Correlations of Sensory Quality Characteristics with Intramuscular Fat Content and Bundle Characteristics in Bovine Longissimus Thoracis Muscle

Young Min Choi1,*, Lyda Guadalupe Garcia2, Kichoon Lee2
Author Information & Copyright
1Department of Animal Sciences, Kyungpook National University, Sangju 37224, Korea
2Department of Animal Sciences, The Ohio State University, Columbus, OH 43210, USA
*Corresponding author : Young Min Choi, Department of Animal Sciences, Kyungpook National University, Sangju 37224, Korea Tel: +82-54-530-1232 Fax: +82-54-530-1229 E-mail:

© Korean Society for Food Science of Animal Resources. This is an Open-Access article distributed under the terms of the Creative Commons Attribution Non-Commercial License ( which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original work is properly cited.

Received: Jan 02, 2019 ; Revised: Feb 12, 2019 ; Accepted: Feb 20, 2019

Published Online: Apr 30, 2019


The aim of this study was to investigate the relation of sensory quality traits of cooked beef to fresh meat quality and histochemical characteristics, especially muscle bundle traits, in the longissimus thoracis muscle of Hanwoo steers. Cooking loss negatively correlated with softness, initial tenderness, chewiness, rate of breakdown (RB), and amount of perceptible residue (AR) after chewing (p<0.05), and drip loss showed negative correlation with RB and AR (p<0.05). All the attributes of tenderness exhibited negative correlation with the Warner-Bratzler shear force value (p<0.05). Marbling score and the intramuscular fat (IMF) content showed positive correlation with all the organoleptic characteristics, including tenderness attributes, juiciness, and flavor (p<0.05). Regarding histochemical characteristics, muscle fiber size did not have a significant correlation with all the sensory quality traits, although the area percentage of type I fiber was related with softness, initial tenderness, and chewiness (p<0.05). On the contrary, the characteristics of muscle bundle were related to all the sensory tenderness attributes (p<0.05), and the sensory tenderness increased with smaller muscle bundle size (p<0.05). These results suggest that the IMF content and bundle characteristics can be used as indicators for explaining the variations in sensory tenderness in well-marbled beef.

Keywords: sensory quality; meat quality; intramuscular fat content; muscle bundle; Hanwoo beef


Beef palatability factors (tenderness, juiciness, and flavor) are considered the most important characteristics that determines consumer satisfaction (Grunert et al., 2004). Sensory quality traits of cooked meat, specifically tenderness, juiciness, and overall acceptability, are influenced by the intramuscular fat (IMF) content (Hocquette et al., 2010; Lee et al., 2018a; Wood et al., 2008). Bovine muscles contain a greater amount of IMF showed tenderer and more juicy compared to bovine muscles contain a low amount of IMF (Hocquette et al., 2010). Thus, most consumers in the US, Japan, and Korea tend to prefer well-marbled beef; thus eating satisfaction is well-associated with the amount of IMF (Cho et al., 2010; Hunt et al., 2014; Hwang et al., 2010), and the minimum content of IMF to achieve acceptable consumer satisfaction of beef in the US is greater than 3%–4% (Savell et al., 1986). In contrast, the European consumers generally prefer the leaner meat due to health concerns (Ngapo et al., 2007). Thus, the consumer acceptability range is wide, primarily due to personal likes/dislikes, culture, religion and family or personal customs. These factors have an impact on eating experiences and purchasing decisions. Owing to these reasons, researches on the effect of IMF percentage on the eating quality traits of cooked beef have generated conflicting results (Hocquette et al., 2010). However, studies have shown that muscle fiber characteristics can impact consumer eating experience by influencing the meat quality characteristics (Choi and Kim, 2009; Rehfeldt et al., 2008).

Muscle fiber characteristics, especially metabolic and morphological properties, could influence the meat quality characteristics and palatability in different species, since the skeletal muscle is primarily composed of muscle fibers (Choi and Kim, 2009; Choi and Oh, 2016). Muscle fibers are classified into various types depending on their contractile and metabolic properties: slow-twitch oxidative (type I), fast-twitch oxido-glycolytic (type IIA) and fast-twitch glycolytic (IIX and IIB) fibers (Schiaffino and Reggiani, 1996). Muscle fibers are grouped together into muscle bundle surrounded by connective tissue, which ensures the muscular contractility in living animals (Schleip et al., 2006). Similar to the fiber number in most terrestrial vertebrates, bundle characteristics of skeletal muscle are also fully developed before or soon after birth with the exception of bundle size (Schleip et al., 2006). With increases in muscle fiber size postnatally, muscle bundle size will also increase. Albrecht et al. (2006) reported no significant difference in the number of fiber per bundle in various cattle breeds aged 2–24 months. In carcasses, muscle bundle characteristics are associated with texture and firmness quality traits evaluated on the exposed muscle cut surface, and thus can influence the sensory quality traits of cooked meat, especially tenderness (Chandraratne et al., 2006; Borgogno et al., 2016). However, the relationship between the muscle bundle, including total bundle number per each muscle, fiber number per each bundle, and bundle size, and organoleptic characteristics of cooked beef remains undefined. The objective of the present study was to determine the correlation of sensory quality traits of cooked beef with fresh meat quality, IMF content and histochemical characteristics of Hanwoo beef.

