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Research Collection Doctoral Thesis Growth performance, carcass characteristics and meat palatability attributes in steers of various beef breeds compared at a similar level of intramuscular fat content Author(s): Chambaz, Alain Publication Date: 2001 Permanent Link: https://doi.org/10.3929/ethz-a-004268179 Rights / License: In Copyright - Non-Commercial Use Permitted This page was generated automatically upon download from the ETH Zurich Research Collection . For more information please consult the Terms of use . ETH Library
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Page 1: dspace cover page - Research Collection24556/eth-24556-02.pdf · La qualité de viande a été comparée dans leLDet bicepsfemoris, regio glutea (aiguillette rumpsteak, BF). La composition

Research Collection

Doctoral Thesis

Growth performance, carcass characteristics and meatpalatability attributes in steers of various beef breeds comparedat a similar level of intramuscular fat content

Author(s): Chambaz, Alain

Publication Date: 2001

Permanent Link: https://doi.org/10.3929/ethz-a-004268179

Rights / License: In Copyright - Non-Commercial Use Permitted

This page was generated automatically upon download from the ETH Zurich Research Collection. For moreinformation please consult the Terms of use.

ETH Library

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Diss. ETHNo. 14316

Growth performance, carcass characteristics and meat

palatability attributes in steers of various beef breeds

compared at a similar level of intramuscular fat content

A dissertation submitted to the

SWISS FEDERAL INSTITUTE OF TECHNOLOGY ZURICH

for the degree of Doctor ofNatural Sciences

presented by

Alain CHAMBAZ

Dipl. Ing.-Agr. ETH

born 3 March 1971

citizen of Arzier (VD)

accepted on the recommendation of

Prof. Dr. M. Kreuzer, examiner

Prof. Dr. N. Künzi, co-examiner

Dr. M.R.L. Scheeder, co-examiner

Zürich 2001

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II

Seite Leer /

Blank leaf

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Ill

Table of contents

1. Summary 1

2. Résumé 3

3. General introduction 6

4. Sources of variation influencing the use of real-time ultrasound to predict intramuscular

fat in live beef cattle 8

Abstract 8

4.1. Introduction 9

4.2. Materials and methods 10

4.2.1. Animals 10

4.2.2. Live animal measurements 10

4.2.3. Carcass measurements 12

4.2.4. Statistical analysis 12

4.3. Results and discussion 13

4.3.1. Test and validation of the prediction equation 13

4.3.2. Comparisons of the new equations developed with results of the literature 19

4.3.3. Possible muscle characteristics influencing intramuscular fat estimation by ultrasound 22

4.4. Implications 23

5. Characteristics of steers of six beef breeds fattened from eight months of age and

slaughtered at a target level of intramuscular fat. I. Growth performance and carcass

