with different capital or lowercase letters are
significantly different at P ≤ 0.01 or P ≤ 0.05 respectively. H, Hubbard
polyethylene film and heated in a water bath at 72 ∘ C for 30 min) JA 957 chickens; A, Ayam Cemani chickens; M, Ayam Cemani × Sussex
and proximate chemical composition (contents of water, protein, hybrids; SEM, standard error of mean; WBC, water-binding capacity.
1731

fat and ash, by standard methods11 ).

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 www.soci.org M Łukasiewicz et al.

Meat and products of animal origin are major components of the

Table 3. Color parameters of chicken meat (♀ + ♂)
human diet. Protein, which is a key component of meat, is needed
Breast muscles Leg muscles by the human body for growth and development. Animal protein
has the highest nutritional value because, unlike vegetable pro-
Group L* a* b* L* a* b* tein, it contains all essential amino acids. Poultry meat is a valuable
H 51.53A 0.83B 7.96B 54.17A 5.57A 12.23B source of protein whose nutritional value is high and comparable
A 35.64B −2.25Aa 12.26A 28.68B −0.62Bb 16.26A
to the FAO/WHO amino acid reference standard. Protein from
poultry meat contains all essential amino acids, including leucine,
M 36.81B −1.08Ab 14.40A 32.07B 0.96Ba 16.26A
isoleucine, valine and arginine. According to Smolińska et al.,12
SEM 1.40 0.35 0.80 1.37 0.43 0.57
meat from landfowl has a higher protein content than meat from
Means in a column with different capital or lowercase letters are waterfowl; however, in older gallinaceous birds, protein concen-
significantly different at P ≤ 0.01 or P ≤ 0.05 respectively. H, Hubbard trations decrease to ca 16–18 g per 100 g owing to an increase
JA 957 chickens; A, Ayam Cemani chickens; M, Ayam Cemani × Sussex in carcass fat content. Grabowski13 demonstrated that the water
hybrids; SEM, standard error of mean. Parameter L* (color brightness)
can assume values from 0 to 100. Parameters a* (redness) and b* content of meat is inversely proportional to its fat content. The
(yellowness) are tri-chromaticity coordinates and can assume positive shelf-life of meat is determined by its moisture content, as water
and negative values: +a* corresponds to red, −a* to green, +b* to is required for microbial growth. The proportion of water in meat
yellow and − b* to blue. decreases as birds grow older owing to an increase in fat content
and processes that occur in muscles. Chicken breast muscles con-
tain significantly less fat than leg muscles, which is a genetically
Significant (P ≤ 0.05) differences in the protein content of leg determined trait. Low-fat meat products are less acceptable in
muscles were found between groups. Group A and M chickens terms of sensory properties but are lower in calories.14
were characterized by a higher protein content of leg muscles than Consumers evaluate meat quality and make their purchasing
group H birds (Table 1). The water content of leg muscles ranged decisions based on the visual appearance and color of the car-
from 71.87 g per 100 g in group H to 72.80 g per 100 g in group cass. The color of fresh meat is determined, among others, by
M. Hybrid chickens had the lowest fat content of leg muscles, the concentrations of heme pigments and pH. The pH value is
while the highest fat content was found in slow-growing Hubbard correlated with changes in the color of poultry meat, in particular
chickens (Table 1). breast muscles. Meat pH is an indicator of the rate of post mortem
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Figure 1. Color of breast and leg muscles and skin of chickens.