Materials and Methods

Animals and muscle samples

Eighty-four Hanwoo steers (aged 26–30 months; mean carcass weight of 458.4±71.1 kg) were obtained from a commercial slaughter house in three batches (20, 32, and 32 animals per day). At 45 min postmortem, muscle pH was measured (pH45 min) from the left carcass side using spear-type portable pH meter and probe (IQ-150 pH meter and PH77-Ssprobe, IQ Scientific Instruments Inc., San Diego, CA). Approximately 20 g samples were removed from the longissimus thoracis muscle at the carcass grading site (13th thoracic vertebrae), and were cut into muscle pieces (0.5 cm×0.5 cm×1.0 cm) for the measurement of fiber and bundle characteristics. Muscle samples then were stored at −80°C until further analyses. In addition, after 24 h of chilling at 4°C, a cross-section of the longissimus thoracis muscle was removed between the 10th to 13th thoracic vertebrae (10 to 13 cm thickness) from the left side of each carcass. Muscle samples were used to determine meat and sensory quality characteristics. Meat quality characteristics studied were muscle pH24 h, objective color measurement, water-holding capacity (WHC), and Warner-Bratzler shear force (WBS). Meat quality measurements were performed along with sampling for sensory quality characteristics of cooked meat. A trained sensory panel was used to evaluate various eating quality attributes, and each sample was cut into steak-shaped chops (1.5 cm thick, approximately 100–120 g), frozen and stored at −20°C until further analysis. Overall fat content was determined by the Soxhlet method using a solvent extraction system (AOAC, 2012). Marbling score was assigned according to the beef marbling standard (BMS) (1, devoid; 9, very abundant) as provided by the Korean Institute of Animal Products Quality Evaluation (KAPE, 2017). Loin-eye area at the 13th thoracic vertebrae from left side of each carcass was provided by the KAPE (2017).

Sensory quality evaluation

Each trained panelists were evaluated a total of 84 samples twice (84 samples×2 replicates). The assessments were performed in separated booths during 33 sessions (5–6 samples per session). Twelve sensory panelists (six women and six men; aged 24–43 years) were trained to assess the meat samples for over 1 year. While training the sensory panelists, muscle samples from different locations and marbling scores of Hanwoo beef were used to obtain consistent and precise results. The sensory panelists were trained according to the previous procedure of Meilgaard et al. (1991) and the American Meat Science Association guidelines (AMSA, 1995). All training and testing were conducted in the Muscle Biology Laboratory at the Kyoungpook National University. A total of 168 samples randomly coded with a three-digit number and selected. Steak-size cuts were roasted in a convection oven (MJ324; LG Electronics, Korea); the temperature was set to 180°C. Steak was turned every 3 min until the core temperature reached 71°C, which was measured using a thermometer (Testo 108; Testo Se & Co., Germany) placed in the geometric center of every sample. Cooked steaks then were sliced into 1.3 cm3 cubes, and then kept in a closed polyethylene bag and maintained in a water bath at 54°C until presented to the sensory panelists (Fortin et al., 2005). Unsalted crackers and sufficient water were supplied to trained panelists to rinse and cleanse the plate at the start of a session and between beef samples. Cooked steaks were assessed for 9 traits including softness (S), initial tenderness (IT), chewiness (C), rate of breakdown (RB), amount of perceptible residue (AR), juiciness (J), flavor intensity (FI), off flavor intensity (OFI), and mouth coating (MC) (AMSA, 1995; Meilgaard et al., 1991).

Meat quality measurements

At 45 min and 24 h postmortem, muscle pH was measured in the longissimus thoracis muscle. At 24 h postmortem, color values of fresh beef were evaluated after blooming time (exposing to air for 30 min at 4°C) using a Minolta chromameter (CR-400, Minolta Camera Co., Osaka, Japan). Objective color values are expressed according to the Commission Internationale de l’Eclairage (CIE, 1978) lightness (L*), redness (a*), and yellowness (b*).

Drip loss, filter-paper fluid uptake (FFU), and cooking loss were evaluated for WHC according to the earlier procedures (Kauffman et al., 1986; Honikel 1998). The cooked beef samples for WBS analysis prepared using a method similar to that of the cooking loss method. Beef samples (about 80 g of initial weight) were freshly cut at 24 h postmortem in a 4°C cold room and weighed. Each sample was kept in a closed polyethylene bag and placed in a temperature-controlled water bath at 80°C until the core temperature reached 71°C. Beef samples were then transferred to an ice water bath and chilled for 15 min. A minimum of ten cores were sliced parallel to the longitudinal direction of the muscle fibers per sample (1.27 cm diameter). Cores were then sheared perpendicular to the muscle fiber orientation using an Instron Universal Testing Instrument (Model 1011, Instron Corp., Norwood, USA) with a Warner–Bratzler shear device (cross-head speed 200 mm/min) (AMSA, 1995).