quality 24

Abstract 24

5.1. Introduction 25

5.2. Materials and methods 26

5.2.1. Animals and experimental design 26

5.2.2. Diet 26

5.2.3. Endpoint of fattening 27

5.2.4. Data and sample collection at slaughter 28

5.2.5. Statistical analysis 29

5.3. Results 29

5.3.1. Growth performance 29

5.3.2. Carcass quality 33

5.4. Discussion 34

5.4.1. Growth development of steers of different breeds fattened to a similar IMF content 35

5.4.2. Carcass characteristics of steers of different breeds fattened to a similar IMF content 37

5.4.3. Dietary energy concentration required for steers of different breeds fattened to a similar IMF content..

38

5.5. Conclusions 39

6. Characteristics of steers of six beef breeds fattened from eight months of age and

slaughtered at a target level of intramuscular fat. II. Meat quality 40

Abstract 40

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6.1. Introduction 41

6.2. Materials and methods 41

6.3. Results 43

6.4. Discussion 48

6.4.1. Realized contents of intramuscular fat in steers fed on a forage-based diet 48

6.4.2. Differences in meat quality of steers of different breed fattened to a target IMF level 49

6.4.3. Fattening series differences in meat quality of steers (tied housing vs loose housing) 52

6.5. Conclusions 53

7. Meat quality of Angus, Simmental, Charolais and Limousin steers compared at the same

intramuscular fat content 55

Abstract 55

7.1. Introduction 56

7.2. Materials and methods 56

7.2.1. Experimental design 56

7.2.2. Experimental procedures performed at slaughter 57

7.2.3. Determination of intramuscular fat content and marbling 57

7.2.4. Analysis of meat quality 58

7.2.5. Sensory evaluation 59

7.2.6. Statistical analysis 59

7.3. Results 60

7.4. Discussion 66

7.4.1. Marbling properties 67

7.4.2. Meat colour 67

7.4.3. Meat texture 68

7.4.4. Flavour, juiciness and water-holding capacity of meat 70

7.4.5. Overall sensory preference of meat 71

7.5. Conclusions 71

8. General discussion 73

9. References 76

10. Appendix 91

11. Remerciements 92

12. Curriculum Vitae 94

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1. Summary

Growth performance, carcass characteristics, and palatability attributes in steers of six different

beef breeds (Angus, Simmental, Charolais, Limousin, Blonde d'Aquitaine, Piedmontese; 22

animals/breed) were compared at a target level of intramuscular fat (IMF) content of 3.5%. The

total mix ration, provided ad libitum, consisted of maize silage, grass silage and concentrate

(52%, 26% and 22% of DM, respectively). Steers were purchased from suckler herds at the same

time and entered the trial at a similar age of approximately 8 months. Series 1 was performed in a

tie-stall barn while a loose-housing system with straw bedding was used in series 2. The animals

were assigned to slaughter when the IMF estimation in the M. longissimus dorsi (LD) according

to the estimation with a real-time ultrasound system in the live animals was approximately 3.5 %

or until 15 months of fattening had passed. This target level was fixed on basis of the results of a

preliminary study investigating the visual preference of marbling which can play an important

role for purchase decision.

The actually measured IMF contents in LD were 3.35 ±1.12, 3.47 ± 0.93, 3.49 ± 1.11 and 3.48 ±

1.08 % (± SD), for Angus, Simmental, Charolais and Limousin, respectively. In spite of a longer

fattening period, Blonde d'Aquitaine and Piedmontese did not reach this target with only 2.34 ±

0.64 and 2.40 ± 0.63 % IMF on average. The high variability between animals of the same breed

group in IMF content was due to the still restricted accuracy of the ultrasound method of IMF

determination in live animals.

Angus, Simmental, Charolais and Limousin reached the target of 3.5 % IMF on average at final

weights of 501 ± 43, 628 ± 60, 693 ± 117 and 668 ± 65 kg, respectively. Blonde d'Aquitaine and

Piedmontese did not reach this target, although the average fattening period lasted 15 months,

i.e., was about three times longer than for Angus, at final weights of 758 ± 93 and 647 ± 64 kg,

respectively. Except in Angus, the average slaughter weights were therefore higher than those

commonly found in Switzerland. The corresponding slaughter ages were 381 ± 25, 509 ± 72, 529

± 104, 610 ± 62, 690 ± 35 and 683 days ± 35 for the Angus, Simmental, Charolais, Limousin,

Blondes d'Aquitaine and Piedmontese, respectively.

Under the conditions of this experimental approach, daily gains were highest in Angus, followed

by Charolais, Simmental, Limousin and Blonde d'Aquitaine and lowest with Piedmontese. The

daily feed intake was significantly lower for Piedmontese than for Charolais, Simmental and

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Angus. As a result, Angus expressed the best feed conversion efficiency over the complete

fattening period while this efficiency was lowest in the Piedmontese. Among the four breeds,

which reached the target IMF content, Limousin steers showed the best carcass quality, namely

the highest dressing percentage and greatest proportion of premium cuts and the highest lean to

fat and lean to bone ratio in the sirloin, followed in descending order by Charolais, Simmental

and Angus. However all four groups were graded as too fat for the Swiss market.

