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Meat Science 77 (2007) 459–466

MEATSCIENCE

Muscle and meat quality characteristics of Holstein and Salers cull cows

C. Jurie a,*, B. Picard a, J.-F. Hocquette a, E. Dransfield b,1, D. Micol a, A. Listrat a

a INRA, UR1213, Unite de Recherches sur les Herbivores, Theix, 63122 Saint Genes Champanelle, Franceb INRA, UR370, Unite de Recherches sur la Qualite des Produits Animaux, Theix, 63122 Saint Genes Champanelle, France

Received 5 December 2006; received in revised form 11 April 2007; accepted 12 April 2007

Abstract

Muscle characteristics and sensory rating of meat were determined in M. longissimus thoracis (LT), M. semimembranosus (SM), M.

semitendinosus (ST) and M. triceps brachii (TB) from seven Holstein (HO, dairy breed) and six Salers (SA, beef breed) cull cows slaugh-tered at 6–7 years of age at the same fat score. Significant differences (P < 0.001) among muscle types were observed: ST was the moreglycolytic and TB the more oxidative; total collagen: ST > SM = TB > LT; initial and overall tenderness: LT > TB = SM > ST, juiciness:TB > LT = SM > ST. Flavour differed only between breeds: HO > SA (P < 0.01). Three tenderness classes (high, intermediate, low) weredetermined from scores for sensory overall tenderness for all 52 meats: the lower total and insoluble collagen contents, the more oxidativemetabolism, the more tender was the meat. Muscle type, and not breed explained most of the variability of meat quality from dairy andbeef cull cows slaughtered at the same age and fat score.� 2007 Elsevier Ltd. All rights reserved.

Keywords: Cows; Breed; Muscle characteristics; Meat quality

1. Introduction

Tenderness is one of the most important organolepticcharacteristics by which consumers judge meat quality.Meat tenderness is influenced by muscle characteristics atslaughter (Maltin, Balcerzak, Tilley, & Delday, 2003) andby the post-mortem changes induced by ageing (Culioli,1999). Few studies have been carried out to characterizecarcass, muscle and meat quality characteristics from cullcows. Recently, Jurie et al. (2006) described carcass andmuscle characteristics from four French breeds of beef cullcows between 4 and 9 years of age. Dransfield et al. (2003)reported meat eating qualities from these cull cows in com-parison with meats from young bulls. However, to ourknowledge, no study has shown relationships between mus-cle and meat quality characteristics from dairy cull cowswhich are the main origin of meat consumed in France.

0309-1740/$ - see front matter � 2007 Elsevier Ltd. All rights reserved.

doi:10.1016/j.meatsci.2007.04.014

* Corresponding author. Tel.: +33 4 7362 4359; fax: +33 4 7362 4639.E-mail address: jurie@clermont.inra.fr (C. Jurie).

1 Present address: WCFS, Wageningen UR, Agrotechnology and FoodInnovations, Bornsesteeg, 59, 6700AA Wageningen, The Netherlands.

In fact, about three-quarters of the beef consumed inFrance derives from females (20% from heifers and 80%from cull cows) or two-thirds from cull cows, 15% fromyoung bulls and about 10% from steers. In addition, meatproduction from cull cows is provided equally from dairyand beef breeds. Beef consumption in France from cullcows is much higher than in the EU or in the US where cullcows represent only about 30% and 15%, respectively, oftotal beef consumption.

Holstein (HO) breed is the premier dairy breed with ahigh potential for milk production and is spread all overFrance whilst the Salers (SA) breed shows excellent mater-nal aptitudes, hardiness, and satisfactory milk yield (Petit& Lienard, 1988) and beef qualities (Dransfield et al.,2003). So, SA breed can be considered as a mixed-breedof which 92% of the French SA herd is located in the Mas-sif Central region, but the animals are the more often raisedfor meat production and so they will be considered as abeef breed in this study.

The aim of this study was to report muscle characteris-tics and sensory qualities of meat from four muscles sam-pled from these two different breeds of cull cows: the

460 C. Jurie et al. / Meat Science 77 (2007) 459–466

Holstein (HO) dairy breed and the Salers (SA) beef breedand was to investigate the relationships between musclecharacteristics and meat tenderness through the determina-tion of meat quality classes.

