Meat Science 66 (2004) 425–436 www.elsevier.com/locate/meatsci

Characterisation of young bulls of the (selected from old Brown Swiss) in relation to carcass, meat quality and biochemical traits

X. Serraa,*, M. Gila, M. Gisperta, L. Guerreroa, M.A. Olivera, C. San˜ udob, M.M. Campob, B. Paneab, J.L. Olletab, R. Quintanillac,1, J. Piedrafitac aInstitut de Recerca i Tecnologia Agroalimenta`ries (IRTA), Centre de Tecnologia de la Carn, Granja Camps i Armet s/n, 17121 Monells, Girona, Spain bDepartamento de Produccio´n Animal y Ciencia de los Alimentos, Universidad de Zaragoza, Miguel Servet, 177, 50013, Zaragoza, Spain cDepartament de Cie`ncia Animal i dels Aliments, Universitat Auto`noma de Barcelona, Campus Universitari, Edifici V, 08193 Bellaterra, Barcelona, Spain

Received 16 July 2002; received in revised form 13 May 2003; accepted 30 May 2003

Abstract Seventy-four young bulls of the Bruna dels Pirineus beef cattle breed were reared in a typical production system and slaughtered at an average age of 381 days and live weight of 541 kg. The animals were evaluated for productive traits, carcass quality, meat quality, eating quality and biochemical characteristics of m. longissimus thoracis (LT). Biochemical measurements included intra- muscular fat (IMF) and collagen proportion, haem pigment concentration, lactate dehydrogenase and isocitrate dehydrogenase activities and type I (MHC I) fibre percentage determined by ELISA. Bruna dels Pirineus bulls achieved high growth rates during the fattening period (1.63 kg/day) and showed good carcass quality, with a high dressing-out proportion (607 g/kg, hot carcass), a good conformation score (U; EUROP) and a moderate fatness score (3;1–5). Carcass composition was estimated from the sixth rib joint dissection (682 g/kg lean proportion, 127 g/kg total dissectable fat and 163 g/kg bone). IMF (24.3 g/kg) and MHC I (27.9%) showed high variabilities (CV>30%). Sensory analysis of LT included beef and livery odour and flavour intensity, and overall tenderness and juiciness assessment of loin samples (14-day ageing). Beef odour and flavour were slightly positively correlated with IMF and carcass fatness score (P<0.05). Fatness, MHC I, insoluble collagen and cooking losses tended to affect the livery flavour intensity positively. This variable was significantly higher in meat from bulls of lower carcass quality (i.e. lower conformation score, lower lean proportion; P<0.05) and higher type I fibre percentage. Loin overall tenderness and juiciness were not affected by the biochemical traits studied, however, they were negatively affected by cooking loss (P<0.05). # 2003 Elsevier Ltd. All rights reserved. Keywords: Beef breed; Carcass quality; Chemical composition; Muscle fibres; Sensory evaluation

1. Introduction E.U. meat quality labels, normally under geographical groupings and with a specific genotype and production The consumption of beef in Europe has decreased in system, conforms with the increasing demand of con- recent decades. Consumers are not only worried about sumers for a quality guarantee (Guerrero, 2001). Many meat quality and its healthiness, but also about its origin local and rustic (non-improved type) beef are and how animals are reared and handled. The creation of reared under traditional systems and designations of origin or quality trade marks (Protected Designation of Origin, Protected Geographical Indication) which pro- * Corresponding author. Tel.: +34-972-630-052; fax: +34-972- mote a diversification of agricultural production and 630-373. E-mail address: [email protected] (X. Serra). specific products. The use of local breeds as an alter- 1 Present address: INRA—Station de Ge´ ne´ tique Quantitative et native system of beef production has the advantage that Applique´ e, 78352 Jouy-en-Josas Cedex, France. these breeds are closely related to the environment and