Meat quality was measured in the LD and the M. biceps femoris, regio glutea (BF). Chemical

composition of both muscles did not reveal important differences between breed groups. Early

and late postmortem muscle pH was relatively similar among breeds in contrast to water-holding

capacity. Angus and Simmental presented the lowest drip losses and simultaneously the highest

thawing and cooking losses in both muscles (Simmental only LD). Globally the Piedmontese

presented the highest water-holding capacity. The LD of Angus and Charolais showed the palest

meat. In line with lightness, heme iron contents were clearly lowest in both muscles in the Angus

steers. No significant differences in collagen solubility and shear force were measured in the LD

between breed groups in contrast to BF and sensory analyses. In series 1 meat of the Angus was

significantly more tender than Blondes d'Aquitaine, Piedmontese and Simmental. But in series 2

Piedmontese were significantly tender than all other breed groups. Piedmontese and Blonde

d'Aquitaine presented a higher flavor intensity than Simmental and with Limousin and Charolais

a more juicy meat compared to Angus and Simmental. It seems that the higher growth rate and/or

movement in series 2 played an important role on meat quality traits, namely clearly better water-

holding capacity and higher tenderness in series 2 (group housing on straw bed) than in series 1

(tied system), especially in Piedmontese.

In conclusion, the present results revealed:

1. The difficulty with pure beef breeds and a forage based ration to reach the desired extent of

marbling and at the same time favourable carcass conformation, carcass size (except Angus)

and fat cover which meet Continental European, particularly Swiss, market demands.

2. Although slaughtered at a similar intramuscular fat content in the LD and reared under the

same conditions, differences in meat quality traits were still found between breed groups.

Nevertheless all breed groups presented meat of good to very good sensory quality even if

meat from Piedmontese was globally preferred relative to meat from Simmental.

3. The IMF content is therefore not the basic cause of the differences in meat quality observed.

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2. Résumé

Les performances d'engraissement, la qualité de carcasse et de viande de bœufs issus de six races

à viande (Angus, Simmental, Charolais, Limousin, Blonde d'Aquitaine, Piémontais; 22

animaux/race) ont été comparées à un taux de graisse intramusculaire cible de 3.5 %.

L'alimentation a été servie ad libitum et consistait pour 52 % d'ensilage de maïs, 26 % d'ensilage

d'herbe et 22 % de concentrés dans la matière sèche. Provenant d'exploitations de vaches

allaitantes, les bœufs ont débuté l'essai à l'âge de 8 mois environ. La première série

d'engraissement s'est déroulée en stabulation entravée et la deuxième en stabulation libre. Les

bœufs ont été abattus lorsque l'estimation du taux de graisse intramusculaire dans le M.

longissimus dorsi (LD), selon une méthode aux ultrasons, avoisinait 3.5 % ou sinon après 15

mois d'engraissement. Utilisée comme critère d'abattage, cette teneur de graisse intramusculaire

résulte d'une étude de préférence évaluant la qualité visuelle du persillé, cette dernière pouvant

jouer un rôle déterminant lors de l'achat.

Les taux de graisse intramusculaire dans le LD effectivement obtenus ont été de 3.35 ±1.12, 3.47

± 0.93, 3.49 ±1.11, 3.48 ± 1.08 % (SD), respectivement pour les Angus, Simmental, Charolais et

Limousin. Malgré une durée d'engraissement prolongée, les Blonde d'Aquitaine et les Piémontais

ont atteint une teneur moyenne de seulement 2.34 ± 0.64 et 2.40 ± 0.63. La variabilité élevée des

taux de graisse intramusculaire entre les animaux de la même race est due à la précision encore

insuffisante de la méthode d'estimation de la graisse intramusculaire aux ultrasons chez l'animal

vivant.

Les Angus, Simmental, Charolais et Limousin ont atteint l'objectif fixé d'environ 3.5 % de

graisse intramusculaire avec des poids vifs finaux en moyenne respectivement de 501 ± 43, 628 ±

60, 693 ± 117 et 668 ± 65 kg. Avec environ 2.4 % en moyenne, les Blonde d'Aquitaine et les

Piémontais n'ont pas atteint le taux cible de graisse intramusculaire malgré une durée

d'engraissement de 15 mois, soit 3 fois plus longue comparée aux Angus et avec des poids vifs

finaux de 758 ± 93 et 647 ± 64 kg. Hormis pour les Angus, les poids à l'abattage obtenus sont

donc plus élevés que ceux rencontrés habituellement en Suisse. Les âges à l'abattage

correspondants ont été de 381 ± 25, 509 ± 72, 529 ± 104, 610 ± 62, 690 ± 35 et 683 ± 35 jours.