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Meat quality of Ayam Cemani chickens
glycolysis in muscles and affects the eating quality, processing
suitability and shelf-life of meat.
The pH of breast muscles was significantly (P ≤ 0.05) higher
in group H (5.80) than in group A (5.67) (Table 2). According
to Gornowicz et al.,15 chicken breast and leg muscles have pH
levels of 6.0–6.1 and 6.2–6.3 respectively. The noted differences
could result from differences in the analyzed breeds and feeding
regimes. The pH values noted in this experiment are comparable
to those reported by other authors,16,17 in whose studies the pH
of chicken breast and leg muscles reached 5.8–6.0 and 6.1–6.5
respectively.
Meat color is an important indicator of its processing suit-
ability. Consumers’ perception of meat color strongly influences
purchasing decisions. Consumers often use visual evaluation of
color to judge meat freshness and quality.18 Acidic pH levels
(5.9–6.2) are required to achieve and stabilize the desired color
of meat. At higher pH levels (6.3–6.4), less oxymyoglobin is
formed and meat turns dark red, which is a typical color of dark,
firm and dry (DFD) meat. When a greater amount of light is
reflected from the surface of meat with a looser texture, its color
becomes brighter, compared with DFD meat with a much firmer
texture.
A statistical analysis revealed that genotype had a significant
(P < 0.01) effect on the values of color parameters L* , a* and b*
of the breast and leg muscles of chickens (Fig. 1). The values of
color coordinates L* , a* and b* are presented in Table 3. According
̇
to Strzyzewski et al.,19 an increase in the pH of meat causes a
decrease in color lightness, and lower pH is correlated with higher
lightness. This correlation was not observed in the present study.
An analysis of L* , a* and b* values revealed differences in color
lightness between the analyzed muscles. The breast muscles of
Hubbard JA 957 chickens were characterized by the highest value
of L* (51.53), whereas the breast muscles of Ayam Cemani chickens
were darkest in color (35.64). A similar trend was noted in leg
muscles, whose color was lightest in Hubbard (54.17) and darkest
in Ayam Cemani (28.68) chickens, the difference being significant
(P ≤ 0.05). The breast and leg muscles of Hubbard chickens were
characterized by the highest value of parameter a* (redness). In
Ayam Cemani and Ayam Cemani × Sussex chickens the values
of color coordinate a* determined for breast and leg muscles
indicated a high contribution of the green component. Parameter
b* represents the position of color between yellow and blue. The
values of coordinate b* were positive, indicating that the color of
breast and leg muscles was shifted toward yellow. According to
Kirkpinar et al.,20 lower values of L* and higher values of a* and b*
indicate the desired color of broiler chicken carcasses.
Table 4 shows the results of muscle microstructure analysis. Sex
had no effect on the histological profile of breast and leg muscles,
but the values of the analyzed parameters tended to be higher in
males. The size of breast muscle fibers varied among genotypes.
In most cases the cross-section of muscle fibers revealed a parallel
fiber arrangement. The breast and leg muscles of slow-growing
Hubbard JA 957 chickens had the largest (P ≤ 0.01) fiber diameter.
The largest fiber diameter of breast muscles was noted in Hubbard
JA 957 chickens (♀♀ 84.42 and ♂♂ 87.59 μm) and the smallest in
Ayam Cemani chickens (♀♀ 45.06 and ♂♂ 47.08 μm). The same
trend was observed with respect to leg muscles, i.e. the largest
fiber diameter was noted in Hubbard chickens (♀♀ 61.15 and ♂♂
63.14 μm) and the smallest in Ayam Cemani chickens (♀♀ 36.30
and ♂♂ 39.03 μm). The above differences were statistically signif-
icant (P ≤ 0.01). A smaller diameter of muscle fibers is beneficial
for meat juiciness. Muscle fiber diameters were comparable in all
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www.soci.org
Table 4. Diameter of fibers (μm) in muscles of chickens
Breast muscles Leg muscles
Group ♀♀ ♂♂ ♀♀ ♂♂
H 84.42B 87.59B 61.15B 63.14B
A 45.06A 47.08A 36.30A 39.03A
M 48.40A 50.48A 41.07A 43.27A
SEM 4.07 4.20 4.23 4.25
Means in a column with different letters are significantly different at
P ≤ 0.01. H, Hubbard JA 957 chickens; A, Ayam Cemani chickens; M,
Ayam Cemani × Sussex hybrids; SEM, standard error of mean.
groups, with 75–90% of fibers in bundles having diameters similar
to the mean value in a given group.
The histological structure of chicken muscles is affected by geno-
type. An abundant supply of blood vessels and a high connective
tissue content are considered to be indicators of normal muscle
structure and have a beneficial influence on meat quality. Muscle
structure affects the tenderness, water-holding capacity, color and
pH of meat,21 – 23 which are important for both processing and con-
sumption.
The histochemical profile of meat is determined by both genetic