0309-1740/03/$ - see front matter # 2003 Elsevier Ltd. All rights reserved. doi:10.1016/S0309-1740(03)00131-1 426 X. Serra et al. / Meat Science 66 (2004) 425–436 helpto maintain biodiversity and sustainable agri- herds all over the Catalan Pyrenean area and originating cultural production, especially in depressed areas. from a minimum of 44 sires. Unfortunately it was not The Bruna dels Pirineus is a medium-sized meat type possible to collect samples from one of the young bulls breed, similar to the American Braunvieh, located in the for meat quality and biochemical analyses and, there- Eastern . The origin of the Bruna dels Pirineus fore, this animal was not included in this paper (n=74) (BP) is a dual purpose breed (dairy and meat) selected although it was considered in another paper focussed on from early crossings between a local Pyrenean breed growth and carcass quality traits (Piedrafita et al., and the old Brown Swiss. BP breed shows an excellent 2003). The animals were evenly distributed over two maternal aptitude and performance in mountain areas consecutive years. The calves were reared in a semi- and could be considered as a fast-growth breed (Campo, extensive regime together with their mothers and started San˜ udo, Panea, Albertı´ , & Santolaria, 1999). Few studies fattening at an average age of 7 months. They were fed a have been carried out to characterise production, car- concentrated meal ad libitum. The diet included pro- cass and meat quality traits of the Bruna dels Pirineus, ducts and by-products both of cereals (wheat, barley also known as Parda Alpina. Recently, Albertı´ , San˜ udo and/or oats) and soybean and was completed with hay and co-workers (Albertı´ , San˜ udo, Campo, Franco, ad libitum (Table 1). Calves were weighed at the start of Lahoz, & Olleta, 1997; Albertı´ et al., 1998; Campo et the fattening period and thereafter every 4 weeks. Dur- al., 1999; San˜ udo, Albertı´ , Campo, Olleta, & Panea, ing the fattening period, which lasted an average of 1998) made a comprehensive study on productive and 170.6 (S.D. 31.3) days, animal average growth rate carcass traits and meat quality characteristics (chemical (AGR, kg/day) was calculated. The afternoon before composition, sensory and instrumental texture analysis) slaughter, the young bulls were weighed (recorded as of seven local Spanish beef cattle breeds, which included slaughter weight, SW) and transported to the abattoir Parda Alpina (in some of these papers quoted as dual- without mixing with animals from different batches. The purpose Brown Swiss) or Bruna dels Pirineus, but with welfare of the animals was taken into account when a small number of animals (12–18 male yearlings per handling them. Animals were slaughtered by captive- breed). However these authors did not study muscle bolt pistol and dressed according to commercial prac- biochemical characteristics. In their study, the calves tice. Carcasses were chilled at 12 C for 3–4 h and stored were reared until weaning in the local conditions of each at 4Æ1 C for 24 h. breed-production system. At the age of 6 months they were fed at a common feedlot with standard feeding and 2.2. Carcass measurements slaughtered at a similar weight for all breeds (460 kg). Their study was intended as a comparison between the The hot carcass weight (without removing the fat and different breeds under the same experimental conditions including testicles, perirenal and pelvic fat) was recor- (i.e. feeding, rearing system and slaughter weight). ded. Dressing-out proportion was calculated from hot The aim of this paper is to describe the variability of the carcass weight and slaughter weight. Perirenal and pel- Bruna dels Pirineus breed-production system regarding vic fat content was expressed as the hot carcass weight carcass traits, meat quality and biochemical character- proportion. Several standard measurements were taken istics of young bulls reared on their typical production on the left half-carcass, according to De Boer, Dumont, system and slaughtered at their usual commercial weight. Pomeroy, and Weniger (1974): carcass length, hindlimb This study is part of a European project (FAIR1 CT95 length, hindlimb width and internal chest width. Also, 0702) that describes both the between and within breed- carcass compactness (carcass weight Ä carcass length) production system meat quality variability of several cattle and hindlimb compactness (hindlimb width Ä hindlimb breeds from France and Spain (Gil et al., 2001; Piedrafita length) were calculated. Carcass conformation score et al., 2003). The knowledge of the relationships among (EUROP classification) with a 15-point scale (1=poor- several productive, carcass quality, meat quality, and bio- est, ..., 15=best) and fatness score (1–5 classification) chemical traits within a breed and its characteristic pro- with a 15-point scale (1=leanest, ..., 15=fattest) were duction system can helpin the improvement by genetic selection, as well as in the understanding of the properties of the final product and its acceptability. Table 1 Diet composition

