Bruising in Slaughter Cattle and Its Relationship with Creatine Kinase Levels and Beef Quality as Affected by Animal Related Factors

Article information

Asian-Australas J Anim Sci. 2014;27(5):717-725
*Corresponding Author: Voster Muchenje. Tel: +27-40-602-2059, Fax: +27-86-628-2967, E-mail:

Department of Livestock and Pasture Science, University of Fort Hare, P. Bag X1314, Alice, Eastern Cape, South Africa

Received 2013 August 08; Revised 2013 October 14; Accepted 2013 November 23.


The objective of the study was to determine the effects of animal related factors on bruising in slaughter cattle, creatine kinase (CK) and beef quality. Three hundred and twenty one cattle from three breeds (108 Bonsmara, 130 Beefmaster and 83 Brahman) were used in this study. The animals were grouped as follows: Group 1 (16 months old), Group 2 (18 months old) and Group 3 (24 months old). At exsanguinations, blood samples for CK determination were collected using disposable vacutainer tubes. Muscularis longissimus thoracis et lumborum (LTL) was collected 24 h after slaughter to determine the colour (L*, a*, and b*) and ultimate pH (pHu) of beef. Breed, sex and age had significant effects (p<0.05) on bruising score, CK levels and beef quality. Bonsmara breed had the highest (80%) bruising score percentage, CK (705.3±80.57 U/L) and pHu (6.3±0.05) values while the Bonsmara had the highest L* (24.8±0.78) a* (17.5±0.53) and b* (12.8±0.53) values. Higher CK levels were also observed in winter compared to summer, spring and autumn respectively. Therefore, animal factors (sex, breed and animal age at slaughter) contribute to the development of bruises and have an effect on the levels of CK and meat quality. It was also concluded that there is no significant relationship between meat parameters (L,* a*, and b*) and CK levels.


Transportation is an important activity of the farming industry. During transportation, beef cattle are grouped by their age, size and breed which are described as intrinsic factors. Grouping of these animals is done by the farm and abattoir workers. This process involves rounding up, kraaling, loading, transportation and off-loading at the abattoir (Bonney, 2006; Chulayo et al., 2012). Transportation is associated with numerous risks which include management, intrinsic and extrinsic factors; of these, intrinsic factors have been found to significantly affect the level of bruising and the quality of meat and carcass characteristics (Hoffman and Lühl, 2012). Bruising in slaughter cattle is described as a superficial discoloration due to haemorrhage of tissue from ruptured blood vessels beneath the surface of the skin (Strappini et al., 2010; Vimiso, 2010; Vimiso and Muchenje, 2013). In the contusions, the blood accumulates in the surrounding tissues, producing pain and swelling (Blood and Studdert, 1988). Bruising occurs at the farm, during transportation and at the slaughterhouse. It is speculated that sex of animals, gender and breed has an effect on animal stress. Male and young animals are at risk of more bruising than female and old animals, due to rough driving and mishandling before and after transportation (Parrot et al., 1994; Jarvis et al., 1995; Averos et al., 2008).

Cattle are prone to repeated rough handling in their lifetime which later on lead to aversive reactions such as stress and bruising immediately prior to transportation (EFSA, 2011). Stressed and bruising in animals results in an increase in the activity of creatine kinase in the blood and this is due to tissue damage and poor muscular tissue reperfusion (Knowles et al., 1999; Vojtic, 2000). In addition, capture, loading and transportation induces permeability of the muscle membrane hence increased creatine kinase (CK) levels in the blood (Warriss et al., 1995; Lopez et al., 2006; Guardia et al., 2009). CK is found in the skeletal muscles of animals and it is responsible for maintaining energy homeostasis at the sites of high adenosine tri-phosphate (ATP) (Diene and Storey, 2009; Chulayo and Muchenje, 2013). This enzyme is generally utilized as an indicator of physical stress and/or muscle damage in animal production. In goats, higher levels of CK in the plasma may also be caused by breed temperament, excitability and fighting against each other (Grzyb and Skorkowski, 2005).

The aforementioned factors, especially transportation, are documented to likely have a greater adverse effect on the welfare of calves than on mature (>17 to 19 months) age animals (Zhang et al., 2005). Grandin (2001) indicated that calf welfare is mostly affected when they are exposed to a multitude of novel and stressful events during handling and transportation. This is also witnessed by the increased incidence of mortality once they arrive at the slaughterhouse (Fike and Spire, 2006). Moreover, old animals experience similar welfare challenges which later reduce carcass quality and meat quality resulting in low economic value (Grandin, 2001). The adverse effects of transportation on welfare vary by animal type. For example, mature cattle (>500 kg) such as transported for about 400 km for slaughter had less shrinkage, deaths and bruising compared to young animals (Grandin, 2007). Therefore, poor beef quality may be influenced by the activity undertaken by the animal prior to slaughter. Beefmaster, Bonsmara, Brahman and Nguni are breeds that can withstand adverse environmental conditions (hot temperatures), long dry periods, diseases and parasites (Beatty et al., 2006; Ndlovu et al., 2008; Marufu et al., 2011). Breed affects meat quality during transportation. Aggressive breeds are more prone to bruising, injuries resulting in carcass damage and poorer meat quality (Voisinet et al., 1997).