Dans les conditions de cet essai, les accroissements journaliers moyens étaient les plus élevés

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pour les Angus, suivis par les Charolais, Simmental, Limousin, Blonde d'Aquitaine et les

Piémontais. L'ingestion journalière des Piémontais était plus faible (P < 0.05) que celle des

Charolais, Simmental et Angus. En conséquence, calculée sur toute la durée de l'engraissement,

les Angus ont présenté la meilleure et les Piémontais la plus mauvaise efficacité alimentaire.

Parmi les quatre races ayant atteint le taux de graisse intramusculaire souhaité, les Limousin ont

présenté la meilleure qualité de carcasse, à la fois concernant le rendement d'abattage, la part de

morceaux nobles et les rapports viande/graisse ou viande/os dans l'aloyau, suivis dans l'ordre

décroissant par les Charolais, Simmental et Angus. Pour le marché suisse, l'objectif d'un taux de

graisse intramusculaire de 3.5 % en moyenne a correspondu à un état d'engraissement exagéré de

la carcasse.

La qualité de viande a été comparée dans le LD et le biceps femoris, regio glutea (aiguillette

rumpsteak, BF). La composition chimique des 2 muscles étudiés ne révèle pas de différence

importante parmi les races. Les mesures de pH à 1 h et 48 h post mortem sont relativement

similaires entre les races contrairement à la capacité de rétention d'eau. Les Angus et les

Simmental présentent les pertes d'exsudats (drip loss) les plus faibles et simultanément les pertes

de décongélation et de cuisson les plus élevées en moyenne (Simmental seulement dans le LD).

Globalement, les Piémontais ont la meilleure capacité de rétention d'eau. Les LD des Angus et

des Charolais sont légèrement pâles avec une teneur en pigments plus basse dans les deux

muscles pour les Angus que pour les autres races. Dans le LD, aucune différence significative

entre les races n'est mesurée concernant la solubilité du collagène et la force de cisaillement,

contrairement au BF et à l'analyse sensorielle. Dans la première série, la viande des Angus est

significativement plus tendre que celle des Blonde d'Aquitaine, Piémontais et Simmental. En

revanche, dans la deuxième série, les Piémontais ont une viande significativement plus tendre que

toutes les autres races. Les Piémontais et les Blonde d'Aquitaine présentent une flaveur plus

intense que les Simmental et avec les Limousin et les Charolais ont une viande significativement

plus juteuse que celle des Simmental et Angus. Il semble que la vitesse de croissance nettement

supérieure et/ou le mouvement dans la deuxième série, en stabulation libre, ait joué un rôle

important sur les caractéristiques de qualité de viande, avec notamment une amélioration de la

capacité de rétention d'eau et de la tendreté, très marquée pour la tendreté chez les Piémontais

comparativement à la première série en stabulation entravée.

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En conclusion, les résultats démontrent:

1. Il est difficile avec des races à viande pures et une ration constituée principalement par des

fourrages de concilier un taux de graisse intramusculaire considéré comme idéal de 3.5 %

dans le faux-filet et en même temps une bonne charnure, un état d'engraissement et/ou un

poids de carcasse acceptables pour le marché suisse actuel.

2. Des différences de qualité de viande dans les deux muscles étudiés entre les groupes raciaux

sont constatées dans cet essai chez des bœufs ayant un taux de graisse intramusculaire

semblable dans le LD et élevés dans les mêmes conditions. Néanmoins, tous les groupes

raciaux ont présenté une qualité de viande jugée globalement bonne à très bonne, même pour

les Simmental, en retrait par rapport aux autres races et particulièrement aux Piémontais.

3. Les différences de qualité de viande observées ne sont donc pas dues directement à la graisse

intramusculaire.