Nutrient

2. Materials and methods Crude protein (g/kg) 160.2 Ether extract (g/kg) 42.3 2.1. Animals and productive traits Crude fibre (g/kg) 50.0 Meat fodder units/kg 1.05 This study was carried out on 75 yearling males of the Calcium (g/kg) 7.7 Phosphorus (g/kg) 4.6 Bruna dels Pirineus cattle breed selected from 22 different X. Serra et al. / Meat Science 66 (2004) 425–436 427 evaluated ((EEC) No 1208/81 and (EEC) No 1026/91; stored at À20 C until analysis. Moisture (ISO 1442), (EEC) No 2930/81 and (EEC) No 2237/91). intramuscular fat (ISO 1443) and protein (ISO 937) The area of m. longissimus thoracis (LT) and its dorsal- proportions were determined. Haem pigment concen- ventral (depth) and medial-lateral (width) diameters were tration was measured according to Hornsey (1956) and measured at the 6/7th-rib level. The 6th rib joint was expressed as micrograms of acid haematin per gram of extracted (24 h post mortem) cutting the length of the bone wet muscle. Collagen determinations were carried out at the limit of the m. serratus dorsalis (Robelin & Geay, on the colour-measurement samples. To estimate total 1975). The weight of the rib joint was recorded and the LT collagen, total hydroxyproline (Hyp) concentration was was weighed and separated for the chemical analyses, while measured according to Bonnet and Kopp (1984). Col- the rest of the rib joint was vacuum packed and frozen at lagen solubility was determined by the method of Hill À20 C until dissection. The rib joints were thawed in cold (1966). water and dissected into lean, total fat (subcutaneous and intermuscular), bone and waste tissues (blood vessels, ten- 2.5. Muscle biochemical analyses dons) to determine the composition of the cut. The contractile traits of the muscle were determined 2.3. Meat quality by enzyme-linked immunosorbent assay (ELISA). The slow myosin heavy chain (MHC-I) percentage was Carcasses with an ultimate pH >6.00 were not included quantified with a specific MHC-I monoclonal antibody in the meat quality, muscle characteristics and sensory (Picard, Leger, & Robelin, 1994). The metabolic traits studies in order to avoid dark, firm and dry meats (DFD). of the muscle were determined by measuring enzyme High ultimate pH affects meat water-holding capacity and activities. The anaerobic glycolytic metabolism was colour (Sornay, Dumont & Fournaud, 1981). Further- assessed by lactate dehydrogenase (LDH) activity (mmol/ more, DFD meat undergoes early spoilage during storage min/g muscle) according to Ansay (1974) and the aerobic (Gill & Newton, 1981), and this negatively affects the oxidative metabolism by isocitrate dehydrogenase sensory characteristics of the meat. Two carcasses out of (ICDH) activity (nmol/min/g muscle) according to the 75 animals studied showed DFD characteristics and Briand, Talmant, Briand, Monin, and Durand (1981). were excluded from the subsequent sample collection Muscle samples for ELISA and enzyme activity ana- and analysis (n=72). Muscle pH measurements were lyses were taken from the LT core at the 7th-rib level at done at the centre of LT (5th rib) at 24 h post mortem 24 h post mortem. Samples were frozen in liquid nitro-  (pHU) and at the 8th-rib level at seven days post mor- gen and stored at À80 C until analysis. Further details tem (pHU7) with a xerolyt pHmeter. about preparation of samples and analyses were descri- A 3 cm-thick chopfrom between the 7th and 8th ribs bed by Gil et al. (2001). of the LT was cut (at 7 days post mortem) and placed in a polystyrene tray, covered with plastic film permeable to 2.6. Sensory analysis oxygen and stored for 24 h at 4 C. Measurements were averaged over four zones of each LT joint, changing the The muscle loin from the 9th to the 11th rib was instrument position each time. Colour measurements removed from the left carcass 24 h post mortem (DFD were carried out with a Minolta CM 2002 spectro- carcasses not sampled) and was vacuum packed and photometer with a C illuminant and a 2 standard stored at 4 C. Seven days after slaughter the LT was observer in the CIELAB space [Lightness (L*), redness taken and 2-cm steaks were cut, individually vacuum (a*), yellowness (b*)] (Commission Internationale de packed and stored at 4 C for another seven-day age- l’E´ clairage, 1976). Hue (H*) and chroma (C*) were cal- ing period. After this period (14 days from slaughter) culated from a* and b* values according to Wyszecki the samples were kept frozen at À20 C until they were and Stiles (1967). analysed. Expressible juice was measured according to a modi- Sixty-nine steaks were thawed overnight at 4–5 C, fication of the method of Grau and Hamm (1953) as afterwards they were wrapped in aluminium foil and described by Sierra (1973). Cooking loss was estimated cooked to an internal temperature of 70 C in a con- as the weight-difference between thawed steaks (14 days vection oven preheated to 200 C. Sample internal tem- ageing) and cooked steaks (oven, 70 C internal tem- perature was monitored using a thermocouple probe, perature, for details see Section 2.6). inserted horizontally to the midpoint of the steak, and a data logger. Thawed and cooked steaks were 2.4. Muscle chemical composition weighed and cooking loss was calculated. The core portion of the steaks was cut into 10 pieces. Each Samples from LT were taken at 24 h post mortem at subsample was immediately wrapped in aluminium the 6th-rib level for chemical composition determina- foil, codified and kept at 60 C. Panellists evaluated tion. These samples were minced, vacuum-packed and the cooked samples in individual booths provided 428 X. Serra et al. / Meat Science 66 (2004) 425–436 with red light and a 60 C heater. A single steak was 3. Results assessed from each animal. Each sensory evaluation session consisted of six randomly selected loin sam- 3.1. Productive and carcass quality traits ples. The six steak subsamples were tasted by each panellist in a different order in each session according Calves began the fattening period at an average age of to a previous design (Guerrero & Gua` rdia, 1997). 209 days and an average weight of 268.2 kg (Table 2). Additionally, within a session each panellist assessed The animals were slaughtered at an average age of 380.6 the same steak-location subsample in order to mini- (S.D. 34.4) days and an average live weight of 541.3 mise any steak-location effects. (S.D. 29.6) kg. Slaughter weight was decided according Panellists were required to rate each sample for inten- to the market demand of the ‘Vedella dels Pirineus sity variation in beef odour, livery odour, beef flavour, Catalans’ (‘Beef of the Catalan Pyrenees’ submitted to livery flavour, overall tenderness and overall juiciness. E.U. recognition for Protected Geographical Indica- Each attribute was rated on a non-structured 10-point tion), i.e. when reaching approximately 550 kg. scale with score 0 equivalent to no attribute intensity The yearlings showed a high AGR during the fatten- and score 10 equivalent to the highest intensity of the ing period of 1.63 (S.D. 0.23) kg/day. The average hot attribute. The mean of the 10 panellists for each steak carcass weight was 329 (S.D. 21.2) kg and the young bulls was used for data analyses. showed a good dressing-out proportion of 607 g/kg, with values ranging from 571 upto 644 g/kg. The perirenal and 2.7. Statistical analysis pelvic fat proportion was 24 g/kg carcass weight. Carcass morphological measurements showed a car- Pearson’s correlation coefficients and principal com- cass length of 126.2 cm and a carcass width of 39.2 cm, ponent analysis (PCA; Proc Factor) for meat quality while hindlimb length and maximum diameter were 79.1 and biochemical properties were obtained by the SAS and 30.6 cm, respectively. Carcass compactness and statistical package (SAS, 1988). hindlimb compactness indices were also calculated: 2.6

Table 2 Means, standard deviations (S.D.) and minimum and maximum values for productive traits and carcass quality characteristics of the Bruna dels Pirineus young bulls (n=74)

Mean S.D. Minimum Maximum

Initial age (day) 208.9 30.1 144 293 Initial live weight (kg) 268.2 40.7 197 372 Fattening period (day) 170.6 31.3 101 242 Age at slaughter (day) 380.6 34.4 334 453 Slaughter weight (kg) 541.3 29.6 440 602 Average growth rate (kg/day) 1.63 0.23 1.04 2.14 Carcass weight (kg) 329.0 21.2 257.6 376.0 Dressing-out proportion (g/kg) 607 16 571 644 Perirenal-pelvic fat proportion (g/kg) 24 6 15 42 Carcass length (cm) 126.2 2.7 120.7 132.3 Internal chest width (cm) 39.2 1.3 35.9 42.5 Hindlimb length (cm) 79.1 2.1 74.0 83.2 Hindlimb width (cm) 30.6 1.4 28.0 34.3 Carcass compactness (kg/cm)a 2.6 0.2 2.1 3.0 Hindlimb compactnessb 0.39 0.02 0.36 0.43

Conformation score 15-point scale 11.2 0.9 10 14 EUROP classification U (E=8.1%, U=91.9%)

Fatness score 15-point scale 6.7 1.5 4 11 1-5 classification 3 (2=44.6%, 3=51.35%, 4=4.05%)

M. longissimus thoracis Area (cm2) 50.5 7.1 34.9 67.8 Medial-lateral diameter (mm) 69.9 8.9 54 91 Dorsal-ventral diameter (mm) 88.5 8.3 70 112

a Carcass weight Ä carcass length. b Hindlimb width Ä hindlimb length. X. Serra et al. / Meat Science 66 (2004) 425–436 429