Several studies indicate that stress lowers post mortem ultimate pH (pHu) values due to depletion of glycogen in ruminants and produces tougher meat. On the other hand pre-slaughter stress results in higher pHu and darker cuts (Devine et al., 2006; Gregory, 2010). However, there is little or no information on the relationship between creatine kinase (CK) and meat quality attributes (ultimate pH and colour). The effects of animal handling and welfare during transportation of animals from farms to commercial abattoirs have been investigated (Gispert et al., 2000; van de Water et al., 2003; Liste et al., 2008; Huertas et al., 2010). In addition, studies (Martinezo-Cerezo et al., 2005; Muchenje et al., 2009) have been conducted to determine the occurrence of bruises in cattle from the farm to the abattoir however, the level of bruising and the relation to CK values was not considered. Therefore the objective of the study was to determine the effects of animal related factors on bruising in slaughter cattle, levels of creatine kinase (CK) and the quality of beef.


Description of the study site

The study was conducted in East London Abattoir which is located in Buffalo City Municipality. The abattoir is situated at longitude and latitude 32.′2° S and 27.′ 5° E in the Amathole District of the Eastern Cape Province of Republic of South Africa (RSA). The area has vegetation that ranges from grassland and thicket to forests and bush veld with Themeda trianda and Digitaria eriantha being the most dominant grass species. The area receives approximately 850 mm of rainfall per year of which most falls during the summer months (November, December and January). The day temperature during for the period of study ranged from 10°C to a high of 36°C, averaging about 22.3°C. The topography of the area is naturally flat with few steep slopes.

Animal management

Data was collected from 321 cattle over four seasons (winter, spring, summer and autumn) between the year 2011 and 2012. The seasons were categorised into the months as follows: winter (June to August), spring (September to October), summer (November to February) and autumn (March to May). All cattle were slaughtered at the East London commercial abattoir in the Eastern Cape Province. This abattoir operates from 0600 to 1700 h on Mondays till Fridays. On arrival at the abattoir the three breeds of cattle; Bonsmara, Brahman and Beefmaster were identified according records from the farmers. The numbers of animals per breed were 108 Bonsmara, 130 Beefmaster and 83 Brahman respectively. The animals used were both male and females and grouped according to age categories as follows: Group 1 (16 months), Group 2 (18 months) and Group 3 (24 months).

The cattle were transported by the owners from the farms to the abattoir. Transportation time (departure, arrival time and distance travelled), lairage duration, number of animals per pen, slaughtered animals per day and daily temperature were all recorded. On arrival at the abattoir, 27 offloading activities were monitored and cattle were given water ad-libitum prior to slaughter. Cattle were then slaughtered using commercial standard procedures of the abattoir. The captive bolt method was used to stun the animals, using a voltage of 300 V, a current of 5 A for 40 to 45 s at a pulse 12 beats/s.

Blood collection and plasma separation

The blood samples were collected from the animals using labelled disposable vacutainer tubes. The blood samples were placed in ice packs prior to creatine kinase (CK) analysis. A centrifuge machine (model: 5403 Centrifuge Geratebay Eppendorf GmbH, Engelsdorp Germany) was used to centrifuge blood samples at 3,550 rpm, 21°C for 10 min. Plasma was then separated and inserted into 1.5 mL Eppendorf tubes using pipettes and stored at −20°C.

Creatine kinase level determination

The serum samples were used to determine the levels of CK at the National Health Laboratory Services (NHLS), Port Elizabeth using Model DXC 600 machine (Beckman Coulter, Ireland). The contents included 2×61 ML CK Reagent, 1 preparation insert and the reactive ingredients included creatine phosphate, disodium Salt 461 mmol/L, nadide 30: 0 mmol/L, adenosine-5-diphosphate, monopotassium salt, dihydrate 36.0 mmol/L, glucose 24.0 mmol/L, glucose-6-phosphate dehydrogenase 46.1 kU/L and hexokinase 136 kU/L. All the ingredients were added for quantitative determination of CK activity of units per litre (U/L) in plasma.

Bruise measurements

Only carcasses that had bruises were considered. The assessment was done using a method based on the Australian Carcass Bruise Score System (Anderson and Horder, 1979). Bruise scores were calculated for each load by multiplying the number of bruises in each size class by a weighting factor: slight 1, medium 3, and heavy 5, and adding these values (Anderson and Horder, 1979). The bruise scores were then divided by the number of cattle per load to give a mean bruise score per animal for each load. Bruise age was estimated using the method of Gracey et al. (1999) as shown in Table 1.

Colour observations used to estimate the age of bruises

Meat quality measurements

The Muscularis longisimus thoracis et Lumborum (LTL) were used for ultimate pH and colour (L*, a*, and b*) measurements. The muscle was removed by cutting a sample between the 4th and 6th ribs of the loin region while the carcass was still hanging, prior to vacuum-packing and storing in a cooler box an hour after collection and further storing in the refrigerator at −4°C.

Colour measurement

Colour of the meat (L* = lightness, a* = redness and b* = yellowness) for the Bonsmara, Beefmaster and Brahman carcasses was determined from the Muscular longissimus thoracis et. Lumborum (LTL) 24 h after slaughter using a colour-guide 45/0 BYK-Gardener GmbH machine with a 20 mm diameter measurement area and illuminant D65-day light, 100 standard observer. Three readings were taken by rotating the Colour Guide 900 machine between each measurement, in order to obtain a representative average value of the colour. This guide was calibrated each day before taking measurements using the green standard.

Determination of meat pH

Meat pH 24 from Bonsmara, Beefmaster and Brahman of carcasses was measured from the Muscularis longissimus thoracis et lumborum muscle at the level of the 10th rib 24 h after slaughter on carcasses that were stored at 0 to 3°C. The measurement was carried out using a portable pH meter (CRISON pH 25, CRISON Instruments, SA Spain). The pH meter was calibrated using pH 4, pH 7 and pH 9 standard solutions (CRISON Instrument, SA Spain) before each day’s measurement. The measurements were then performed with a sharpened metal sheath to prevent probe breakage and meat contamination.