2. Materials and methods

2.1. Animals

Seven HO cull cows and six SA cull cows were finishedat the experimental farm of the Herbivore Research Unit ofthe National Institute for Agricultural Research (INRA,Institut National de la Recherche Agronomique) of Cler-mont-Ferrand/Theix, France. The animals were slaugh-tered at the same fat score assessed manually on the backof the live animal (average score 3.5 on a scale from 1, verythin, to 5, very fat; Agabriel, Giraud, & Petit, 1986). Thefinishing periods were of 7.5 ± 1.1 and 6.1 ± 0.6 weeksfor HO and SA cull cows, respectively. At slaughter, theanimals were 6.6 ± 0.2 years old. All the animals receivedthe same diet, ad libitum, consisting of 850 g/kg overpres-sured sugar-beet pulp silage and hay, and supplementedwith maize, soya-bean cake and urea. During the finishingperiod, the animals were housed in loose-housing by breed,and weighed individually every 2 weeks and double-weighed on two consecutive days.

Animals were slaughtered at the experimental slaughter-house of the Research Centre in compliance with ethicalguidelines for animal care. The final and carcass weightswere recorded. At 24 h post-mortem the sixth ribcut wasdissected into muscle, fat and bone, to determine the carcasscomposition (muscle, fat and bone) using the regressionequations as described by Robelin and Geay (1975). Musclesamples were removed from the centre of the M. triceps bra-

chii (TB) and of the M. longissimus thoracis (LT) at the sixthribcut, and also in the centre of the superficial portion of M.

semimembranosus (SM) and M. semitendinosus (ST). Mus-cle samples were taken within 15 min after exsanguination:for histochemical analysis, they were frozen in isopentanechilled in liquid nitrogen, and for determination of meta-bolic enzyme activities they were immediately frozen inliquid nitrogen. Both types of samples were stored at�80 �C until analysed. The carcasses were placed in a coldroom (2 �C) and muscle samples from the same muscleswere taken at 24 h post-mortem for the determination ofcollagen characteristics: they were cut in pieces (4–5 cmcross-section), sealed under vacuum, and stored at �20 �Cuntil analysis. For meat quality evaluation, samples werealso taken at 24 h post-mortem and stored in vacuum-packsat 4 �C for 14 days. The samples were collected from thesame side of the carcass (the left) for both muscle fiber char-acteristics and sensory traits.

2.2. Muscle histochemical analysis

Serial cross-sections of 10 lm thickness were cut witha cryotome (HM 500M, MICROM, France) at �20 �C.

The fibres were stained with azorubine to delineate theiroutline. The fibre contractile type was determined bydemonstrating myofibrillar mATPase activity after acidpre-incubation at pH 4.2 for 15 min, according to Guthand Samaha (1970) and adapted for bovine muscle(Picard, Duris, & Jurie, 1998). The metabolic type wasdetermined by demonstrating succinate dehydrogenase(SDH) activity according to Pearse (1968). Fibres wereidentified according to their ATPase and SDH reactivitiesand classified according to Peter, Barnard, Edgerton, Gil-lepsie, and Stempel (1972) as slow oxidative (SO), fastoxidative glycolytic (FOG) or fast glycolytic (FG). Thecross-sectional area (lm2) and the frequency (expressedas the percentage number of each muscle fiber type tothe analysed total fiber number) were quantified in tworandomly selected areas in each series of section withan image analysing software program (Visilog, NOESIS,France) for the four muscles. In addition, the mean fibrearea for each sample [(SO% frequency * SO area +FOG% frequency * FOG area + FG% frequency * FGarea)/3] and the area percentage (expressed as the areapercentage of each muscle fibre type relative to the ana-lysed total fibre area, for example SO% area = (SO% fre-quency * SO area)/(SO% frequency * SO area + FOG%frequency * FOG area + FG% frequency * FG area) werecalculated. A minimum of 200 fibres per sample wasexamined.

2.3. Measurement of enzyme activities

Two hundred mg of frozen muscle sample was groundand homogenized with a polytron in 4 ml of 140 mMsucrose and 50 mM triethanolamine pH 7.5 buffer and cen-trifuged at 6000g for 15 min at 4 �C. Enzyme activities weremeasured in the supernatant. Anaerobic glycolytic metabo-lism was studied by measuring lactate dehydrogenase(LDH, EC 1. 1. 1. 27) activity in the transformation ofpyruvate into lactate according to Ansay (1974). Aerobicoxidative metabolism was studied by measuring isocitratedehydrogenase (ICDH, EC 1. 1. 1. 42) NADP activityaccording to Briand, Talmant, Briand, Monin, & Durand(1981). These two protocols were described in Jurie, Rob-elin, Picard, Renand, & Geay (1995). Enzyme activities(means of triplicate) were expressed in lmole per min pergram of wet muscle.