Histochemical analysis

Muscle sections (10 μm) were obtained from each muscle sample using a cryostat (CM1860, Leica, Nussloch, Germany) at −25°C. All histochemical images for measuring the muscle fiber and determining bundle characteristics were taken using an image analysis system consisting of an optical microscope (DM500, Leica Microsystems, Wetzlar, Germany) fitted with a high-resolution digital camera (ICC50, Leica Microsystems, Milton Keynes, England) connected to a LAS EZ software (Leica Microsystems, Heerbrugg, Switzerland) for a standard workstation computer. Image-Pro Plus software (Media Cybernetics, Rockville, USA) were used for the image analysis. The histochemical images for the fiber and bundle characteristics were taken at 100× and 40× magnifications, respectively. For muscle fiber characteristics, a staining method examining myofibrillar ATPase activity was used. The myofibrillar ATPase activity was detected following pre-incubation with acid (pH 4.3) (Lind and Kernell, 1991). Muscle fibers were classified as type I, IIA, or IIX fiber (Brooke and Kaiser, 1970). The mean of muscle fiber cross-sectional area (CSA), mean CSA of each fiber type, and total fiber number were calculated. The area percentage of muscle fiber was obtained as the ratio of total area of each fiber type to the total measured fiber area multiplied with 100. To evaluate the characteristics of muscle bundle, each muscle section was performed with Aniline Blue and Orange G (Albrecht et al., 2006; Krichesky, 1931). The mean size of muscle bundle, mean numbers of fiber per each bundle, and total number of muscle bundle were calculated like those of muscle fiber. More than 30 bundles per sample were used.

Statistical analysis

Data for sensory quality characteristics were analyzed using the SAS software (SAS, 2014) to calculate simple means. All data were checked for normality using a Shapiro-Wilk test, and data for sensory quality traits were found not to be normally distributed. Thus, the Spearman correlation coefficients were calculated using the PROC CORR procedure of SAS software (SAS, 2014) to determine the correlation between sensory quality, meat quality and histochemical characteristics. The principal component analysis (PCA) was performed by PRINCOMP procedure (SAS, 2014) based on the correlation matrix.


Correlation between sensory and meat quality characteristics

Table 1 presents the mean, standard deviation, and overall range (maximum and minimum) for all the sensory quality traits. The variables tested, especially five attributes of tenderness, showed broad ranges. The correlation between sensory and meat quality characteristics are presented in Table 2. Muscle pH at 45 min and 24 h postmortem did not have a significant correlation with any of the sensory quality traits (p>0.05). Lightness of meat was positively correlated with the five tenderness-related attributes (p<0.05), including scores of S (r=0.35), IT (r=0.32), C (r=0.29), RB (r=0.37), and AR (r=0.33). Redness and yellowness were positively correlated with RB score (r=0.23 and 0.27, respectively). However, it did not correlate significantly with J, FI, and MC (p>0.05). Further, drip loss exhibited a negative correlation with the RB (r=−0.29) and AR (r=−0.22). Cooking loss was negatively correlated with all the eating quality characteristics (p<0.05), except the C and FI (p>0.05). The WBS showed a negative correlation with scores for tenderness attributes (p<0.05), whereas the marbling score was positively related with all the sensory quality characteristics including tenderness, juiciness, and flavor (p<0.05). In particular, the content of IMF correlated with MC (r=0.62) and the tenderness attributes including S (r=0.40), IT (r=0.41), C (r=0.46), RB (r=0.33), and AR (r=0.50).

Table 1. Sensory quality traits of the longissimus thoracis muscle (10th to 13th thoracic vertebrae) of the Hanwoo steer
Mean SD Minimum Maximum
Tenderness attribute
 Softness1) 5.56 1.24 2.63 7.75
 Initial tenderness2) 5.40 1.33 2.00 7.80
 Chewiness3) 5.15 1.39 2.00 7.63
 Rate of breakdown4) 5.03 1.11 2.13 7.13
 Amount of perceptible residue5) 5.38 0.87 3.25 7.70
Juiciness6) 5.31 0.88 3.25 7.25
Flavor intensity7) 5.72 0.53 4.50 6.75
Off flavor intensity8) 6.23 0.42 5.25 7.25
Mouth coating9) 5.19 1.11 2.50 7.80

1) Scale: 1, very hard; 9, very soft.

2) Scale: 1, very tough; 9, very tender.

3) Scale: 1, very chewy; 9, very tender.

4) Scale: 1, very slow; 9, very fast.

5) Scale: 1, very abundant; 9, none.

6) Scale: 1, not juicy; 9, extremely juicy.

7) Scale: 1, very weak; 9, very strong.

8) Scale: 1, very strong; 9, very weak.

9) Scale: 1, none; 9, very high.