Table 3 teristics of LT are presented in Table 4. The expressible Means, standard deviations (S.D.) and minimum and maximum juice proportion of the LT was 210 (S.D. 26) g/kg and values for sixth rib joint dissection composition the cooking loss was 229 (S.D. 51) g/kg, showing a high Mean S.D. Minimum Maximum variability (CV=22.3%). Objective (instrumental) measurements of muscle Muscle (g/kg) 682 35 571 782 Overall dissectable fat (g/kg) 127 21 82 195 colour were performed at 8 days post mortem. The Subcutaneous fat (g/kg) 30 11 12 59 lightness (L*) and redness (a*) values for LT muscle Intermuscular fat (g/kg) 97 15 67 137 were 36.0 and 21.3, respectively. The calculated hue Bone (g/kg) 163 25 111 235 and chroma values were 27.4 and 24.2, respectively. Waste (g/kg) 28 7 9 44 Concerning LT muscle chemical composition, moist- ure and protein concentrations were 746 and 217 g/kg and 0.39 kg/cm, respectively. Carcasses of BP yearlings wet muscle, respectively, while intramuscular fat showed a good conformation with an 11.2 average (IMF), with 24.3 g/kg wet muscle, showed a high assessment score (15-point scale), equivalent to a U variability between animals (CV=32.9%). The con- (91.9%) on the EUROP system. Carcass fatness scores centration of muscle haem pigments in BP yearlings were moderate with 6.7 on a 15-point scale, equivalent was 148 mg acid haematin/g muscle (CV=16.1%). The to 3 (51.35%) on the 1–5 system. The LT area (7th rib amount of total collagen in LT was 3.14 mg/g wet level) was 50.5 (S.D. 7.1) cm2, with a muscle width weight and the soluble collagen proportion was 40% of (medial-lateral diameter) and muscle depth (dorsal-ven- total collagen. tral diameter) of 69.9 and 88.5 mm, respectively. The contractile and metabolic characteristics of LT Carcass tissue proportions for BP young bulls are are also presented in Table 4. The percentage of slow presented in Table 3. The dissection of the 6th rib joint oxidative fibres estimated by quantification of the MHC gave 682 g/kg of muscle, 127 g/kg of dissectable fat and I isoform was 27.9 (S.D. 8.9)%, showing a high varia- 163 g/kg of bone. bility (CV=31.9%). Muscle glycolytic metabolism assessed by LDH activity was 1171 mmol/min/g 3.2. Meat quality and biochemical characteristics of m. (CV=12.5%) and the oxidative activity ICDH was longissimus thoracis 1642 nmol/min/g (CV=23.9%). The muscle glycolytic capacity, given as the ratio between LDH and ICDH Means, standard deviations and minimum and max- activities (LDH/ICDH=0.76 mmol/nmol), also showed imum values for meat quality and biochemical charac- high variability (CV=28.9%).

Table 4 Means, standard deviations (S.D.) and minimum and maximum values for meat quality characteristics, chemical composition and biochemical properties of m. longissimus thoracis (n=72)

Mean S.D. Minimum Maximum pHU 5.56 0.09 5.35 5.93 pHU 7 days post mortem 5.53 0.11 5.24 5.89 Expressible juice (g/kg) 210 26 142 269 Cooking loss (g/kg) 229 51 112 352 L*a*b* Colour Minolta 2002 L* 36.0 2.7 28.2 41.2 a* 21.3 2.4 15.7 26.1 b* 11.2 3.2 6.5 23.5 Hue () 27.4 5.4 18.6 45.4 Chroma 24.2 3.1 17.4 33.0 Moisture (g/kg wet weight) 746.0 9.4 715.3 768.2 Total protein (g/kg wet weight) 217.0 8.4 175.2 230.6 Intramuscular fat (g/kg wet weight) 24.3 8.0 9.5 57.0 Haem pigments (mg haematin/g wet weight) 148 24 85 201 Total collagen (mg/g wet weight) 3.14 0.55 1.96 4.05 Insoluble collagen (mg/g wet weight) 1.86 0.38 0.96 2.93 Soluble collagen (% total collagen) 39.97 8.72 22.20 55.58 Myosin heavy chain I percentage 27.9 8.9 12.3 56.7 Lactate dehydrogenase (mmol/min/g muscle) 1171 146 865 1473 Isocitrate dehydrogenase (nmol/min/g muscle) 1642 393 881 2684 Ratio LDH/ICDH (mmol/nmol) 0.76 0.22 0.36 1.40 430 X. Serra et al. / Meat Science 66 (2004) 425–436