Statistical analysis

The PROC GLM of Statistical Analysis System (SAS) Institute (2003) was used to analyse the effect of breed, age, gender and season on the activity of creatine kinase and bruising score. The model used was as follows:



  • Yijkl = response variable (creatine kinase, bruising score and bruise age)

  • μ = constant mean common to all observations

  • αi = effect of breed

  • βj = effect of age group (Group 1, 2, and 3)

  • λk = effect of season, Gl = effect of gender and ɛijklm = random error

Significant differences between least square group means for CK and animal factors were compared using the Duncan’s procedure. Pair-wise comparison of LS means for creatine kinase concentration, sex, breed and animal age at slaughter were done using PDIFF.

The effect of breed, age group and season on CK, pHu, L,* a*, and b*, were analysed using Proc GLM of SAS (2003). The stastical model used as follows:



  • Yijkl = Response variables (CK, pHu, L, a, b and WBSF)

  • μ = Overall mean

  • αi = breed effect (Bonsmara, Beefmaster and Brahman)

  • βj = age group effect (Group 1, 2, and 3)

  • λk = seasonal effect (summer, winter, spring and autumn)

  • ɛijkl = random error term

The significant difference between least square group means for CK levels and beef quality parameters were compared using Duncan multiple range procedure. Correlations were computed using PROC GLM of Statistical Analysis System (SAS) (2003) to determine the relationship between pre-slaughter condition on CK and meat quality attributes.


The effect of breed on the levels of plasma creatine kinase

The Bonsmara (Table 2) breed had significantly higher (705.3±80.57 U/L) creatine kinase (CK) levels than Brahman (461.8±80.63 U/L), and Beefmaster breed (657.4±80.63 U/L) respectively. The higher CK activity in Bonsmara indicate that the breed was more susceptible to stress before slaughter and also could have been due to rough driving and mishandling before and after transportation (Broom et al., 1996; Averos et al., 2008). Warries et al. (1995) and Early et al. (2010) highlighted that animals transported for periods of between 5 to 15 h are susceptible to stress which increases the activity of creatine kinase. The reaction of this breed to longer transportation is perplexing because it has been highlighted that Bonsmara are able to withstand harsh and unfamiliar conditions (Ndlovu et al., 2008). However, if animals are transported for longer hours they have reduced plasma CK levels if they are given feed and water at the lairages.

Least square means (±SE) for the effect of breed on activity of creatine kinase cattle at slaughter

The lower the CK levels in Brahman and Beefmaster showed that these two breeds were not susceptible to stress during transportation as compared to Bonsmara breed. This is because these are synthetic breeds which can adapt well to handling and adverse environmental conditions (Beatty et al., 2006). The activity of CK increases in the blood after tissue damage, poor muscular tissue reperfusion, decreased heat and fatigue; apparently as a result of an increase in the permeability of the muscle membranes induced by capture, loading and transportation (Lopez et al., 2006; Guardia et al., 2009). Furthermore, other animals also are susceptible to transport stress leading to highest levels of CK in the blood (Yu et al., 2009).

Effect of sex and age on plasma creatine kinase levels

Male animals had the highest CK values (p<001) compared to female animals as shown in Table 3. Males were more stressed than females and this agrees with the study by Mota-Rojas et al. (2006) which indicated that the sex of the animal has an effect on the response of the animal to stress. This is because male animals are more excitable, aggressive and stress sensitive than females. Therefore, that could be the reason of increased plasma activity of CK found in steers transported for 12 to 16 h. Similar results were reported by Neal et al. (2009) who reported that male humans had higher levels of CK.

Least square means (±SE) for the effect of sex and age of animal on the activity of creatine kinase levels of cattle at slaughter

Creatine kinase levels were higher (p<0.05) in 18 months old animals than in 16 and 24 months old animals as shown in Table 3. According to Smith et al. (1998) the different levels of CK in the older animals is not well understood although it might be due to muscle fatigue as a result of strenuous exercise. Similarly, the CK levels in 16 month old animals was higher (p<0.05) than in 24 month old. Young animals are poorly adapted to cope with transportation related stress thus it results in high rates of injuries. Young animals respond to transportation stress by increasing their body temperature, heart rate and plasma cortisol concentration (Steinhardt and Thielscher, 1999). The findings of the current study are similar to those reported by Parrot et al. (1994), who observed that young animals are at risk of more bruising than older animals resulting in high levels of CK. This is also linked with CK in muscles as it is expressed in higher levels in males and younger animals (Neal et al., 2009).

Effects of season on bruising score and the levels of plasma creatine kinase

Results on the effect of season on the bruising score and CK levels are shown in Table 4. Creatine kinase (CK) was higher during the winter season (704.1±43.85 U/L) and lower in the summer season (181.3±124.09 U/L), while spring (698.3±59.19 U/L) and autumn (658.0±116.25 U/L) had intermediate CK levels. Knowles et al. (1998) observed similar results and found that CK levels were higher in winter season due to poor pre-slaughter conditions. However, other authors reported an increased CK values in lambs as a result of physical stress during summer (Lowe et al., 2002; Miranda-de la Lama et al., 2011). Wilson et al. (1990) and Liu et al. (2008) reported that vigorous physical activity and high temperatures had an effect on CK level.