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Table 2. Correlation coefficient (r) between sensory quality characteristics of cooked beef and meat quality characteristics in the longissimus thoracis muscle of Hanwoo steer
pH45 min pH24 h L* a* b* Drip loss FFU Cooking loss WBS Marbling score IMF content
Tenderness attribute
 Softness1) −0.15 −0.04 0.35** 0.13 0.20 −0.18 −0.08 −0.33** −0.56*** 0.53*** 0.40***
 Initial tenderness2) −0.16 −0.01 0.32** 0.12 0.18 −0.20 −0.06 −0.23* −0.54*** 0.54*** 0.41***
 Chewiness3) −0.14 −0.02 0.29** 0.08 0.12 −0.19 −0.03 −0.21 −0.52*** 0.53*** 0.46***
 Rate of breakdown4) −0.13 −0.01 0.37** 0.23* 0.27* −0.29** −0.23* −0.24* −0.47*** 0.40*** 0.33**
 Amount of perceptible residue5) −0.04 −0.06 0.33** 0.18 0.12 −0.22* −0.15 −0.25* −0.43*** 0.51*** 0.50***
Juiciness6) 0.11 0.08 0.12 −0.02 −0.01 0.09 0.06 −0.29** −0.41*** 0.53*** 0.53***
Flavor intensity7) 0.16 0.04 −0.04 −0.04 −0.05 0.21 0.21 −0.16 −0.28** 0.30** 0.27*
Off flavor intensity8) −0.09 0.01 0.15 0.13 0.12 −0.01 −0.03 −0.40*** −0.27* 0.44*** 0.24*
Mouth coating9) 0.08 0.09 0.19 −0.05 −0.02 0.11 0.13 −0.30** −0.49*** 0.68*** 0.62***

1) Scale: 1, very hard; 9, very soft.

2) Scale: 1, very tough; 9, very tender.

3) Scale: 1, very chewy; 9, very tender.

4) Scale: 1, very slow; 9, very fast.

5) Scale: 1, very abundant; 9, none.

6) Scale: 1, not juicy; 9, extremely juicy.

7) Scale: 1, very weak; 9, very strong.

8) Scale: 1, very strong; 9, very weak.

9) Scale: 1, none; 9, very high.

* p<0.05;

** p<0.01;

*** p<0.001.

FFU, filter-paper fluid uptake; WBS, Warner-Bratzler shear force; IMF, intramuscular fat.

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Relationship between sensory quality and histochemical characteristics

There was no correlation between muscle fiber area and sensory quality of cooked beef (p>0.05; Table 3); however, the total fiber number positively correlated with the scores of AR (r=0.27) and MC (r=0.38). In the composition of fiber type, the area percentage of type I fiber showed positive correlation with S (r=0.23), IT (r=0.24), C (r=0.24) and J (r=0.23) scores, whereas the percentage of type IIX fiber negatively correlated with the score for softness (r=−0.22). However, no significant correlation was observed between type IIA percentage and sensory quality traits (p>0.05). Table 4 presents the relationships between the sensory quality traits and muscle bundle characteristics. Bundle size and fiber number per bundle exhibited similar correlation tendencies. They correlated negatively with the scores of S (r=−0.43 and −0.38, respectively), IT (r=−0.41 and −0.36, respectively), C (r=−0.44 and −0.38, respectively), RB (r=−0.44 and −0.39, respectively), and AR (r=−0.30 and −0.27, respectively). Total bundle number positively correlated with all the tenderness attributes (p<0.05). On the contrary, no significant correlation was observed between the characteristics of muscle bundle and all the traits (p>0.05), except for the tenderness attributes (p<0.05).

Table 3. Correlation coefficient (r) between sensory quality characteristics of cooked beef and muscle fiber characteristics in the longissimus thoracis muscle of Hanwoo steer
Muscle fiber area Total fiber number Fiber area percentage
Mean Type I Type IIA Type IIX Sum Type I Type IIA Type IIX Type I Type IIA Type IIX
Tenderness attribute
 Softness1) −0.03 0.08 0.03 0.01 0.18 0.21 0.13 0.04 0.23* 0.08 −0.19
 Initial tenderness2) 0.01 0.10 0.04 0.02 0.19 0.21 0.13 0.05 0.24* 0.05 −0.17
 Chewiness3) 0.01 0.09 0.02 0.02 0.19 0.20 0.14 0.05 0.24* 0.04 −0.16
 Rate of breakdown4) −0.04 0.07 0.02 −0.04 0.21 0.19 0.20 0.09 0.15 0.11 −0.14
 Amount of perceptible residue5) −0.07 −0.02 0.01 −0.04 0.27* 0.27* 0.20 0.12 0.13 0.08 −0.11
Juiciness6) 0.05 0.03 0.07 0.10 0.10 0.18 0.08 −0.01 0.23* 0.05 −0.13
Flavor intensity7) 0.06 0.09 −0.02 0.09 0.01 0.12 0.05 −0.04 0.20 −0.02 −0.04
Off flavor intensity8) −0.09 0.02 −0.05 −0.06 0.21 0.21 0.20 0.15 0.09 0.03 −0.08
Mouth coating9) −0.06 0.02 −0.07 0.01 0.24* 0.38** 0.16 0.11 0.21 −0.03 −0.07

1) Scale: 1, very hard; 9, very soft.

2) Scale: 1, very tough; 9, very tender.

3) Scale: 1, very chewy; 9, very tender.

4) Scale: 1, very slow; 9, very fast.

5) Scale: 1, very abundant; 9, none.

6) Scale: 1, not juicy; 9, extremely juicy.

7) Scale: 1, very weak; 9, very strong.

8) Scale: 1, very strong; 9, very weak.