2.4. Collagen characteristics

Frozen muscle was homogenized in a household cutter,freeze-dried for 48 h, pulverized in a horizontal blade milland stored at �20 �C in stoppered plastic flasks until ana-lysed. Total collagen and collagen in the insoluble partwere measured as described by Listrat et al., 1999 andupdated by Listrat & Hocquette (2004). The data (meansof triplicates) were expressed in lg of hydroxyproline(OH-proline) per mg of dry matter for both total and insol-uble collagen.

Table 1Slaughter characteristics from Holstein (HO) and Salers (SA) cull cows

Slaughter characteristics HO SA s.e.a Breed effect P-values

No. of cows 7 6Final live weight (kg) 666 626 28.9 (31.2) 0.36Carcass weight (kg) 356 362 14.6 (15.8) 0.79

Carcass compositionb:Fat (kg) 109 84 11.4 (12.3) 0.17Muscle (kg) 190 225 6.9 (7.4) 0.006Bone (kg) 57 53 1.4 (1.5) 0.08

Offal fat (kg) 43 28 5.0 (5.4) 0.07Total fatty tissue (kg) 152 112 16.3 (17.6) 0.13

a s.e. for HO cull cows out of brackets and for SA cull cows betweenbrackets.

b Carcass composition is based on an estimation from the dissection ofthe sixth ribcut in muscle, fat and bone (Robelin & Geay, 1975).

C. Jurie et al. / Meat Science 77 (2007) 459–466 461

2.5. Meat quality evaluation

After 14 days at +4 �C, the chilled muscles were cut intosteaks (about 1.5 cm thick), packed under vacuum andplaced at �20 �C until sensory evaluation. For each sen-sory session, eight frozen steaks (4 muscles · 2 breeds) werethawed at room temperature and then grilled between twocontact plates heated to 310 �C and the steaks grilled for2 min giving an internal temperature between 55 and60 �C. A total of seven sessions were performed. The grilledsamples were presented two at a time (2 breeds · 1 muscle)and the quality evaluated as five attributes (‘initial tender-ness’ assessed over the first few bites, ‘overall tenderness’juiciness’, ‘residue’ after chewing and ‘flavour intensity’)following the protocol defined by Dransfield et al. (2003).The five attributes described above rated by 9–11 experi-enced panellists on non-structured line scales marked atthe extremities ‘low’ and ‘high’ and subsequently scoredas the distance in units of 1, from 0 to 10. As the composi-tion of the panel changed throughout the different sessions,only panel average ratings were analysed.

2.6. Statistical analysis

Data were analysed using the GLM procedure of SAS(1996). For slaughter characteristics, the effect of breedwas included in the model. For muscle characteristics andquality evaluation, the effects tested in the model includedbreed, animal nested within breed, muscle and breed ·muscle interaction. The breed effect was tested against ani-mal within breed. The residual mean square was used as theerror term for other effects. All results were presented asadjusted means with appropriate standard errors of means(s.e.). Differences between adjusted means were comparedusing the PDIFF option of SAS. No breed · muscle inter-actions were observed, so only main effects are reported intables. In addition, a disjoint (three solutions) cluster anal-ysis was realized using the FASTCLUS procedure of SASto produce three quality classes from overall tendernessof all meats (tenderness classes). The values of all variableswithin each of the three classes were analysed by GLM pro-cedure and their significance between classes was deter-mined by the PDIFF option.

3. Results

3.1. Breed effect on slaughter and muscle characteristics

The final live weights and carcass weights averaged 646and 359 kg, respectively and did not (P > 0.10) differbetween breeds. The SA cows had 35 kg more muscle thanHO cows (P < 0.01). In contrast, HO cows tended to havemore bone (+4 kg, P = 0.08), offal fat (+16 kg, P = 0.07),total fat (+25 kg, P = 0.17) and total fatty tissue (+40 kg,P = 0.13) than SA cows (Table 1).

No significant effect of breed was observed for any of thestudied muscle characteristics (% frequency, area and %

area of the different types of fibre, metabolic enzyme activ-ities and total and insoluble collagen contents) (Table 2).