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3. General introduction

Over the last decades the media have pointed out a lot of problems concerning the production of

meat in general, as for example bovine spongiform encephalopathy, foot and mouth disease,

environment pollution and lack of animals' care (Bonny, 2000). Economic pressures due to

steadily declining beef consumption have challenged the livestock and beef sectors to produce

meat that will regard the consumers' expectations. To be consistent with consumers' preferences,

farmers have attempted to optimize breed, fattening regime and housing. This encouraged the

development of a multitude of beef label programs (Dufey and Chambaz, 1999a). Generally, one

goal of these label programs is to achieve a clear differentiation from the conventional production

as for instance by prescribing a certain type of diet, avoidance of artificially produced feed

ingredients, access to fresh air, fattening of steers instead of bulls and/or the use of beef breeds or

crosses with beef breeds. The beef production in Switzerland is largely a by-product of dairy

husbandry but it has been noted in the last few years a clear shift towards using beef breeds.

There is therefore a need from the producers and the labels sides to be able to rely on

scientifically obtained data about the growth performance, carcass quality and meat quality of

different beef breeds fattened under typical Swiss production conditions. Carcasses and meat of

improved quality are anticipated from these labels particularly in sensory respect.

One component of beef meat quality is marbling, which describes the visible proportion and

distribution of intramuscular fat (IMF). The role of IMF or marbling on palatability traits is a

contentious issue. An abundance of research stretching over the last 30 years indicates that

tenderness and juiciness rating improve slightly as marbling/IMF increases, however

marbling/IMF explains only approximately 5 to 10 % of the variation in tenderness of the

longissimus dorsi (Blumer, 1963; Dikeman, 1996). Wheeler et al. (1994) and Rymill et al. (1997)

concluded that degree of doneness was considerably more important in producing tender and

juicy steaks than was IMF content. Actually IMF seems to be more an attribute which may

accompany high tenderness under certain circumstances rather than causing a direct effect on

tenderness, talking about amounts in the common range for Western beef production (Scheeder,

1998). Nevertheless IMF and palatability of beef meat remains tightly linked together in the mind

ofmost butchers and gastronomy circles. Therefore marbling is often subjectively equated by the

consumers with quality and eye appeal of IMF, i.e. its amount and distribution, can play an

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important role on purchase decision. This is specially the case in branded beef programs. A

preliminary study investigating the visual preference of marbling was conducted in Switzerland

using photographs of loin eye cuts. Excepted the class without marbling (27 % of the responses),

the most frequently selected marbling classes were the ones with 3 % and 4 % IMF with 47 % of

the responses. Going from the principle that the potential consumer of label meat wants a product

which is different from the conventional one, he should not focus on the absence of fat. On this

basis an IMF content of between 3 and 4 % was adopted in this study as slaughter end point.

Selecting a similar IMF content in the longissimus dorsi as slaughter criterion has raised the need

of an appropriate determination tool. Ultrasound technology has already widely been used and

evaluated for estimation of body composition in live beef cattle (Tschümperlin, 1996). In the

United States a new ultrasound method has been developed for about 10 years to permit also the

estimation of IMF in live animal. American farmers were interested in the possibility to estimate

early in the feedlot phase the IMF content of the animals as carcass quality in beef is determined

almost entirely by marbling score. This method was used in this study to determine the slaughter

end point of each animal.

Meat production has often been reproached to be not targeted on consumers' needs but on

carcass, an intermediate product. The concept of this project was therefore to center firstly the

interest on the recognized consumers' needs and to evaluate only secondly the resulting effects

upstreaming from the consumer level, i.e. on the butcher and farmer levels. This approach tries to

live up to consumers' expectations. It must be kept in mind that the volume of meat production is

finally determined in response of the degree of satisfaction aroused by the product.

The objectives of this study were to compare steers of six different beef breeds for a similar

intramuscular fat content of 3.5 % and to determine:

1. which differences can be observed in sensory traits?

2. which slaughter weight will be reached, for a given energy density ofthe diet?

3. which relationship exists between this IMF content and fatness of the carcass?