Least square means (±SE) for effects of season on bruising score and CK levels of cattle

Autumn had the highest (p<0.05) bruising score (1.86±1.07), while winter (1.44±0.13) and spring (1.32±0.54) had intermediate scores and the summer season had the least bruising score (1.12±0.11) as shown in Table 3 and 4. Seasonal factors mainly temperature and humidity also seem to play a role on the prevalence of bruises. Results from this study concur with Gosalvez et al. (2006) and dalla Costa et al. (2007), who found that cold weather combined with precipitation, increased the mean number of bruises per animal, although the reasons are still unclear.

Effect of breed on meat quality

Ultimate pH was significantly affected by the breed of cattle as shown in Table 5. Beef from Beefmaster and Brahman breeds had significantly higher (p<0.05) pHu values of 6.3±0.05 and 6.1±0.06, respectively compared to that of the Bonsmara breed which had an average value of 5.6±0.05. The pHu levels of Bonsmara were within the optimal range and similar to those reported by Muchenje et al. (2008). Different temperament found within an individual breed can influence meat quality due to inherent genetic parameters. The findings of this study are similar to the findings by Grandin (1997), Mormede et al. (2002), Fazio and Ferlazo (2003) who reported that breed temperament influence the degree to which animals respond to stress.

Effects of breed on ultimate pH, colour of meat and the activity of creatine kinase from beef cattle (±SEM)

The higher pH values obtained from Beefmaster and Brahman could be due to stress prior to slaughter, rough handling, inclement temperatures or anything that causes the animal to draw on its glycogen reserves before slaughter. These two breeds are known to have aggressive behaviour and are more prone to injuries resulting in carcass damage and poorer meat. The current results are similar to observations by Voisinet et al. (1997) and King et al. (2006) who found that cattle which are highly temperamental had a high incidence of borderline dark cutters than calm cattle. Brahman breed had a significant effect (p<0.05) on L* values, with darker meat than Bonsmara and Beefmaster breeds. The darker meat produced by the Brahman breed in comparison to the Bonsmara show similar trends to the findings by O’Neill et al. (2006). O’Neill et al. (2006) observed that indigenous breeds such as Bonsmara released more catecholamines than exotic breeds during the pre-slaughter period, causing the depletion of glycogen.

Bonsmara and Beefmaster had the highest a* values with Brahman having the least a* values. The a* values ranged from 17.5±0.53 in meat from Bonsmara to 13.2±0.66 in meat from Brahman. The a* values from these breeds were within the normal range. This is because pHu values were fairly normal and colour is not greatly affected by pre-slaughter conditions such as transportation. Bonsmara and Beefmaster breeds had the highest b* values (12.8±0.53) while the lowest b* values (9.6±0.61) were found from the Brahman breed. This could be due to the extent of stress during handling and transportation depending on the temperament, handling during the journey, condition of the animal, and duration of the journey resulting in pale colour of animals.

Effect of age on meat quality

The effect of age on meat quality attributes is shown in Table 6. Meat from 16, 18, and 24 months old animals had no effect on pHu. The pHu values were 6.1 in meat from 16, 18, and 24 months old. The pHu values from 16, 18, and 24 months old cattle were above the normal pH range. Meat with an ultimate pH that is above 6.0 is categorised as a dark firm dry (DFD) meat (Bartos et al., 1993; Mounier et al., 2006) with increased tenderness (Silva et al., 1999) and poor palatability (Viljoen et al., 2002; Wulf et al., 2002). This can also be attributed to the fact that an animal’s behaviour differs with its age (Terlouw, 2005). The age of the group had an effect (p<0.05) on L* values (Table 3 and 6). The L* values ranged from 25.2±0.71, 26.6±0.83, and 20.9±0.73 for 16, 18, and 24 months old cattle respectively.

Effect of age on ultimate pH, colour of meat and the activity of creatine kinase from cattle slaughtered at the abattoir (±SEM)

Darker L* values were observed in 24 months old animals indicating meat becomes darker with increasing animal age. These results are similar to findings by Funghi et al. (1994) who found that L* values decreased with an increase in slaughtering age. Zhang et al. (2005) found that high pH meat had lower L* (lightness), a* (redness) and b* (yellowness) values than normal pH meat, indicating that high pH meat is darker and less brown than in normal pH meat. Meat from 16 month old animals had the highest a* and b* (17.9±0.53) and (12.9±0.49) and the meat from 24 months old animals had the least values of (12.9±0.54) and (9.3±0.51) respectively. Differences in meat colour has been associated with variations in intramuscular fat, moisture content and is age dependent on the muscle myoglobin content (Lawry, 1974).

Effect of breed on bruising scores of beef animals

Effect of breed on the bruising scores of slaughtered beef animals are shown in Figure 1. The Bonsmara breed had the highest percentage of non-bruised meat (88%) and the Brahman had the lowest none bruised meat (49%). Bonsmara also had highest percentage 20% of slightly bright red bruised meat (0 to 10 h old) while Brahman had lowest percentage of bright red bruised meat 4%. Susceptibility to bruising and temperament differs between individual animals. Fordyce et al. (1985) concluded that bruising score was beyond the breed differences. It appears that breed differences can be attributed to differences in behaviour and either being horned or hornless (Minka and Ayo, 2007).

Figure 1

Effect of breed on bruising scores of slaughtered beef animals. None = 0 h; slightly = 0 to 10 h old, red and hemorrhagic (bright-red); medium = 24 h old, dark- red colour; severe = 24 to 38 h old, watery consistency.