3.2. Characteristics of different muscle types

Fibre type characteristics, metabolic enzyme activitiesand collagen contents differed significantly among theLT, SM, ST and TB muscles (Table 2).

The FOG fibre type differed less between muscles thandid the SO and FG fibre types. FOG area was not differentamong muscles, and FOG% frequency and % area werelower (P < 0.05) in LT than in SM, ST and TB muscles.On the other hand SO and FG characteristics were signif-icantly different among muscles, and these differences werein accordance with the differences in oxidative (ICDH) andglycolytic (LDH) enzyme activities. In fact, the SO% fre-quency and % area, and ICDH activity were lower(P < 0.001) in ST than in SM, LT and TB muscles butthe SO area was greater (P < 0.05) for ST than for theother muscles. Indeed, the SO area was 1.3-fold greaterand the SO% frequency was 2.3-fold lower, so SO% area(expressed as the area percentage of SO fibre type to thetotal fibre area) was 2.4-fold lower in ST than in LT, SMand TB muscles. Furthermore, the FG fibre type character-istics (% frequency, area, % area), and LDH activity werehigher (P < 0.001) in ST than in LT, SM and TB muscles.Inversely, ICDH activity was the highest, and the FG% fre-quency and area, and LDH activity were the lowest in TBmuscle. The SO% frequency and area were the highest bothin TB and LT muscles. Total and insoluble collagen con-tents were the lowest in LT compared to the other threemuscles. Total collagen content was the highest in STand insoluble collagen content was not different amongSM, ST and TB muscles.

3.3. Sensory rating of meat from cull cows

The average ratings for the five sensory attributes ofeach breed and for each muscle are given Table 3. There

Table 2Influence of breed and muscle on muscle characteristics of cull cows

Muscle characteristics Breeda Muscleb Significance of effect P-values

HO SA s.e. LT SM ST TB s.e. Breed Muscle

No. of observations 28 24 13 13 13 13Fibre characteristicsc

% Frequency

SO 19.0 20.6 1.2 (1.3) 24.6a 18.9b 10.3c 25.5a 1.7 0.37 0.001FOG 22.4 22.9 0.9 (1.0) 18.6b 24.2a 24.1a 23.8a 1.4 0.73 0.02FG 58.6 56.5 1.5 (1.6) 56.8b 56.9b 65.6a 50.8c 1.9 0.36 0.001

Area (lm2)

SO 2662 2524 188 (203) 2364b 2423b 3074a 2510b 175 0.63 0.03FOG 3293 3013 184 (198) 3146 2923 3471 3071 171 0.32 0.16FG 4033 3757 169 (182) 3816b 3466b 4762a 3538b 212 0.29 0.001Mean area 3623 3301 124 (134) 3314b 3107b 4250a 3175b 161 0.11 0.001

% Area

SO 14.1 15.7 1.0 (1.1) 17.9ab 14.2b 7.3c 20.2a 1.6 0.33 0.001FOG 20.5 20.8 0.9 (1.0) 17.5b 22.8a 19.5ab 22.9a 1.5 0.80 0.04FG 65.4 63.5 1.9 (2.0) 64.6b 62.9b 73.2a 56.9c 2.1 0.50 0.001

Metabolic enzyme activities (lmol min�1 g�1)d

ICDH 1.49 1.48 0.08 (0.09) 1.37b 1.76a 0.79c 2.01a 0.10 0.97 0.001LDH 816 837 26.0 (28.1) 835b 821b 893a 757c 19.7 0.59 0.001

Collagen contents (lg OH-proline mg�1 dry matter)

Total 4.48 4.10 0.15 (0.16) 3.21c 4.63b 5.19a 4.15b 0.18 0.10 0.001Insoluble 3.22 3.39 0.13 (0.14) 2.62b 3.57a 3.71a 3.32a 0.16 0.40 0.001

LSmeans in the same row for muscle effect not followed by a common letter (a–c) differ significantly (P < 0.05).a Breed designation: HO = Holstein; SA = Salers; s.e. for HO cull cows out of brackets and for SA cull cows between brackets.b Muscle designation: LT = longissimus thoracis; SM = semimembranosus; ST = semitendinosus; TB = triceps brachii.c Fibre designation: SO = slow oxidative, FOG = fast oxidative glycolytic, FG = fast glycolytic.d Enzyme designation: ICDH = isocitrate dehydrogenase; LDH = lactate dehydrogenase.