3.3. Relationships among meat quality variables and had a noticeable influence on the four first PCs biochemical characteristics of LT (loadings<0.45), i.e. they did not contribute to the total variation of these PCs. The relationships among meat quality and biochem- ical variables, i.e. the location of the variables in the 3.4. Correlations with sensory attributes multivariate space, were studied by principal compo- nent analysis (PCA). Table 5 shows the coefficients of Correlation coefficients (r) between sensory attributes correlation (r) between the variables used in the analy- and carcass quality, meat quality and LT biochemical sis. In Fig. 1 (a and b) the results of the four first prin- characteristics of BP young bulls are presented in cipal components (PCs) that accounted for 63% of the Table 6. Beef odour and beef flavour showed a positive total variation of meat quality and biochemical char- correlation. These attributes were also positively correl- acteristics are plotted. The first axis (PC 1 with 21.2% of ated (P<0.05) with intramuscular fat content and fat- total variance; Fig. 1a) included colour variables: b*, ness score, although the coefficients were low. Livery chroma and hue and expressible juice, all correlated, flavour was the sensory attribute that showed the and with lower loadings pHU and a*. Lying near the majority of significant correlations (P<0.05). On one first PC on the opposite side, the LDH activity was hand, livery flavour was positively correlated with peri- negatively correlated with pHU, b* and Hue. The sec- renal and pelvic fat, overall dissectable fat, cooking loss, ond PC (17.5%) included traits related to muscle oxi- MHC I% and insoluble collagen content. On the other dative capacity: haem pigments and ICDH activity, as hand, it showed negative correlations with carcass con- well as total and insoluble collagen and the redness col- formation score and lean (muscle) proportion. More- our component, a*. PCs three and four representing, over, livery flavour intensity was positively correlated respectively, 15.9 and 8.6% of the total variability in with overall tenderness (r=+0.38, P<0.05) and livery meat quality and biochemical characteristics are shown odour (r=+0.55, P<0.05). Overall tenderness and in Fig. 1b. The important variables in the third axis, overall juiciness were positively correlated (r=+0.58, placed farthest from the origin, were muscle glycolytic P<0.05), and both attributes were negatively correlated capacity (LDH/ICDH ratio and LDH activity), muscle with cooking loss (r=À0.25 and r=À0.69, respectively). redness and protein content which were negatively cor- related with those parameters on the opposite side, moisture proportion, ICDH activity, MHC I percentage 4. Discussion and pHU. Finally, the total and insoluble collagen and hue contributed to the variability of the fourth PC. It is The high AGR observed in BP young bulls indicated noteworthy that neither IMF nor muscle lightness (L*) a good growth-potential of the breed, it was one of the

Table 5 Correlation coefficients (r) between meat quality and biochemical characteristics of m. longissimus thoracis used in the PC analysis

pHU L* a* b* C* Hue EXP Moisture IMF Protein Pigment LDH ICDH MHC I LDH/ TOT JUICE ICDH COL pHU 1 L* À0.21 1 a* 0.00 À0.45* 1 b* 0.22 À0.18 0.39* 1 C* 0.13 À0.41* 0.86* 0.81* 1 Hue 0.21 0.08 À0.01 0.90* 0.48* 1 EXPJUICE À0.05 0.11 0.11 0.40* 0.29* 0.38* 1 Moisture 0.23 À0.12 À0.34* 0.06 À0.18 0.20 0.00 1 IMF 0.17 0.04 0.23 0.05 0.17 À0.03 À0.09 À0.44* 1 Protein À0.03 0.01 0.10 À0.06 0.03 À0.10 À0.02 À0.15 0.08 1 Pigment 0.07 À0.28* 0.43* À0.09 0.21 À0.35* À0.19 À0.36* 0.29* À0.10 1 LDH À0.36* 0.14 0.01 À0.37* À0.20 À0.40* À0.15 À0.46* À0.02 0.02 0.33* 1 ICDH 0.05 0.04 À0.12 À0.23 À0.21 À0.20 À0.37* 0.01 0.10 À0.40* 0.46* 0.23* 1 MHC I 0.13 0.07 À0.10 0.16 0.03 0.22 0.11 0.05 0.20 À0.21 0.16 À0.11 0.32* 1 LDH/ICDH À0.32* 0.07 0.16 À0.03 0.08 À0.08 0.23* À0.35* À0.07 0.37* À0.20 0.43* À0.73* À0.33* 1 TOTCOL À0.08 À0.09 0.06 0.05 0.07 0.02 À0.06 À0.19 0.14 À0.11 0.27* 0.20 0.35* 0.00 À0.25* 1 INSCOL À0.20 0.02 0.02 À0.01 0.00 0.00 À0.10 À0.09 0.13 À0.05 0.17 0.32* 0.37* 0.16 À0.12 0.67* pHU,pH24h;L*, a*, b*, colour CIE Lab (Minolta 2002) 8 days post mortem; C*, Chroma; EXPJUICE, expressible juice; IMF, intramuscular fat; LDH, lactate dehydrogenase; ICDH, isocitrate dehydrogenase; MHC I, slow myosin heavy chain (isoform mhc1); LDH/ICDH, ratio between LDH and ICDH activities; TOTCOL, total collagen; INSCOL, insoluble collagen. * Significant at the 0.05 level. X. Serra et al. / Meat Science 66 (2004) 425–436 431

Fig. 1. Relationships among meat quality variables and biochemical characteristics of LT obtained by principal component (PC) analysis (Proc Factor): (a) projection of the variables in the plane defined by the first two PCs, (b) projection of the variables in the plane defined by PCs three and four. Principal Component (%), percentage of total variance explained by the PC; EXPJUICE, expressible juice; ICDH, isocitrate dehydrogenase; IMF, intramuscular fat; INSCOL, insoluble collagen; L*, a*, b*, colour CIE Lab (Minolta, 2002) 8 days post mortem; LDH, lactate dehydrogenase;

LDH/ICDH, ratio between LDH and ICDH activities; MHC I, slow myosin heavy chain (isoform mhc1); pHU, pH 24 h; TOTCOL, total collagen.

Table 6 Correlation coefficients (r) between sensory attributes and carcass quality, meat quality and biochemical characteristics of m. longissimus thoracis

Odour Flavour Overall

Beef Livery Beef Livery Tenderness Juiciness

Beef odour 1 Livery odour 0.10 1 Beef flavour 0.61* À0.07 1 Livery flavour À0.08 0.55* À0.06 1 Overall tenderness 0.08 0.01 0.09 0.38* 1 Overall juiciness À0.06 0.02 À0.07 0.05 0.58* 1 Perirenal and pelvic fat À0.09 0.03 À0.02 0.26* À0.03 À0.12 Conformation score À0.05 À0.16 À0.01 À0.33* À0.05 0.17 Fatness score 0.27* 0.23 0.16 0.11 À0.09 À0.40* Muscle À0.17 À0.16 À0.09 À0.25* À0.08 0.08 Overall dissectable fat 0.17 0.15 0.22 0.31* 0.11 À0.16 Cooking loss 0.02 0.10 0.15 0.29* À0.25* À0.69* Intramuscular fat 0.27* 0.15 0.24* 0.16 0.02 À0.20 Myosin heavy chain I 0.09 0.11 0.20 0.24* 0.06 0.07 Insoluble collagen 0.03 0.16 0.00 0.33* 0.20 0.09