Bonsmara had the highest percentage 10% medium-bruised dark red (24 h old) with Brahman having the lowest bruised dark red meat 5%. This could have been due several injuries associated with road transportation. The level of bruising might increase with the length of journey travelled by the animals (Weeks et al., 2002). Brahman and Beefmaster breeds had the lowest percentage of severe deep bleeding bruising or injuries (24 to 38 h old).

Effect of age on bruising scores of beef animals

The highest number of slightly bright red bruises was observed in 16 month cattle while the 24 month old cattle had the lowest slightly bright bruised meat (Figure 2). The highest medium dark red bruises was observed in 16 month and lowest medium dark red bruise was observed in the 24 month old cattle. Young animals are more prone to bruising than old animals and this might have been due to increased poor handling (Strappini et al., 2009). Similar results were reported by von Holleben et al. (2009) indicating that the number heifers that had severe bruises were higher than either bulls or cows. However, contradicting results reported by Wythes and Shorthose (1991) highlighted that heavy animals had more bruising than young animals and mature ones and between old cows and old steers of the group. This therefore could mean that cattle bruising does not depend on the age of the animal but is mainly determined by the type of vehicle, duration of transport and handling methods used prior and after transport.

Figure 2

Effect of age on the bruising scores of cattle. None = 0 h; slightly = 0 to 10 h old (red and hemorrhagic bright-red); medium = 24 h old (dark-red colour); severe = 24 to 38 h old (watery consistency).

Effect of sex on bruising scores of slaughtered beef animals

Highest levels of slightly bruised meat and dark bruised meat were observed in female animals, while male animals had the lowest slightly bright bruised meat and medium dark bruised meat (Figure 3). In the literature there is evidence that female animals had higher chances for bruising than male animals (Yeh et al., 1978; Javis et al., 1995). These results are similar with Yeh et al. (1978) who found that heifers had significantly more bruises than steers and also when animals kept as separate groups, cows bruise significantly more than steers and bulls. Cows were the only class of animal where the amount of bruising increased with duration of the journey. Both male and female animals had a lower percentage of severe deep bruises.

Figure 3

Effect of sex on bruising scores of cattle. None = 0 h; slightly = 0 to 10 h old (red and hemorrhagic bright-red); medium = 24 h old (dark- red colour); severe = 24 to 38 h old (watery consistency).


Results from this study showed that animal factors such as sex, breed and animal age at slaughter contribute to the development of bruises, and have an effect on creatine kinase (CK) levels and meat quality. The highest percentage of slightly bruised meat was observed in Bonsmara breed, while Beefmaster and Brahman had intermediate bruising scores. Bonsmara breed had higher CK levels than Beefmaster and Brahman breeds. In addition seasonal changes also had an effect on creatine kinase and meat quality. Creatine kinase levels were higher during winter months than in spring and autumn. On the other hand, distance had a negative relationship on beef quality (L*, a*, and b*), however no correlation was observed between distance and CK levels. It was also concluded that there is no significant relationship between meat parameters (L,* a*, and b*) and CK levels.


The authors wish to acknowledge the support received from the Meat Industry Trust fund (MIT) and Project C263. This study would not be successful without the assistance from the East London abattoir and National Health Laboratory Services.