Table 3Influence of breed and muscle on sensory quality of meat from cull cows

Sensory attributes Breeda Muscleb Significance of effect P-values

HO SA s.e. LT SM ST TB s.e. Breed Muscle

No. of observations 28 24 13 13 13 13Initial tenderness 5.49 5.27 0.17 (0.17) 6.80a 5.18b 4.27c 5.27b 0.24 0.36 0.001Overall tenderness 5.23 4.95 0.18 (0.18) 6.52a 4.86b 3.87c 5.11b 0.23 0.30 0.001Residue 4.66 5.19 0.25 (0.25) 3.68b 5.23a 5.66a 5.13a 0.22 0.17 0.001Juiciness 5.35 5.29 0.10 (0.10) 5.41ab 5.32b 4.68c 5.88a 0.17 0.65 0.001Flavour 5.78 5.17 0.12 (0.12) 5.48 5.69 5.17 5.55 0.16 0.005 0.17

Values are the LSmeans of sensory panel averages for each attribute (scored from 0 to 10). LSmeans in the same row for muscle effect not followed by acommon letter (a–c) differ significantly (P < 0.05).

a Breed designation: HO = Holstein; SA = Salers; s.e. for HO cull cows out of brackets and for SA cull cows between brackets.b Muscle designation: LT = longissimus thoracis; SM = semimembranosus; ST = semitendinosus; TB = triceps brachii.

462 C. Jurie et al. / Meat Science 77 (2007) 459–466

were no significant effects of breed for initial tenderness,overall tenderness, residue and juiciness, only flavour wasdifferent between breeds: it was higher for meat from HO(rating 5.78, on a scale from 0 to 10) than SA (rating5.17) cows (P < 0.01). Combining both breeds, importantvariations on initial and overall tenderness, residue andjuiciness were consistent across muscles (P < 0.001), onlyflavour was not different between muscle types (on average,rating 5.47). The LT was the most tender (rating 6.80 and6.52 for initial and overall tenderness, respectively) of thefour muscles and the ST was the toughest (rating 4.27

and 3.87 for initial and overall tenderness, respectively).The TB and LT were the juiciest (rating 5.88 and 5.41,respectively) and the ST the least juicy (rating 4.68).

3.4. Tenderness classes

Only the sensory rating for the overall tenderness fromall meats (including all animals and the four different mus-cle types) was used to produce three tenderness classes. Thelowest tenderness class (Class 1, containing 27% of themeats) had an average rating (on a scale from 0 to 10) of

Fig. 1. Percentage distribution of meats within their tenderness class(Class 1 = lowest, Class 2 = intermediate, Class 3 = highest tendernessclass. Breed designation: HO = Holstein; SA = Salers. Muscle designa-tion: LT = longissimus thoracis; SM = semimembranosus; ST = semitendi-

nosus; TB = triceps brachii).

C. Jurie et al. / Meat Science 77 (2007) 459–466 463

3.63 (±0.46). The intermediate tenderness class (Class 2,containing 53% of the meats) had an average rating of5.18 (±0.62). The highest tenderness class (Class 3, contain-ing 20% of the meats) had an average rating of 6.86(±0.49). Classes were significantly different at P < 0.001for the overall tenderness.

The types of meats within the three tenderness classesare given in Fig. 1. The lowest tenderness class (Class 1)was mainly composed of ST (62%) and did not containany LT muscle samples. SM and TB muscle samples repre-sented 23% and 15% of the meats in Class 1, respectively.The opposite order was found in the highest tendernessclass (Class 3), which was mainly composed of LT samples(70%), and did not contain any ST muscle. SM and TBmuscle samples represented 20% and 10% of the meats inClass 3, respectively. The intermediate tenderness class(Class 2) was composed of TB (35%), SM (31%), LT(19%) and ST (15%) muscle samples. The distributionswithin the tenderness classes with respect to breed wereone third for HO breed and two-thirds for SA breed, in

Table 4Muscle characteristics among the three tenderness classesa

Muscle characteristicsb Class 1

Mean s

Insoluble collagen content (lg OH-proline mg�1 dry matter) 3.78a 0Total collagen content (lg OH-proline mg�1 dry matter) 4.87a 0LDH/ICDH ratio 894a 8SO% frequency 14.6b 2LDH (lmol min�1 g�1) 872a 2ICDH (lmol min�1 g�1) 1.17b 0FOG% frequency 24.9a 1SO area (lm2) 2949a 2

a The class average values and standard errors, F ratio and significance, foclasses (1 = lowest, 2 = intermediate, 3 = highest tenderness class) are given. Wdiffer significantly (P < 0.05).