9) Scale: 1, none; 9, very high.

* p<0.05;

** p<0.01.

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Table 4. Correlation coefficient (r) between the sensory quality characteristics of cooked beef and muscle bundle characteristics in the longissimus thoracis muscle of Hanwoo steer
Muscle bundle characteristics
Bundle area Fiber number per bundle Total bundle number
Tenderness attribute
 Softness1) −0.43*** −0.38** 0.50***
 Initial tenderness2) −0.41*** −0.36** 0.50***
 Chewiness3) −0.44*** −0.38** 0.51***
 Rate of breakdown4) −0.44*** −0.39*** 0.50***
 Amount of perceptible residue5) −0.30** −0.27* 0.39***
Juiciness6) −0.17 −0.15 0.24*
Flavor intensity7) 0.14 −0.13 0.10
Off flavor intensity8) −0.05 −0.06 0.18
Mouth coating9) 0.18 −0.13 0.27*

1) Scale: 1, very hard; 9, very soft.

2) Scale: 1, very tough; 9, very tender.

3) Scale: 1, very chewy; 9, very tender.

4) Scale: 1, very slow; 9, very fast.

5) Scale: 1, very abundant; 9, none.

6) Scale: 1, not juicy; 9, extremely juicy.

7) Scale: 1, very weak; 9, very strong.

8) Scale: 1, very strong; 9, very weak.

9) Scale: 1, none; 9, very high.

* p<0.05;

** p<0.01

*** p<0.001.

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PCA of the experiments

The PCA was performed to understand the correlation between the variables, including sensory quality traits of cooked beef, meat quality and histochemical characteristics (Fig. 1). The first three of these PCs accounted for 55.6% of the variance observed in the 34 variables (PC1=30.8%, PC2=14.5% and PC3=10.3%). All sensory quality traits were distinguishable by the positive loading of PC1 and PC2 (Fig. 1A). In Fig. 1B, initial tenderness, chewiness and AR were distributed in the area with negative loading of PC3. All tenderness attributes were located close to one another indicating that they were positively correlated with each. The IMF content, marbling, and total number of bundle were distinguishable by the positive loading of PC1, while WBS, bundle size, and fiber number per bundle showed the negative loading of PC1. However, IMF content, bundle size, and fiber number per bundle were distributed in the area with negative loading of PC2, but positive loading of PC3.

Fig. 1. Principal component analysis (PCA) plots for PC1 versus PC2 (A) and PC1 versus PC3 (B). The percentage variance explained by the three PCs was 55.6%. S, softness; IT, initial tenderness; C, chewiness; RB, rate of breakdown; AR, amount of perceptible residue; J, juiciness; FI, flavor intensity; OFI, Off flavor intensity; MC, mouth coating; DL, drip loss; FFU, filter-paper fluid uptake; CL, cooking loss; WBS, Warner-Bratzler shear force; MS, marbling score; IMF, intramuscular fat content; CSA, muscle fiber cross-sectional area; CSA-I, type I fiber CSA; CSA-IIA, type IIA fiber CSA; CSA-IIX, type IIX fiber CSA; TFN, total fiber number; TFN-I, TFN of type I fiber; TFN-IIA, TFN of type IIA fiber; TFN-IIX, TFN of type IIX fiber; AP-I, area percentage of type I fiber; AP-IIA, AP of type IIA fiber; AP-IIX, AP of type IIX fiber; BA, bundle area; FNB, fiber number per bundle; TBN, total bundle number.
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Eating quality characteristics of cooked beef are associated with the quality characteristics of fresh meat. Muscle pH at the early postmortem period is an important indicator of the postmortem glycolytic rate, and the ultimate pH can influence the meat tenderness (Marsh et al., 1987). Lomiwes et al. (2013), showed that beef with a higher ultimate pH (≥C6.2) showed higher scores for sensory tenderness and lower values of WBS than beef with normal (6.19 to 5.8) or lower pH (≤5.79), which might be attributable to the extent of myofibril structural degradation by endoprotease, and activities of endoprotease were greater at higher pH due to their isoelectric point (Pulford et al., 2008). However, in the present study, no significant correlation was observed between sensory quality traits and muscle pH, a discrepancy which might arise due to the curvilinear relationship between the ultimate pH and sensory tenderness (Silva et al., 1999). Furthermore, muscle pH can influence the development of meat color (Jeong et al., 2010). In the current study, muscle pH45 min was significantly correlated with lightness (r=−0.23; data not shown), and values of softness (r=0.34), initial tenderness (r=0.31) and AR (r=0.29) were positively correlated with lightness. Generally, the WHC of meat is positively correlated with tenderness. Gil et al. (2008) reported that drip loss is correlated with hardness and juiciness. Porcine muscles with lower drip loss were previously shown to exhibit lower values of WBS compared to those with higher drip loss (Jeong et al., 2010). Results from the present study corroborate with this observation; low cooking loss was associated with tender and juicy beef. As expected, the WBS positively correlated with the tenderness attributes as well as juiciness and flavor intensity (p<0.05).