Anderson B, Horder JC. 1979;The Australian carcass bruising scoring system. Queensl Agric J 105:281–287.
Averos A, Martin S, Riu M, Serratosa J, Gosalvez LF. 2008;Stress response of extensively reared young bulls being transported to growing-finishing farm under Spanish summer commercial conditions. Livest Sci 119:174–182.
Bartos L, France C, Rehak D, Stipkova L. 1993;A practical method to prevent dark cutting (DFD) in beef. Meat Sci 34:275–282.
Beatty DT, Barnes A, Taylor E, Pethick D, McCarthy M, Maloney SK. 2006;Physiological responses of Bos Taurus and Bos indicus cattle to prolonged, continuous heat and humidity. J Anim Sci 84:972–985.
Blood DC, Studdert VP. 1988;Bailliere’s Comprehensive Veterinary Dictionary. BailliereTindall :24–28.
Bonney RJ. 2006;Farm animal welfare at work. Appl Anim Behav Sci 100:140–147.
Broom DM, Goode JA, Hall SJG, Lloyd DM, Parrot RF. 1996;Hormonal and physiological effects of a 15- hour road journey in sheep: comparison with the responses to loading, handling and penning in the absence of transport. Br Vet J 152:593–604.
Chulayo AY, Muchenje V. 2013;The effects of pre-slaughter stress and season on the activity of plasma creatine kinase and mutton quality from different sheep breeds slaughtered at a smallholder abattoir. Asian Australas J Anim Sci 26:1762–1772.
Chulayo AY, Tada O, Muchenje V. 2012;Research on pre-slaughter stress and meat quality: A review of challenges faced under practical conditions. Appl Anim Husb Rural Dev 5:1–6.
Dalla Costa OA, Fauctitano L, Coldebella A, Ludke JV, Pelso JV, Dalla Roza D, Parranhos da Costa MJR. 2007;Effect of the season of the year, truck type, and location on truck on skin bruises and meat quality in pigs. Livest Sci 107:29–36.
Devine CE, Lowe TE, Wells RW, Edwards NJ, Edwards JEH. 2006;Pre-slaughter stress arising from on farm handling and its interactions with electrical stimulation on tenderness of lambs. Meat Sci 73:304–312.
Diene CA, Storey KB. 2009;Creatine kinase regulation by reversible phosphorylation in frog muscle. Comp Biochem Physiol Part B Biochem Mol Biol 152:405–412.
Early B, Murray M, Prendiville DJ. 2010;Effect of road transport for up to 24 hours followed by twenty-four hour recovery on live weight and physiological responses of bulls. BMC Vet Res 6:38.
EFSA Panel on Animal Health and Welfare (AHAW). 2011;Scientific opinion concerning the welfare of animals during transport. EFSA J 2011. 91966. :125.
Fazio E, Ferlazzo A. 2003;Evaluation of stress during transport. Vet. Res. Commun. 27(Suppl):519–524.
Fike K, Spire MF. 2006;Transportation of cattle. Vet Clin North Am Food Anim Pract 22:305–320.
Fordyce G, Goddard ME, Tyler R, William C, Toleman MA. 1985;Temperament and bruising of Bos Indicus cross cattle. Aust J Exp Agric 25:283–288.
Funghi R, Pugliese C, Bozzi R, Lucifero M. 1994. Evolution of performances of Chianina young bull from 6–24 months. Quality parameters of meat. 81–86, Production. In : 7th International Congress on Italian beef cattle contest. Perugia, Italia.
Gosalvez LF, Averos X, Valdevira JJ, Herranz A. 2006;Influence of season distance and mixed loads on physical and carcass integrity of pigs transported to slaughter. Meat Sci 73:553–558.
Gispert M, Faucitano L, Oliver MA, Guárdia MD, Coll C, Siggens K, Harvey K, Diestre A. 2000;A survey of pre-slaughter conditions, halothane gene frequency, and carcass and meat quality in five Spanish pig commercial abattoirs. Meat Sci 55:97–106.
Gracey JF, Collins DS, Huey RJ. 1999. Meat Hygiene 10th editionth ed. Elsevier Health Science Publishers. United Kingdom:
Grandin T. 1997;Assessment of stress during handling and transport. J Anim Sci 75:249–257.
Grandin T. 2001. Livestock handling and transport 2nd editionth ed. CAB International. Wallingford, Oxford, United Kingdom:
Grandin T. 2007. Livestock Handling and Transport 3rd Editionth ed. CAB International. Wallingford Oxford, United Kingdom:
Gregory NG. 2010;How climatic changes could affect meat quality. Food Res Int 43:1866–1873.
Grzyb K, Skorkowski EF. 2005;Characterization of creatine kinase isoforms in herring (Clupea harengus) skeletal muscle. Comp Biochem Physiol Part B Biochem Mol Biol 140:629–634.
Guardia MD, Estany J, Balasch S, Oliver MA, Gispert M, Diestre A. 2009;Risk assessement of skin damage due to pre-slaughter conditions and RYRL gene in pigs. Meat Sci 81:745–751.
Hemsworth PH, Rice M, Karlen MG, Collega L, Barnett JN, Coleman GJ. 2011;Human-animal interactions at abattoirs: Relationships between handling and animal stress in sheep and cattle. Appl Anim Behav Sci 135:24–33.
Hoffman L, Lühl J. 2012;Causes of cattle bruising during handling and transport in Namibia. Meat Sci 92:115–124.
Huertas SM, Gil AD, Piaggio JM, van Eerdenburg FJCM. 2010;Transportation of beef cattle to slaughterhouses and how this relates to animal welfare and carcase bruising in an extensive production system. Anim Welf 19:281–285.
Jarvis AM, Selkirk L, Cockram MS. 1995;The influence of source, sex class and pre-slaughter handling on the bruising of cattle at two slaughterhouses. Livest Prod Sci 43:215–224.
Kannan G, Terrill YH, Kouakow B, Gazal OS, Gelaye S, Amoah EA, Samake S. 2000;Transportation of goats: effects on physiological stress responses and live weight loss. J Anim Sci 78:1450–1457.
King DA, Schuehle Pfeiffer CE, Randel RD, Welsh TH Jr, Oliphint RA, Baird BE, Curley KO Jr, Vann RC, Hale DS, Savell JW. 2006;Influence of animal temperament and stress responsiveness on the carcass quality and beef tenderness of feedlot cattle. Meat Sci 74:546–556.
Knowles TG. 1999;A review of the road transport of cattle. Vet Rec 144:197–201.
Knowles TG, Warriss PD, Brown SN, Edward JE. 1998;Effects of stocking density on lambs being transported by road. Vet Rec 142:505–509.
Lawry RA. 1974. Meat science 2nd editionth ed. Oxford: Pergamon Press.
Liu ZL, Chen P, Li JM, Lin SB, Wang DM, Zhu LP, Yang DP. 2008;Conjugated linoleic acids (CLA) moderate negative responses of heat stressed cows. Livest Sci 118:255–261.
Liste MG, María GA, García-Belenguer S, Chacón G, Gazzola P, Villarroel M. 2008;The effect of transport time, season and position on the truck on stress response in rabbits. World Rabbit Sci 16:229–235.
Lopez O, Marco I, Montane J, Lavin S. 2006;Transport stress in Southern chamois (rupricapra pyrenaica) and its modulation by acepromazine. Vet J 172:347–355.
Lowe TE, Gregory NG, Fisher AD, Payne SP. 2002;The effects of temperate evaluation and water deprivation on lamb physiology welfare and meat quality. Aust J Agric Res 53:704–714.