b Fibre designation: SO = slow oxidative, FOG = fast oxidative glycolytic;dehydrogenase.

both Class 1 and Class 3. The opposite distribution wasfound for Class 2 with two-thirds for HO breed and onethird for SA breed. In addition, it is noteworthy that100% of ST samples and 83% of LT samples from SA cowswere found in Class 1 and Class 3 respectively, whereas STand LT samples from HO breed were found both in Class 1(33%) and Class 2 (67%) for ST samples, and both in Class2 (67%) and Class 3 (33%) for LT samples.

The distributions of the muscle characteristics were cal-culated across the three tenderness classes. Those whichdiffered significantly are given in Table 4. The insoluble col-lagen content was the only muscle characteristic which dis-tinguished the three tenderness classes: Class 1 > Class2 > Class 3. This order was the same for the total collagencontent but the Classes 2 and 3 did not differ significantlyfor this parameter. The lowest tenderness class (Class 1)was characterized by a higher LDH/ICDH ratio and alower SO% frequency compared to the other classes. It alsocontained muscles with higher FOG% frequency and largerSO area compared to the highest tenderness class (Class 3).

4. Discussion

4.1. Differences between breeds

Few differences in slaughter characteristics wereobserved between HO and SA breeds: the SA cows hadmore muscle in carcass and the HO cows were fatter thanSA cows in association with numerically higher weightsof fats in carcass, of offal fat and of total fatty tissue,although these differences in fats were not significant. Sim-ilarly, differences in carcass composition can be foundwhen comparing certain breeds, particularly between dairy(Holstein) and beef (Blue Belgian) breeds (Istasse et al.,1990). The proportion of muscle in the carcass was lower,whereas the proportions of adipose tissue and bone werehigher in the Holstein than in the Blue Belgian bulls. Somedifferences in carcass composition were also observedbetween beef breeds only (Aubrac, Charolais, Limousin,

Class 2 Class 3 F ratio P-values

.e. Mean s.e. Mean s.e.

.18 3.25b 0.13 2.77c 0.20 7.10 0.002

.26 4.32b 0.18 3.67b 0.30 4.62 0.018 576b 63 628b 101 4.41 0.02.3 20.5a 1.6 24.1a 2.6 4.10 0.026 791b 19 845a 30 3.54 0.04.15 1.66a 0.11 1.42b 0.17 3.52 0.04.5 23.1a 1.1 19.5b 1.8 2.74 0.0704 2418b 144 2359b 232 2.67 0.08

r those characteristics differing significantly (by GLM procedure) amongithin a muscle characteristic, means not followed by a common letter (a–c)

Enzyme designation: ICDH = isocitrate dehydrogenase; LDH = lactate

464 C. Jurie et al. / Meat Science 77 (2007) 459–466

Salers) for both young bulls (Jurie et al., 2005) and cullcows (Jurie et al., 2006). At a same age of slaughter, thefinal live weight of the Charolais animals was higher thanthat of the other breeds. The Limousin animals were char-acterized by a higher muscle weight and lower fat and boneweights. In the current study, the proportion of muscle incarcass was 62% for SA cows and only 53% for HO cows.The differences in the muscle proportion in the carcass canbe explained, at least in part, by variability in fibre hyper-trophy and/or hyperplasia. Indeed, Bellman, Wegner, Teu-scher, Schneider, & Ender (2004) reported differences inmuscle type cross-sectional area between Charolais andHolstein bulls in M. semitendinosus. These authors con-cluded that the nutrients transformed into protein in accre-tion-types of cattle, such as Charolais, during growth leadto a higher enlargement of muscle fibres than in secretion-types of cattle, such as Holstein. But in the current study,the mean fibre area, as well as areas of SO, FOG, FG fibreswere not different between SA and HO cows. So, we maysuppose that HO cattle may have a lower number of fibresat birth than SA cattle. In fact, during postnatal growth,both hyperplasia (DNA accumulation by the fusion ofsatellite cells) and hypertrophy cause an increase in musclesize (Jurie et al., 1995), while the number of muscle fibresremains constant, the total number of fibres being fixedfrom the end of the second third of gestation in bovine(Picard, Lefaucheur, Berri, & Duclos, 2002). Although thishypothesis has not yet been verified, it would explain thesimilar fibre mean area and a different muscle proportionin carcass between HO and SA cows. Differences in thenumber of fibres at birth were described by Wegner et al.(2000): Blue Belgian bulls, so-called double muscled cattle,have twice the number of fibres of other breeds (Angus,Galloway and Holstein), and that was responsible forhigher muscle weights. In accordance, Maltin et al. (2001)reported a higher number of fibres and a higher whole areaof M. longissimus lumborum for Charolais compared withHolstein steers, whereas the mean fibre area was not differ-ent between breeds. The differences in muscle weights incarcass between HO and SA breeds may have resulted froma higher number of muscle fibres for SA animals, the meanfibre area being not different between breeds.