The extent of marbling plays a key role in determining organoleptic characteristics (Hocquette et al., 2010; Lee et al., 2019). Similar to Japanese Black cattle (Albrecht et al., 2011), Hanwoo cattle are genetically predisposed to depositing a higher content of IMF than German Angus and Galloway (Albrecht et al., 2006; Cho et al., 2010). The IMF content of the longissimus thoracis muscle at the 13th thoracic vertebra ranges from approximately 5% to 21% with marbling scores from 1 to 9 (Cho et al., 2010; KAPE 2017). In the present study, the IMF content and marbling score of the longissimus thoracis muscle ranged from 6.63% to 22.8% and 2 to 9, respectively (data not shown). The tenderness attributes of cooked beef correlated with the IMF content and marbling score (p<0.05). A similar result has been reported by Nishimura (2015), who suggested that the IMF content was strongly related to tenderness evaluated by sensory panel in Japanese Black steer, as the consumers in Japan preferred a well-marbled beef.

It is well known that type I fiber is predominantly energized by the oxidative metabolism, and therefore lipids and fatty acids are mainly used as sources of metabolic fuel in living animals (Schiaffino and Reggiani, 1996). However, in the present study, no significant correlations were found between area percentage of type I fiber and IMF content in this study (p>0.05), as a higher IMF content is achieved by long-term feedlot programs with high energy in well-marbled beef (Cho et al., 2010; Hocquette et al., 2010). On the other hand, the metabolic and contractile characteristics of muscle are associated with sensory quality, especially meat tenderness (Choi et al., 2013; Picard et al., 2014). According to Jeong et al. (2010), the fiber mean CSA and type I fiber CSA were positively correlated with sensory tenderness of cooked pork. Several studies have investigated the effects of fiber characteristics on sensory quality traits of cooked beef. However, collectively, researchers are divided on whether fiber characteristics affect the sensory quality traits of cooked beef. For instance, some studies showed a negative correlation between fiber area and tenderness of cooked beef (Seideman et al., 1986), while others reported no correlation between these two attributes (Vestergaard et al., 2000; Wegner et al., 2000). In the present study, a limited correlation was observed between the morphological characteristics of muscle fiber and the sensory quality traits of the cooked beef. On the contrary, the tenderness attributes, except the RB and AR, were positively correlated with the area percentage of type I fiber (p<0.05).

A muscle bundle is a group of muscle fibers enveloped by connective tissue layer (known as the perimysium) and can be classified into primary and secondary bundles (Purslow, 2005). Muscle fascicle and perimysium maintain passive stiffness of the muscle for accurate contraction and prevent overstretching of muscle fibers in living animal (Schleip et al., 2006). Thus, muscle bundle area is related to the graininess of muscle surface, which can be observed in carcasses (Lee et al., 2018b; Purslow, 2005). Although the bundle CSA increases with in advanced maturity, significant differences in bundle characteristics, including bundle CSA, have been observed in various cattle breeds of similar age (Albrecht et al., 2006). Cooper et al. (1968) reported that the bundle area positively correlated with WBS value (r=0.39, p<0.01), and negatively with the sensory tenderness (r=−0.30, p<0.05) in the bovine longissimus muscle, in accordance with the results of the present study. Muscles with s smaller bundle area and lower fiber number per bundle exhibited lower initial force to penetrate beef, easily to break into fragments, and leave lower amount of residue compared to muscles with larger bundle area and higher fiber number per bundle (p<0.05).

In the PCA, the first seven and ten PCs were extracted explaining 81.5% and 91.9% of the total variance (data not shown). The first two and three PCs described 45.3% and 55.6% of the total variance in the 34 variables, as there were many variables in this study. Mwove et al. (2018) extracted the first seven PCs explaining 95.5% of the total variance in the 24 beef quality characteristics, including sensory quality, WBS and proximate composition, and Liu et al. (2004) reported that the first two and seven PCs accounted for 36.8% and 69.2%, respectively, of the total variance for 24 variables including sensory quality traits, color and physical characteristics of chicken breast. In the current study, sensory quality characteristics, especially tenderness attributes, were positively correlated with marbling score, IMF content and total bundle number. On the other hand, tenderness attributes were negatively correlated with WBS, total bundle number and bundle area, which were located on the opposite quadrant.


The present study found that bovine muscles with smaller bundle size exhibited more tender after initial compression and less chewing force than bovine muscles with larger bundle size. As expected, the IMF content showed a positive effect on tenderness attributes. Therefore, the characteristics of muscle bundle can influence the tenderness attributes of cooked beef as the IMF content.


Conflict of Interest

The authors declare no potential conflict of interest.


This research was supported by the National Research Foundation of Korea (NRF-2017R1D1A3B03029840).


Author Contributions

Conceptualization: Choi YM. Data curation: Choi YM, Garcia LG, Lee K. Formal analysis: Choi YM. Methodology: Choi YM. Software: Choi YM. Validation: Choi YM, Garcia LG, Lee K. Investigation: Choi YM. Writing - original draft: Choi YM, Garcia LG, Lee K. Writing - review & editing: Choi YM, Garcia LG, Lee K.


Ethics Approval

This article does not require IRB/IACUC approval because there are no human and animal participants.