Marufu MC, Qokweni L, Chimonyo M, Dzama K. 2011;Relationships between tick counts and coat characteristics in Nguni and Bonsmara cattle reared on semiarid rangelands in South Africa. Ticks Tick Born Dis 2:172–177.
Minka NS, Ayo JO. 2007;Effects of loading behaviour and road transport stress on traumatic injuries in cattle transported by road during hot dry season. Livest Sci 107:91–95.
de la Lama GC, Sepulveda WS, Villarroel M, Maria GA. 2011;Livestock vehicle accidents in Spain: causes, consequences, and affects on animal welfare. J Appl Anim Welf Sci 14:109–123.
Miranda-de la Lama GC, Villarroel M, Liste G, Escos J, Maria GA. 2010;Critical points in the chain of (Please check again!) lamb in Spain that may compromise the animal’s welfare. Small Rumin Res 90:174–178.
Mormede P, Courvoisier H, Ramos A, Marissal-Avry N, Ousova O, De ‘saute’s C. 2002;Molecular genetics approaches to investigate individual variations in behavioural and neuroendocrine stress responses. Phychoneuroendocrinology 27:563–583.
Morta-Rojas D, Becerril M, Lemus C, Sanchez P, Gonzalez M, Olmos SA, Ramirez R, Alonso-Spilsbury M. 2006;Effects of mid-summer duration on pre- and post-slaughter performance and pork quality in Mexico. Meat Sci 73:404–412.
Mounier L, Dubroeucq H, Andanson S, Veissier I. 2006;Varriations in meat pH of beef bulls in relation to conditions of transfer to slaughter and previous history of the animals. J Anim Sci 84:1567–1576.
Muchenje V, Dzama K, Chimonyo M, Raats JG, Strydom PE. 2008;Meat quality of Nguni, Bonsmara and Aberdeen Angus steers raised on natural pasture in the Eastern Cape, South Africa. Meat Sci 79:20–28.
Muchenje V, Dzama K, Chimonyo M, Strydom PE, Raats JG. 2009;Relationship between pre-slaughter stress responsiveness and beef quality in three cattle breeds. Meat Sci 81:653–657.
Ndlovu T, Chimonyo M, Okoh AI, Muchenje V. 2008;A comparison of stress hormone concentration at slaughter in Nguni, Bonsmara and Angus steers. Afr J Agric Res 3:96–100.
Neal RC, Ferdinand KC, Ycas J, Miller E. 2009;Relationship of ethnic origin, gender, and age to blood creatine kinase levels. Am J Med 122:73–78.
O’Neill HA, Webb EC, Frylinck L, Strydom PE. 2006. The stress responsiveness of three different beef breed types and the effect on ultimate pH and meat colour. In : Proceedings 52nd International Congress of Meat Science and Technology. p. 181–182. 13–18 August 2006; Dublin, Ireland.
Parrot RF, Misson BH, De la Riva CF. 1994;Differential stressors effects on the concentration of cortisol, prolactin and catecholamines in the blood of sheep. Res Vet Sci 56:234–239.
Statistical Analysis System. 2003. SAS/STATT Guide to Personal Computers, Version 6 Statistical Analysis System Institute Inc. Cary, NC, USA:
Silva JA, Patarata L, Martins C. 1999;Influence of ultimate pH on bovine meat tenderness during ageing. Meat Sci 52:453–459.
Smith SA, Montain SJ, Matott RP, Zientara GP, Jolesz FA, Fielding RA. 1998;Creatine supplementation and age influence muscle metabolism during exercise. J Appl Physiol 85:1349–1356.
Steinhardt M, Thielscher HH. 1999;Maturity of suckler calves and dairy calves at the second and third week of postnatal age and forms of reaction of the animals to transport by road. Landbauforsch Volk 49:70–89.
Strappini AC, Metz JHM, Gallo G, Kemp B. 2009;Origin and assessment of bruises in beef cattle at slaughter. Animal 3:728–736.
Strappini AC, Frankena K, Metz JHM, Gallo G, Kemp B. 2010;Prevalence and risk factors for bruises in Chilean bovine carcasses. Meat Sci 86:859–864.
Terlouw C. 2005;Stress reactions at slaughter and meat quality in pigs: genetic background and prior experience. A brief review of recent findings. Livest Prod Sci 94:125–135.
de Water G, Verjans F, Geers G. 2003;The effect of short distance transport under commercial conditions on the physiology of slaughter calves; pH and colour profiles of veal. Livest Prod Sci 82:171–179.
Viljoen HF, De Kock HL, Webb EC. 2002;Consumer acceptability of dark, firm and dry (DFD) and normal pH beef steaks. Meat Sci 61:181–185.
Vimiso P. 2010. Effects of marketing channel on bruising, ultimate pH and colour of beef; and stakeholder perception on quality of beef from cattle slaughtered at a small holder abattoir. MSc Thesis University of Fort Hare; South Africa:
Vimiso P, Muchenje V. 2013;A survey on the effect of transport method on bruises, pH and colour of meat from cattle slaughtered at a South African commercial abattoir. S Afr J Anim Sci 43:105–111.
Voisinet BD, Grandin T, O’Connor SF, Tatum JD, Deesing MJ. 1997;Bos indicus cross feedlot cattle with excitable temperaments have tough meat and a higher incidence of borderline dark cutters. Meat Sci 46:367–377.
Von Holleben K, Henke S, Schmidt T. 2009. Effect of transport time up to 8h on physiological and biochemical stress indicators and resulting carcass/meat quality in cattle - an integrated approach. In : Theatre presentation, 25th August, 60th Annual Meeting of EAAP. Barcelona, Spain. Book of Abstracts No 15. p. 243.
Vojtic I. 2000;Macro CK type 1 as a major component of serum creatine kinase activity in pregnant sheep. Small Rumin Res 35:249–253.
Warriss PD, Brown SN, Knowles TG, Kestin SC, Edwards JE, Dolan SK, Philips AJ. 1995;The effects on cattle of transport by road for up to fifteen hours. Vet Rec 136:319–323.
Weeks CA, McNally PW, Warriss PD. 2002;Influence of the design of facilities at auction markets and animal handling procedures on bruising in cattle. Vet Rec 150:743–748.
Wilson BW, Nieberg PS, Buhr RJ, Kelly BJ, Shultz FT. 1990;Turkey muscle growth and focal myopathy. Poult Sci 69:1553–1562.
Wulf DM, Emnett RS, Leheska JM, Moeller SJ. 2002;Relationship among glycolytic potential, dark cutting (dark, firm and dry) beef, and cooked beef palatability. J Anim Sci 80:1895–1903.
Wythes JR, Shorthose WR. 1991;Chronological age and dentition of effects on carcass and meat quality of cattle in Northern Australia. Aust J Exp Agric 31:145–152.
Yeh E, Anderson B, Jones PN, Shaw FD. 1978;Bruising in cattle transport over a long distance. Vet Rec 103:117–119.
Yu J, Tang S, Bao E, Zhang M, Hao Q, Yue Z. 2009;The effect of transportation on the expression of heat shock proteins and meat quality of M. longissimus dorsi in pigs. Meat Sci 83:474–478.
Zhang SX, Farouk MM, Young OA, Wieliezco KJ, Podmore C. 2005;Functional stability of frozen normal and high pH beef. Meat Sci 69:765–772.