The amount of total tissue fat in the carcass, whichshowed the opposite effect to the proportions of musclein the carcass, was higher for HO cows (30%) than SA cows(23%). Correspondingly, for all 4 muscles pooled, HO hada slightly more intense flavour than meat from SA cows.Thus, this observation would suppose some differences inintramuscular fat contents between breeds and could thusexplain the observed significant difference in meat flavourbetween breeds. In fact, a correlation coefficient ofr = +0.35 was reported by Renand, Picard, Touraille,Berge, & Lepetit (2001) between meat flavour intensityand intramuscular lipid content. Although the intramuscu-lar fat content was not determined in this study, differencesin lipid content among muscles would be expected, withLT = TB > ST = SM (Stolowski et al., 2006) and more fla-

vour in muscles with a higher fat content (Dransfield et al.,2003). But, in addition, there were no significant differencesin flavour among muscles. So, we can suppose that the dif-ferences in flavour between SA and HO breeds may beexplained by differences in other compounds than lipid.Indeed, not only lipids contribute to meat flavour but othercompounds such as heterocyclic, phenolic and S-containingcompounds are important flavour-producing endproducts(Gandemer, 1998). It is noteworthy that the other sensoryattributes of meat (initial and overall tenderness, residue,juiciness), determined after 14 days of ageing, were not dif-ferent between breeds. These results are in line with thoseof Monson, Sanudo, & Sierra (2004) who showed thatthe differences in Warner–Bratzler shear between breedsdisappeared after 14 days of ageing, and concluded thatageing time tended to eliminate the differences in meat tex-ture between breeds.

Our results failed to detect any differences in musclecharacteristics (fibre characteristics, metabolic enzymeactivities and collagen contents) between HO and SAbreeds for cull cows slaughtered at the same age and fatten-ing state. In contrast, metabolic enzyme activities and col-lagen contents differ among four beef breeds (Aubrac,Charolais, Limousin and Salers) in muscles of cull cows(Jurie et al., 2006). Salers and Charolais cull cows exhibitedhigher total and insoluble collagen contents on average forthree muscles than Aubrac and Limousin cull cows. Mon-son et al. (2004) reported higher total and insoluble colla-gen contents for Holstein and Old Brown Swiss than forLimousin and Blonde d’Aquitaine bulls. These resultsmay be explained by differences in maturity as earliermaturing breeds tend to deposit more collagen with agreater proportion of insoluble collagen. Similarly, HO(dairy) breed, known as early maturing breed, and SA(beef) breed, known as earlier maturing beef breed thanother beef breeds (Charolais and Limousin) (Lienardet al., 2002), were not different in collagen contents.

4.2. Differences among muscle types

The studied muscles in these HO and SA cows differed intheir characteristics (fibre characteristics, metabolic enzymeactivities and collagen contents), as previously described inbeef bulls (Jurie et al., 2005) and cows (Jurie et al., 2006).Indeed, in beef bulls, beef or dairy cows, compared to LTand TB muscles, the ST muscle presented the largest fibrecross-sectional area and was considered as the fastest gly-colytic muscle according to its highest FG% frequencyand LDH activity, and its lower SO% frequency and ICDHactivity. The TB muscle was a more oxidative muscle thanLT according to higher FOG% frequency and ICDH activ-ity and lower FG% frequency and LDH activity. The SMmuscle, not studied in these previous studies, is a motormuscle, localized deeper in the hindquarter than the STmuscle. SM muscle was more oxidative as indicated byhigher SO% frequency and % area, a higher ICDH activity,and inversely lower FG% frequency and % area, and a

C. Jurie et al. / Meat Science 77 (2007) 459–466 465

lower LDH activity than ST muscle. These observations fitwith the fact that type I (slow oxidative) fibres becomemore abundant from superficial to deep regions, while typeIIB (fast glycolytic) fibres have the opposite distribution(Totland & Kryvi, 1991). In fibre characteristics and meta-bolic enzyme activities, SM muscle was intermediatebetween LT and TB muscles, and similar to TB muscle incollagen contents. LT muscle presented the lowest totaland insoluble collagen contents compared with the otherstudied muscles in agreement with previous studies (Jurieet al., 2005, 2006; Stolowski et al., 2006).