Albrecht E, Gotoh T, Ebara F, Xu JX, Viergutz T, Nurnberg G, Maak S, Wegner J. 2011; Cellular conditions for intramuscular fat deposition in Japanese Black and Holstein steers. Meat Sci. 89:13-20


Albrecht E, Teuscher F, Ender K, Wegner J. 2006; Growth- and breed-related changes of muscle bundle structure in cattle. J Anim Sci. 84:2959-2964


American Meat Science Association. 1995 Research guidelines for cookery, sensory evaluation, and instrumental tenderness measurements of fresh meat. American Meat Science Association. Chicago, IL, USA: .


AOAC. 2012 Official methods of analysis of AOAC international. 19th edAOAC International. Gaithersburg, MD, USA: .


Borgogno M, Sacca E, Corazzin M, Favotto S, Bovolenta S, Piasentier E. 2016; Eating quality prediction of beef from Italian Simmental cattle based on experts’ steak assessment. Meat Sci. 118:1-7


Brooke MH, Kaiser KK. 1970; Three “myosin adenosine triphosphatase” systems: The nature of their pH lability and sulfhydryl dependence. J Histochem Cytochem. 18:670-672


Chandraratne MR, Samarasinghe S, Kulasiri D, Bickerstaffe R. 2006; Prediction of lamb tenderness using image surface texture features. J Food Eng. 77:492-499


Cho SH, Kim J, Park BY, Seong PN, Kang GH, Kim JH, Jung SG, Im SK, Kim DH. 2010; Assessment of meat quality properties and development of a palatability prediction model for Korean Hanwoo steer beef. Meat Sci. 86:236-242


Choi YM, Kim BC. 2009; Muscle fiber characteristics, myofibrillar protein isoforms, and meat quality. Livest Sci. 122:105-118


Choi YM, Nam KW, Choe JH, Ryu YC, Wick MP, Lee K, Kim BC. 2013; Growth, carcass, fiber type, and meat quality characteristics in Large White pigs with different live weights. Livest Sci. 155:123-129


Choi YM, Oh HK. 2016; Carcass performance, muscle fiber, meat quality, and sensory quality characteristics of crossbred pigs with different live weights. Korean J Food Sci Anim Resour. 36:389-396


Commission Internationale de l’Eclairage (CIE). 1978 Recommendations on uniform color spaces – Color differences equations, psychrometic color terms, Supplement 2 to CIE publication 15 (E1.3.1) 1971/(TC1.3). Central Bureau of the Commission Internationale de l’Eclairage. Vienna, Austria: .


Cooper CC, Breidenstein BB, Cassens RG, Evans G, Bray RW. 1968; Influence of marbling and maturity on the palatability of beef muscle. II. Histological considerations. J Anim Sci. 27:1542-1546


Fortin A, Robertson WM, Tong AKW. 2005; The eating quality of Canadian pork and its relationship with intramuscular fat. Meat Sci. 69:297-305


Gil M, Delday MI, Gispert M, Furnols MF, Maltin CM, Plastow GS, Klont R, Sosnicki AA, Carrion D. 2008; Relationships between biochemical characteristics and meat quality of longissimus thoracis and semimembranosus muscles in five porcine lines. Meat Sci. 80:927-933


Grunert KG, Bredahl L, Brunso K. 2004; Consumer perception of meat quality and implications for product development in the meat sector-A review. Meat Sci. 66:259-272


Hocquette JF, Gondret F, Baeza E, Medale F, Jurie C, Pethick DW. 2010; Intramuscular fat content in meat-producing animals: Development, genetic and nutritional control, and identification of putative markers. Animal. 4:303-319


Honikel KO. 1998; Reference methods for the assessment of physical characteristics of meat. Meat Sci. 49:447-457


Hunt MR, Garmyn AJ, O’Quinn TG, Corbin CH, Legako JF, Rathmann RJ, Brooks JC, Miller MF. 2014; Consumer assessment of beef palatability from four beef muscles from USDA choice and select graded carcasses. Meat Sci. 98:1-8


Hwang YH, Kim GD, Jeong JY, Hur SJ, Joo ST. 2010; The relationship between muscle fiber characteristics and meat quality traits of highly marbled Hanwoo (Korean native cattle) steers. Meat Sci. 86:456-461


Jeong DW, Choi YM, Lee SH, Choe JH, Hong KC, Park HC, Kim BC. 2010; Correlations of trained panel sensory values of cooked pork with fatty acid composition, muscle fiber type, and pork quality characteristics in Berkshire pigs. Meat Sci. 86:607-615


Kauffman RG, Eikelenboom G, van der Wal PG, Merkus G, Zaar M. 1986; The use of filter paper to estimate drip loss of porcine musculature. Meat Sci. 18:191-200


Korea Institute of Animal Products Quality Evaluation [KAPE]. The beef carcass grading. Available from: at Feb 1, 2017


Krichesky B. 1931; A modification of Mallory’s triple stain. Stain Technol. 6:97-98


Lee B, Yoon S, Lee Y, Oh E, Yun YK, Kim BD, Kuchida K, Oh HK, Choe J, Choi YM. 2018a; Comparison of marbling fleck characteristics and objective tenderness parameters with different marbling coarseness within longissimus thoracis muscle of high-marbled Hanwoo steer. Korean J Food Sci Anim Resour. 38:606-614.