* Significant at the 0.05 level. highest among the seven local Spanish beef cattle breeds AGR as the breed, a meat breed of similar age participating in the European project (Piedrafita et al., and weight at slaughter (382.8 days and 551.6 kg, 2003): Asturiana de los Valles (AV, double-muscled respectively), and the A-NI, but the latter had different breed); Asturiana de las Montan˜ as (AM, small- to ages and weights at slaughter (363.6 days and 481 kg). medium-sized non-improved breed); Pirenaica (PI, Albertı´ et al. (1995) reported no significant differences in medium- to large-sized meat purpose breed); Avilen˜ a- AGR between Parda Alpina (PA) (synonymous with Negra Ibe´ rica (A-NI, medium-sized non-improved Bruna dels Pirineus) and PI breeds sacrificed at 470 kg breed); Morucha (MO, small- to medium-sized non- live weight and fed a concentrate diet. In another study improved breed); and Retinta (RE, large-sized non- with several local Spanish beef breeds: AV, A-NI, MO, improved breed). The BP yearlings showed the same PA/BP, PI, RE and (RG, a large-sized 432 X. Serra et al. / Meat Science 66 (2004) 425–436 meat breed) intensively reared, Albertı´ et al. (1997) found Carcass lean proportion is directly affected by the that meat breeds: PI, RG and PA showed the highest number and diameter of muscle fibres. LT area is rela- growth rate. Similarly, Campo et al. (1999), in a study with ted to fibre diameter and fibre length as both increase the same breeds on a common feeding regime found that muscle girth (Swatland, 1994). Muscle morphology PA animals (quoted as dual-purpose Brown Swiss) showed measurements showed that BP yearlings had a smaller the highest growth rate, together with the PI breed. LT area than meat breeds such as PI and AV, with The commercial value of carcasses is determined by values over 63 cm2, while less improved breeds (AM, weight, grading and tissue composition, and carcass A-NI, MO and RE) showed lower values (Piedrafita et weight depends upon slaughter weight and dressing-out al., 2003). In the BP breed, Font et al. (1997) reported percentage (Kempster, Cuthbertson & Harrington, an LT area of 71.2 cm2 but it was measured at the 10th 1982a). The high dressing-out percentage of BP young rib level. The LT width and depth were 765 and 1121 bulls was similar to the PI breed and lower, as expected, mm, respectively. than the AV breed with double muscled animals (Pie- Carcass composition is important in the evaluation of drafita et al., 2003). Albertı´ et al. (1995) reported similar carcass quality. A good method of estimation of carcass dressing proportions between PA and PI breeds slaugh- composition is through the dissection of rib-cuts. Oli- tered at the same live weight. In their study with seven va´ n, Martı´ nez, Garcı´ a, Noval, and Osoro (2001) found local Spanish breeds, Albertı´ et al. (1997) did not find that dissection of the 6th rib joint gives a better predic- significant differences in dressing percentages in PA, PI tion of tissue composition, i.e. a better estimation of and RG (meat breeds) slaughtered at 471, 460 and 472 carcass fat, than 10th rib joint dissection. Piedrafita et kg live weight, respectively. al. (2003) found the lean proportion of BP young bulls Carcass morphological measurements for BP under- was lower than PI and AV breeds, as expected, and in taken according to the recommendations of De Boer et agreement with the lower LT area of BP animals. al. (1974) showed medium–big-framed animals in com- However, Albertı´ et al. (1995) found no differences parison with the other Spanish breeds participating in between PI and PA breeds in tissue composition of the the project. BP hindlimb length and maximum diameter 10th rib joint (SW=470 kg). The joint composition were similar to those of PI, MO and RE breeds (Pie- obtained for PA yearlings was 670 g/kg muscle, 160 g/ drafita et al., 2003). In general, a high carcass compact- kg fat and 170 g/kg bone. Albertı´ et al. (1998) obtained ness index corresponds with good conformation and the following composition for PA breed: 718 g/kg mus- high dressing percentage. In a study with BP breed cle, 90 g/kg fat and 191 g/kg bone. Their results showed Font, Garcia-Macias, Guerrero, and Oliver (1997) that PA lean proportion was significantly lower than obtained a much lower carcass compactness index, that of meat breeds (PI and AV). On the other hand, in which could be explained by the lower carcass weight comparison with our results, the lower fat proportion of (249 kg) of their animals. On the other hand, Albertı´ et PA could be explained by the higher slaughter weight of al. (1998) described a 2.3 kg/cm carcass compactness BP young bulls in our study (541 vs. 460 kg), since fat index in PA young bulls (quoted as dual-purpose Brown proportion increases with age (Kempster et al., 1982b). Swiss), the same index as PI and RG (fast-growth and In addition, rib joint dissection overestimates the fat well-conformed breeds), sacrificed at 460 kg. proportion in comparison with half carcass dissection Concerning the fatness scores, the moderate fatness (Oliva´ n et al., 2001). level observed in the present study would make it pos- With regard to meat quality characteristics, the LT sible to rear BP young bulls to higher weights while expressible juice percentage found in BP young bulls maintaining good carcass quality. Conformation and was slightly lower than reported by Albertı´ et al. (1995) fatness scores in PI were one point below the BP scores for PA and PI in seven-day-aged LT. San˜ udo et al. according to the results presented by Piedrafita et al. (1998) did not find significant differences in expressible (2003). Font, Pi, Garcia-Macias, Guerrero, and Oliver juice between PA and other local Spanish beef breeds, (1995) found conformation and fatness scores of 8.7 and intensively reared. Similarly, Oliva´ n et al. (1999),ina 6.6, respectively. The lower conformation scores com- study with AM and AV breeds intensively reared, pared to our study could be explained by the lower car- reported significantly lower LT expressible juice for AM cass weight (249 vs. 329 kg) and the presence of females, (non-improved type breed) than for AV (a meat breed which reach lower weights and higher fatness, at a given with double-muscled animals). These results suggest age, than males (Kempster, Cuthbertson, & Harrington, that leaner breeds have lower water holding capacity, as 1982b). Albertı´ et al. (1997) found lower conformation occurs in leaner pigs as a consequence of the presence of and fatness scores for PA/BP yearlings than our BP the halothane gene (Oliver, Gispert, & Diestre, 1993). young bulls, probably due to their lower slaughter Regarding cooking loss, San˜ udo et al. (1998) did not find weight (471 vs. 541 kg). Also, Campo et al. (1999) found significant differences between seven local Spanish beef a lower conformation score for PA yearlings (SW=473 breeds intensively reared: AV, A-NI, MO, PA, PI, RE kg), but with the same fatness value as in our study. and RG. Vestergaard, Therkildsen, Henckel, Jensen, X. Serra et al. / Meat Science 66 (2004) 425–436 433