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4. Sources of variation influencing the use of real-time ultrasound to predict

intramuscular fat in live beef cattle

Based on:

A. Chambaz, P.-A. Dufey, M. Kreuzer, and J. Gresham, 2001

(submitted to Canadian Journal ofAnimal Science)

Abstract

A total of 123 steers of six European breeds (Angus, Simmental, Charolais, Limousin, Blonde

d'Aquitaine, Piedmontese) were used i) to evaluate the precision of the ultrasound predicted

intramuscular fat (USIMF) and its sources of variation using the current Pie QUIP technology ii)

to develop improved models for predicting USIMF. Steers were slaughtered when they reached

the target value of 3.5 % USIMF. Hide samples were obtained 3 d before slaughter by shot-

biopsy. After slaughter, a sample of the longissimus muscle was used for determination of actual

chemical intramuscular fat (EEIMF), collagen content and solubility. Among the variables

available during a chute-side scanning session, hide thickness and ultrasound subcutaneous fat

thickness at the 12th and 13th rib were shown to be significantly correlated with EEIMF. These

two variables were selected as possible independent variables to evaluate the construction of new

models. The model with the best fit included USIMF, hide thickness and live weight and had a

standard error of prediction of 0.96 % which is similar to other published technologies. Breed

group and collagen related traits did not influence USIMF estimation. Finally, the revised Pie

QUIP technology should be considered as one technology of choice to predict EEIMF content in

live animals.

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4.1. Introduction

Marbling, i.e. the visible proportion and distribution of intramuscular fat determined in the

longissimus dorsi muscle, is the most important factor affecting quality grade in the United States

(Boggs et al., 1998) and in Canada (Newman et al., 1994) which even opens markets for

unconventional solutions such as the use of Wagyu crossbred cattle (Mir et al., 1999). In Europe,

the carcass value is solely determined by its conformation and fatness, and no such system exists

for marbling as in the USDA quality grade system (USDA, 1989). Although the influence of

intramuscular fat percentage on the palatability of beef is a contentious issue (Wheeler et al.,

1994; Dikeman, 1996; Rymill et al., 1997), also in Europe marbling is often cited and demanded

as a primary quality attribute of beefby gourmets or in label productions. Marbling influences the

visual quality of beef which is a major component affecting positively or negatively purchase

decision of the consumers. Therefore it would be desirable to be able to accurately assess the

amount of intramuscular fat in the live beef animal prior to harvest to meet the demands of

different markets (Hassen et al. 1999; Brethour 2000) for instance a range of 3 to 4%

intramuscular fat which was the preferred range in a Swiss assessment (Chambaz et al., 2001a)

corresponding to the USDA grade 'Slight Degree of Marbling'. Real-time ultrasound technology

is a non-invasive method of determining on the live animal the percentage of intramuscular fat in

the longissimus dorsi (loin eye) muscle (Kriese, 1996; Herring et al., 1998). Considerable

improvement has been made in the last few years in different ultrasound technologies to estimate

intramuscular fat in live beef animals (Kriese, 1996; Herring et al., 1998; Brethour, 2000; Hassen

et al., 2001; Gresham, 2001). One technology that has not been documented in the literature as

extensively as others is known as the Pie QUIP technology (Gresham, 1997). This technology has

the advantage to provide instant chute-side results but was judged less accurate than four other

real-time ultrasound systems (Herring et al., 1998). But since then the Pie QUIP technology has

been modified.

The objectives of this study were: 1) to determine by live animal real-time ultrasound evaluation

the precision and accuracy of utilizing the current Pie QUIP technology in determining time of

harvest animals when between 3.0 and 4.0% intramuscular fat is reached, 2) to determine and

evaluate live animal and muscle tissue characteristics that might influence the precision and

accuracy of using ultrasound to estimate intramuscular fat in live beef animals, 3) to develop new

prediction models for improving the accuracy and precision of the current Pie QUIP technology.

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For this purpose steers of very different breed and highly varying in live weight were used in

order to ensure applicability for a broad range of animals.

4.2. Materials and methods

4.2.1. Animals

Two replicated finishing trials included each 12 steers out of six breeds yielding a total of 144

carcasses. Steers were either purebred Simmental, Charolais, Limousin, Blonde d'Aquitaine,

Piedmontese or Angus (% Angus obtained from grading up of native Swiss dairy breeds). All

steers were purchased from suckler herds at the same time and entered the trial at a similar age of

8.0 ± 0.8 mo (mean ± SD). The animals within each trial were finished under the same conditions

and had ad libitum access to a diet consisting, on a dry matter basis (g kg-1), of maize silage

(520), grass silage (260) and concentrate (220). The complete diet contained 135 g kg-1 crude

protein and 11.2 MJ kg-1 metabolizable energy. Trial 1 was performed in a tie-stall barn while a

loose-housing system with straw bedding was applied in Trial 2. The steers were cared for under

guidelines comparable to those laid down by the Canadian Council on Animal Care. Further

details on the animals and the experimental procedures are described elsewhere (Chambaz et al.,

2001a).