Lee B, Yoon S, Choi YM. 2019; Comparison of marbling fleck characteristics between beef marbling grades and its effect on sensory quality characteristics in high-marbled Hanwoo steer. Meat Sci. 152:109-115


Lee Y, Lee B, Kim HK, Yun YK, Kang SJ, Kim KT, Kim BD, Kim EJ, Choi YM. 2018b; Sensory quality characteristics with different beef quality grades and surface texture features assessed by dented area and firmness, and the relation to muscle fiber and bundle characteristics. Meat Sci. 145:195-201


Lind A, Kernell D. 1991; Myofibrillar ATPase histochemistry of rat skeletal muscles: A “two-dimensional” quantitative approach. J Histochem Cytochem. 39:589-597


Liu Y, Lyon BG, Windham WR, Lyon CE, Savage EM. 2004; Principal component analysis of physical, color, and sensory characteristics of chicken breasts deboned at two, four, six, and twenty-four hours postmortem. Poult Sci. 83:101-108


Lomiwes D, Farouk MM, Frost DA, Dobbie PM, Young OA. 2013; Small heat shock proteins and toughness in intermediate pHu beef. Meat Sci. 95:472-479


Marsh BB, Ringkob TP, Russell RL, Swartz DR, Pagel LA. 1987; Effects of early-postmortem glycolytic rate on beef tenderness. Meat Sci. 21:241-248


Meilgaard M, Civille GV, Carr BT. 1991 Sensory evaluation techniques. 3rd edCRC Press Inc. Boca Ranton, FL, USA: .


Mwove JK, Gogo LA, Chikamai BN, Omwamba M, Mahungu SM. 2018; Principal component analysis of physicochemical and sensory characteristics of beef rounds extended with gum Arabic from Acacia senegal var. kerensis. Food Sci Nutr. 6:474-482


Ngapo TM, Martin JF, Dransfield E. 2007; International preferences for pork appearance: II. Factors influencing consumer choice. Food Qual Prefer. 18:139-151


Nishimura T. 2015; Role of extracellular matrix in development of skeletal muscle and postmortem aging of meat. Meat Sci. 109:48-55


Picard B, Gagaoua M, Micol D, Cassar-Malek I, Hocquett JF, Terlouw CEM. 2014; Inverse relationships between biomarkers and beef tenderness according to contractile and metabolic properties of the muscle. J Agric Food Chem. 62:9808-9818


Pulford DJ, Fraga Vazques S, Frost DF, Fraser-Smith E, Dobbie P, Rosenvold K. 2008; The intracellular distribution of small heat shock proteins in post-mortem beef is determined by ultimate pH. Meat Sci. 79:623-630


Purslow PP. 2005; Intramuscular connective tissue and its role in meat quality. Meat Sci. 70:435-447


Rehfeldt C, Henning M, Fiedler I. 2008; Consequences of pig domestication for skeletal muscle growth and cellularity. Livest Sci. 116:30-41


SAS. 2014 SAS/STAT software for PC. Release 9.4. SAS Institute Inc. Cary, NC, USA: .


Savell JW, Cross HR, Smith GC. 1986; Percentage ether extractable fat and moisture content of beef longissimus muscle as related to USDA marbling score. J Food Sci. 51:838-839


Schiaffino S, Reggiani C. 1996; Molecular diversity of myofibrillar proteins: Gene regulation and functional significance. Physiol Rev. 76:371-423


Schleip R, Naylor IL, Ursu D, Melzer W, Zorn A, Wilke HJ, Horn FL, Klingler W. 2006; Passive muscle stiffness may be influenced by active contractility of intramuscular connective tissue. Med Hypotheses. 66:66-71


Seideman SC, Crouse JD, Cross HR. 1986; The effect of sex condition and growth implants on bovine muscle fiber characteristics. Meat Sci. 17:79-95


Silva JA, Patarata L, Martins C. 1999; Influence of ultimate pH on bovine meat tenderness during ageing. Meat Sci. 52:453-459


Vestergaard M, Oksbjerg N, Henckel P. 2000; Influence of feeding intensity, grazing and finishing feeding on muscle fibre characteristics and meat colour of semitendinosus, longissimus dorsi and supraspinatus muscles of young bulls. Meat Sci. 54:177-185


Wegner J, Albrecht E, Fiedler I, Teuscher F, Papstein HJ, Ender K. 2000; Growth- and breed-related changes of muscle fiber characteristics in cattle. J Anim Sci. 78:1485-1496


Wood JD, Enser M, Fisher AV, Nute GR, Sheard PR, Richardson RI, Hughes SI, Whittington FM. 2008; Fat deposition, fatty acid composition and meat quality: A review. Meat Sci. 78:343-358

Journal Title Change

We announce that the title of our journal and related information were changed as below from January, 2019.


Before (~2018.12)

After (2019.01~)

Journal Title

Korean Journal for Food Science of Animal Resources

Food Science of Animal Resources

Journal Abbreviation

Korean J. Food Sci. An.

Food Sci. Anim. Resour.







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