and environmental factors. A fast growth rate of birds, a high
carcass dressing percentage and a high muscle yield contribute
to an increase in muscle fiber size, which often deteriorates meat
quality.24,25 Prentis et al.26 reported that an increase in muscle
weight in chickens reflects an increase in muscle fiber diameter. In
poultry the cross-sectional area of muscle fibers increases with age,
and fast-growing chickens are characterized by a larger diameter
of muscle fibers than slow-growing chickens. According to Brocka
et al.27 and Guernec et al.,28 an increase in muscle fiber diameter
and length can result from intensive selection and changes in the
size and shape of muscle fibers.29 The above increase is also associ-
ated with a higher proportion of giant fibers whose cross-sectional
area is typically four to five times larger than that of normal fibers.30
Skeletal muscle is a heterogeneous tissue consisting of diverse
types of muscle fibers responsible for structural differences
between muscles.31 The characteristics of muscle fibers are
affected by a variety of factors such as sex,32 age,33 breed34 and
physical activity.35 Previous research has shown that outdoor
access can also influence muscle fiber characteristics.36 – 38 The
number and type of muscle fibers are already determined in
prenatal life.39,40 Sobolewska et al.41 demonstrated that in broiler
chickens raised until 35 days of age the most rapid increase in mus-
cle fiber diameter was observed between days 8 and 21. In chick-
ens, breast muscle fibers are fully developed at around 8 weeks
of age.
The effect of muscle fiber characteristics on meat quality has
been studied for years. Muscle fiber characteristics are deter-
mined genetically and vary among breeds. The histological and
biochemical properties of muscles, including the type, number,
proportions, diameter and metabolic profile of fibers, had a signif-
icant effect on the pH and water-binding capacity of meat, thus
affecting its quality.42 The proportions and diameters of different
muscle fiber types affect the sensory properties of meat, which
is an important consideration for consumers. A high number
of small- and average-diameter fibers improves meat quality.43
Increased thickness of white muscle fibers has a positive effect on
meat tenderness and a negative effect on juiciness. A reverse trend
1733
was noted in red muscle fibers.44,45 Chicken breast muscles are
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 www.soci.org
composed almost entirely of white fibers. The number of red
muscle fibers decreases with age and with increasing body
weight.46
In addition to muscle fiber structure, another important trait
of muscle microstructure responsible for meat tenderness is the
number of fibers in bundles. In a study by Scheuermann et al.,47
male broiler chickens were characterized by a denser network of
breast muscle fibers than females. Differences in the number of
muscle fibers and muscle fiber size between males and females
result also from different levels of physical activity. Muscle fiber size
varies among breeds. Burke and Henry48 reported that the number
of myofibers in M. semimembranosus was twofold higher in broiler
strain chickens than in bantam chickens. In a study by Remignon
et al.49,50 the total number of fibers in M. anterior latissimus dorsi
was 15–20% higher in male chickens of the fast-growing line than
in those of the slow-growing line.
A histochemical analysis revealed that the breast muscles
of slow-growing Hubbard JA 957 chickens had a larger fiber
diameter than the breast muscles of Ayam Cemani and Ayam
Cemani × Sussex chickens. A higher proportion of large-diameter
muscle fibers contributed to a significant increase in cooking loss
(P ≤ 0.01). The lowest cooking loss in both breast and leg muscles
was noted in Ayam Cemani chickens (Table 2). The physicochemi-
cal properties of breast and leg muscles of chickens are shown in
Table 2. The pH24 of breast muscles was similar in all groups (ca
5.7) and comparable to that reported by Fiłonik and Niemiec.51
The water-binding capacity of muscle proteins decreases with
decreasing pH levels and increases with increasing pH values. The
muscles of Ayam Cemani chickens were characterized by higher
water-binding capacity compared with the other two groups. The
lowest cooking loss was also noted in group A. Genotype had a
significant (P ≤ 0.01) effect on water-holding capacity, determined
as forced drip loss, which was lower in Ayam Cemani chickens.
Water-holding capacity is a key indicator of meat technological
quality.52
CONCLUSIONS
1 The results of this study show that genotype (Ayam Cemani,
Ayam Cemani × Sussex, Hubbard JA 957) affected the quality
and color of meat and the histological profile of chicken breast
and leg muscles.
2 Meat from Ayam Cemani chickens is a rich source of protein and
could be highly valued by gourmet consumers, connoisseurs
and dieticians for its rarity and originality.
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