4.3. Relationships between muscle characteristics and meat

quality

All muscles were pooled and then classified as high,intermediate and low in overall tenderness to analyse rela-tionships between muscle characteristics and meat qualitytraits. There were no important differences in tendernessclasses between the two different breeds. This observationis similar to that of Dransfield et al. (2003), who comparedfour beef breeds. On the other hand, the distribution ofmuscles varied among the tenderness classes. Thus, the cur-rent study is in accordance with the studies of Dransfieldet al. (2003) and Stolowski et al. (2006) which showed thatthe muscle type played the greatest role in determining ten-derness classes. But among the muscle characteristics, totaland insoluble collagen contents were best in differentiatingthe tenderness classes. Previous work had shown that tri-glyceride and collagen contents were the best discrimina-tors of quality classes (Dransfield et al., 2003).Insolublecollagen content also distinguished the three tendernessclasses in this current work. It is known that connective tis-sue does not change during ageing (Monson et al., 2004),but determined the basal toughness of beef. So the differ-ences in collagen contents between muscles were still pres-ent after 14 days of ageing and are likely to explain a largepart of the variability in tenderness. In addition, LDH/ICDH ratio and FOG% frequency were lower, and SO%frequency were higher in the higher tenderness class, andinversely lower in the lower tenderness class. So, it wouldappear that more oxidative muscles are more tender. Sim-ilarly, Dransfield et al. (2003) found that the lowest eatingquality class had the highest FOG% frequency and lowestSO% frequency. Ouali (1990) reported that rate of proteol-ysis post-mortem was higher in the muscles containing pre-dominantly ‘white’ or fast glycolytic fibres. So, the meatageing rate depends of the proportion of white fibres. Butfrom this study, overall tenderness, evaluated at 14 dayspost-mortem (i.e. when ageing is almost completed) wouldbe positively associated with oxidative muscle metabolismthan with glycolytic muscle metabolism. This work sup-ports the finding by Maltin et al. (1998) and Dransfieldet al. (2003) who demonstrated a positive relationshipbetween SO frequency and tenderness. In addition, it isnoteworthy that SO area was higher in the lowest tender-ness class. This result is in accordance with the studies of

Renand et al. (2001) who showed a negative correlationcoefficient between tenderness and muscle fibre area, andDransfield et al. (2003) who showed that muscle fibre areawas lowest in the best eating quality class.

In conclusion, this study has shown that meat tender-ness was not different between Holstein and Salers cullcows, which are dairy and beef breeds respectively, whenslaughtered at the same age and fattening. These resultsare of importance since cull cows represent two-thirds ofbeef consumption in France, and meat production fromthese animals is equally provided by dairy and beef breeds.Moreover the findings are very important for consumerswho generally have a more negative image of meat fromdairy breeds compared to that from beef breeds. Neverthe-less, meats from these cull cows can be distributed into dif-ferent tenderness classes, which rely mainly on the differentproperties of the four studied muscle types. Furthermore,differences between muscles in insoluble and total collagencontents (and to a lower extent in oxidative metabolismand in muscle fibre area) explain, for the greatest part,the repartition of meat samples in the different tendernessclasses. So at 14 days post-mortem, the lower total andinsoluble collagen contents, the more oxidative metabo-lism, and to a lesser extent the smaller the fibre area, themore tender was the meat.

Acknowledgements

The experiment was financed by the ‘Fonds Nationalpour l’Amenagement et le Developpement du Territoire’,France (study 99/000144 December 12th, 1999). Theauthors express their thanks to Rt. Jailler, S. Rudel andthe farm staff for animal management, Rd. Jailler and theslaughter-house staff for animal slaughter, M.C. Bayleand H. Dubroeucq for organisation of sensory sessionsand the staff of the ‘Croissance and Metabolisme du Mus-cle’ team for technical assistance.

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