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Figure 1

Effect of breed on bruising scores of slaughtered beef animals. None = 0 h; slightly = 0 to 10 h old, red and hemorrhagic (bright-red); medium = 24 h old, dark- red colour; severe = 24 to 38 h old, watery consistency.

Figure 2

Effect of age on the bruising scores of cattle. None = 0 h; slightly = 0 to 10 h old (red and hemorrhagic bright-red); medium = 24 h old (dark-red colour); severe = 24 to 38 h old (watery consistency).

Figure 3

Effect of sex on bruising scores of cattle. None = 0 h; slightly = 0 to 10 h old (red and hemorrhagic bright-red); medium = 24 h old (dark- red colour); severe = 24 to 38 h old (watery consistency).

Table 1

Colour observations used to estimate the age of bruises

Observable colour of the bruise Estimated age of the bruise in hours
Red and haemorrhagic (bright-red) 0 to 10 h old
Dark- red colour Approximately 24 h old
Watery consistency 24 to 38 h
Rusty orange colour, soapy to touch, clear yellow mucus +72 h (3 days old)

Table 2

Least square means (±SE) for the effect of breed on activity of creatine kinase cattle at slaughter

Breed CK (U/L)
Bonsmara (n = 108) 705.3±80.57a
Beefmaster (n = 130) 657.4±73.85ab
Brahman (n = 83) 461.8±80.63b

Means in the same column with different superscripts are significantly different at * p<0.05.

n = Total number of animals. CK = Creatine kinase. U/L = Units per litre.

Table 3

Least square means (±SE) for the effect of sex and age of animal on the activity of creatine kinase levels of cattle at slaughter

Variables N CK (U/L)
 Females 222 579.3±13.65b
 Males 91 724.6±21.31a
 16 months 116 636.6±76.83b
 18 months 85 729.6±89.75a
 24 months 112 524.1±78.19c

Means in the same column with different superscripts are significantly different at * p<0.05.

N = Total no. of animal per breed. CK = Creatine kinase.

U/L = Units per litre.

Table 4

Least square means (±SE) for effects of season on bruising score and CK levels of cattle

Season Bruising score Creatine kinase (U/L)
Spring 1. 32±0.54b 698.3±59.19a
Summer 1.12±0.11c 181.3±124.09b
Winter 1.44±0.13b 704.1±43.85a
Autumn 1.86±1.07a 658.0±116.25a

Means in the same column with different superscripts are significantly different at * p<0.05.

U/L = Units per litre.

Table 5

Effects of breed on ultimate pH, colour of meat and the activity of creatine kinase from beef cattle (±SEM)

Parameters Breed

Bonsmara Beefmaster Brahman
N 108 130 83
CK( U/L) 705.3±80.57a 657.3±73.85ab 461.8±80.63b
pHu 5.6±0.05b 6.3±0.05a 6.1±0.06a
L* 24.8±0.78a 24.6±0.71a 21.7±0.89b
a* 17.5±0.53a 16.8±0.53a 13.2±0.66b
b* 12.8±0.53a 11.0±0.49a 9.6±0.61b

Means in the same row with different superscripts are significantly different at * p<0.05.

pHu = Ultimate pH. L* = Lightness. a* = Redness. b* = Yellowness.

CK = Creatine kinase. U/L = Units per litre. N = Total number of animals per breed.

Table 6

Effect of age on ultimate pH, colour of meat and the activity of creatine kinase from cattle slaughtered at the abattoir (±SEM)

Parameters Age

16 months 18 months 24 months
N 120 87 114
CK (U/L) 636.9±76.83b 729.6±89.75a 524.1±0.63c
pHu 6.1±0.05a 6.1±0.06a 6.1±0.05a
L* 25.2±0.71a 26.6±0.83a 20.9±0.73b
a* 17.9±0.53a 17.7±0.62a 12.9±0.54b
b* 12.9±0.49a 11.5±0.58b 9.3±0.51c

Means in the same row with different superscripts are significantly different at * p<0.05.

pHu = ultimate pH.

L* = Lightness. a* = Redness. b* = Yellowness.

CK = Creatine kinase. U/L = Units per litre.