Andersen, and Sejrsen (2000) studied the effect of two Gil et al. (2001) found the BP breed had a haem pig- different rearing systems on meat quality of Friesian ment concentration similar to PI (a meat breed) and calves at two slaughter weights (360 and 460 kg) and significantly lower than non-improved type breeds such found lower cooking losses in intensively reared animals as AM, RE and MO. San˜ udo et al. (1998) reported (135 g/kg), which also showed significantly higher similar pigment contents for PA, RE and A-NI breeds, intramuscular fat contents, compared with extensively with the MO breed showing a significantly higher con- reared animals (150 g/kg). However, no effect of weight tent (P<0.05), whereas the lowest values (P<0.05) were at slaughter was observed on cooking loss. found in AV, RG and PI, large-sized meat breeds. Muscle colour depends on the pigment concentration Collagen solubility of BP young bulls was lower than and its physico-chemical state, as well as on the muscle found by Campo (1999) in PA animals (49.8%; 473 kg structure (Renerre, 1982). As discussed previously (Gil et live weight). As collagen solubility decreases with age al., 2001), meat from our BP young bulls was significantly (Hill, 1966; Sims & Bailey, 1981), this could explain the darker (lower L* values) than meat from PI and AV (meat low values of soluble collagen (25.2%) found by type breeds) and A-NI (medium-sized non-improved Renand et al. (1997, 2001) in Charolais calves slaugh- breed), although a* values (redness component) were not tered at 17 months in comparison with the BP breed. In significantly different between these breeds. In a study with addition, these authors reported a lower IMF propor- seven local Spanish beef breeds intensively reared Albertı´ tion (17.8 g/kg) than that of the BP breed, despite the et al. (1999) classified PA meat as the darkest and reddest age differences, because the Charolais is a later-matur- (LT) together with the non-improved breeds (A-NI, MO ing meat breed and is leaner and fattens later than the and RE). These four breeds differed from meat breeds AV, BP breed. PI and RG which gave lighter and less red meat. In agree- According to the biochemical characteristics and the ment with these findings, Insausti, Beriaı´n, Purroy, colour variables the BP was placed in an intermediate Albertı´, Lizaso, and Herna´ ndez (1999) also described PA position in relation to the other local Spanish beef cattle and MO as producing the darkest meat colour 1 h after breeds (Gil et al., 2001). The MHC I% variability of BP cutting in a study that also included AV, PI and RG. breed observed was in agreement with the results BP young bulls showed an intermediate IMF level reported by several authors on MHC I isoform quanti- with respect to the other local Spanish beef cattle breeds fication with ELISA (Jurie, Robelin, Picard, Renand, & (Gil et al., 2001). Font et al. (1997) reported an IMF Geay, 1995; Picard et al., 1994; Renand et al., 1997, value similar to that in the present study, although their 2001). In their study with the Charolais breed (animals animals (bulls and heifers) were younger. Greater dif- aged 15–18 months), Picard et al. (1994) found 38.1% ferences in IMF concentration would have been expec- MHC I with a CV of 29.3%. On the other hand, Jurie et ted between the two studies, as IMF increases with age al. (1995) described a similar MHC I percentage in (Maltin et al., 1998; Renand, Picard, Touraille, Berge, & Limousin young bulls (27%, CV=30%; slaughter Lepetit, 2001; Renand, Touraille, Geay, Berge, Lepetit, weight 591 kg) to that of the BP breed. Renand et al. & Picard, 1997; Vestergaard et al., 2000). However, the (1997, 2001), in a study with Charolais bulls (aged small IMF differences observed could be explained by between 15 and 19 months), reported a similar MHC I the presence of heifers (25%) in their study, since isoform percentage (25.3%, CV=30.5%) in LT muscle, females have a higher IMF level than males at the same while both enzymatic activities were higher than in the age (Kempster et al., 1982b). In other studies with BP breed (LDH=1302 mmol/min/g, CV=10.3% and intensively reared young bulls from local Spanish beef ICDH=1960 nmol/min/g, CV=22.6%). However, the breeds, higher IMF concentrations were found for the LDH/ICDH ratio was lower (0.66 mmol/nmol) showing PA breed (Campo et al., 1999; San˜ udo et al., 1999). In a higher oxidative capacity of LT muscle from the both studies the PA slaughter weights (460 and 473 kg) Charolais animals. were lower than in our study, and therefore would not The study of the relationships between meat quality explain their higher IMF contents. This may relate to and biochemical variables showed that the most impor- the different energy levels in the diets used. Some studies tant variables, which explained the greatest portion of with dual purpose and indigenous cattle breeds have the total variance, were the colour variables (b*, chroma reported lower IMF levels than BP. Aass (1996) found and hue), the expressible juice and the glycolytic activity 12.5 g/kg wet muscle IMF concentrations in dual pur- LDH in the first PC. The muscle oxidative capacity pose Norwegian cattle (13.4 months age; 400 kg live represented by haem pigments content and ICDH weight), while Strydom, Naude, Smith, Scholtz, and van activity, together with muscle redness and total and Wyk (2000) reported IMF levels ranging from 5.0 to 6.1 insoluble collagen contents were found in the second g/kg wet muscle in indigenous African breeds (Afrika- PC. While the most important variables in the third PC ner and Ngumi, respectively) compared with 16.2 g/kg were the muscle glycolytic capacity and muscle redness in Brown Swiss (with young bulls slaughtered at 300– which correlated negatively with moisture content and 460 kg and aged 413 months). oxidative characteristics (ICDH and MHC I). Jurie et 434 X. Serra et al. / Meat Science 66 (2004) 425–436 al. (1995) obtained a similar distribution for LT from in Portuguese autochthonous Maronesa bulls. Vester- (slaughtered at 16 months), with a PCA gaard et al. (2000) described significantly lower tender- showing ICDH activity and MHC I% (oxidative meta- ness and juiciness panel scores (P<0.001 and P<0.006, bolism) negatively correlated to LDH activity (glycoly- respectively) in extensively reared Friesian bulls com- tic metabolism). In this regard, Talmant, Monin, pared with intensively reared bulls which showed sig- Briand, Dadet, and Briand (1986) reported, by means of nificantly lower cooking losses (P<0.003). PCA, a clear distinction between glycolytic metabolism The lack of significant correlations between overall (myofibrillar ATPase and glycolytic potential) and oxi- tenderness and juiciness with IMF and other biochem- dative metabolism (citrate sintase and haem iron) in a ical characteristics of LT would indicate that meat sen- study with 18 bovine muscles (including longissimus sory quality is affected, to a large extent, by such ante dorsi) from Charolais and Friesian cattle (females, males and post mortem factors as ageing and cooking regime. and castrated males, with ages ranging between 2 and 10 In this respect, Gregory et al. (1995), in a study with sev- years). eral purebred and composite cattle breeds, found low In the present study intramuscular fat content was positive correlations between IMF and the sensory attri- positively correlated with beef odour and beef flavour, butes of tenderness, juiciness and flavour (r=+0.17, although it only explained 5–7% of the variance of these +0.20 and +0.12, respectively). Campo et al. (1999) attributes. However, this observation is in agreement observed that the IMF effect on sensory attributes was with the importance of the lipid fraction in meat, which less important than ageing time. is responsible for the characteristic species-specific fla- vours and aromas (Pearson & Young, 1989). In this respect, Renand et al. (2001) also described positive 5. Conclusions correlations between lipid content and flavour intensity in steaks at 6 and 15 days post mortem. In their study, The BP young bulls showed a high average growth Charolais young bulls (slaughtered at 17 months) rate and good carcass quality traits (i.e. dressing pro- showed a lower IMF content (17.8 g/kg, with a high portion, conformation and fatness scores) like other CV=56%) in comparison to BP yearlings. local breed-production systems, e.g. Pirenaica and On the other hand, livery flavour intensity showed a Rubia Gallega (meat breeds). The BP moderate carcass positive correlation with overall tenderness. Moreover, fatness would enable the rearing of bulls to higher livery flavour related positively with several carcass weights while still maintaining good carcass quality. traits that indicated a lower carcass quality (i.e. higher Carcass fatness score and intramuscular fat propor- fat content and lower conformation score and lean tion were the variables that positively affected meat beef proportion). However, no significant correlations were odour and flavour. On the other hand, livery flavour observed for livery odour. A reason for these results intensity was higher in meat from carcasses of a lower could be the greater difficulty in the evaluation of this quality (i.e. lower conformation score, lower muscle attribute by the panellists, although livery odour proportion, higher overall dissectable fat). Undoubt- showed a significant correlation with livery flavour. edly, it will be important to ascertain consumer pre- The two texture attributes, overall tenderness and ferences with respect to livery flavour in meat, and overall juiciness were positively correlated. This rela- therefore we suggest a consumers’ study be carried out tionshiphas been reportedin several studies ( Campo, to consider their preferences when selecting animals to 1999; Crouse, Cross, & Seideman, 1985; Destefanis, improve meat quality. Barge, Brugiapaglia, & Tassone, 2000; Gregory, Cun- diff, & Koch, 1995; Silva, Patarata, & Martins, 1999) with correlation coefficients ranging between r=+0.42 Acknowledgements and +0.88. The more tender the meat, the more quickly the juices are released by chewing and the more juicy the The authors thank Mr A. Quintana and Mrs M. J. meat appears (Cross, 1988). Bautista for their helpin the collection of carcass data Cooking loss was found to negatively affect overall and samples, Mrs. M.D. Gua` rdia for her collaboration tenderness and juiciness. Such findings are in agreement in the sensory analysis and Mr. R.M. Pellicer for animal with other studies (e.g. Destefanis et al., 2000; Silva et care and supervision. This research was financed by the al., 1999). In their study with several hypertrophied and EU–FAIR1CT950702 project. normal beef breeds, Destefanis et al. (2000) described negative correlations between cooking loss and tender- ness (P<0.01). On the other hand, Silva et al. (1999) References reported negative correlations between cooking loss and Aass, L. (1996). Variation in carcass and meat quality traits and their sensorial tenderness (P<0.001) at 1, 6 and 13 days post relations to growth in dual purpose cattle. Livestock Production mortem (r=À0.84, r=À0.64 and r=À0.76, respectively) Science, 46, 1–12. X. Serra et al. / Meat Science 66 (2004) 425–436 435

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