In order to develop and test a reliable prediction equation, two independent sets of experimental

animals were required. One group was needed to develop the equation, and an independent set of

animals was needed to evaluate the accuracy and precision of the equation (Gresham et al.,

1994). In this study, the data from every fourth animal harvested sequentially was designated as

part of the validation set. Animals with incomplete data for variables to be utilized in the analysis

were deleted from the total data set resulting in n = 123. As a result, there were 96 animals used

for developing the prediction equations (test data set) and 27 animals were used to validate the

prediction equations (validation data set).

4.2.2. Live animal measurements

Intramuscular fat content was estimated with a real-time ultrasound system in the live animals.

Images were captured with a Pie Medical scanner 200 (Maastricht, NL) equipped with a 3.5

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11

MHz, 18-cm transducer (Model ASP-18). Ultrasound estimation of intramuscular fat content

(USIMF) was accomplished with the machine internal analysis program and a software

prediction program available from Classic Ultrasound Equipment, Tequesta, FL, USA (Gresham,

1997). This technology, called Pie QUIP (Quality Ultrasound Indexing Program) technology, is

based primarily on a characterization of regions of analysis in the dorsal and ventral areas of the

tri th

longissimus dorsi muscle at the 12 and 13 rib (Gresham unpublished data). Then a

mathematical analysis is conducted by a proprietary software system that is a part of the

ultrasound scanner to analyze relationships of the regions of analysis by applying a mathematical

model influenced by attenuation and scattering of the ultrasound beam (Haumschild and Carlson,

1983; Brethour, 1990; Park et al., 1994). This program is recommended for beef animals up to 24

months of age (Gresham, 1996). Hair was clipped and linseed oil was applied to the hide surface

before scanning to ensure acoustical contact between surface of transducer probe and hide surface

(Gresham, 1999). The probe was centered directly above the 12th and 13th rib and placed

longitudinal to the mid-line of the animal at a point to approximate a mid-point between the

medial and lateral ends of the longissimus dorsi muscle. This site would then create an ultrasound

image in the area of the 12th/13th rib and 1st lumbar region (Gresham, 1997). Both subcutaneous

fat thickness (USSCF12/13) and QUIP index for USIMF calculations were measured at the same

time without need of a contour-fitting stand-off. Each steer was scanned in 14 d intervals until it

reached the targeted USIMF of about 3.5% as determined by real-time ultrasound evaluation or

after a maximum of 15 months of fattening if 3.5% was not reached. At least three scans were

obtained on each individual animal per scanning session for analysis to increase the precision of

the estimation (Hassen et al., 1999).

Linear live animal measurements included live weight, average daily gain, days on feed and age

at slaughter. As a variable which may influence the result of the ultrasonic measurements, hide

thickness was determined. Cores (8.3 mm diameter) of hide samples were obtained by shot-

biopsy (Biotech, Nitra, Slovakia) 3 d before slaughter. Biopsy was performed at the same site as

the ultrasound measurements on the shaved hide of the live animal. The core sample was

separated from the muscle and subcutaneous fat and the thickness of the hide was determined by

use of an electronic caliper square.

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4.2.3. Carcass measurements

Steers were transported for approximately 1 h to a commercial harvesting facility. Carcasses were

weighed at about 1 h post mortem, then moved to a chilling cooler and allowed to chill for 48 h at

2°C. Carcass subcutaneous fat measurements were obtained two days post mortem between the

9th and the 10th (SCF9/10) and between the 12th and the 13th rib (SCF12/13) to the nearest

millimeter with an electronic caliper square. Carcass grading in terms of fatness was performed

on a five-point scale (1 = low; 5 = high fatness) by experienced official staff according to the

Swiss beef classification grid (Proviande, 2001). A section of the longissimus dorsi muscle was

excised from the left side of each carcass and stored at 2°C until chemical analysis. A slice of

approximately 300 g of longissimus dorsi muscle (region of 12th and 13th rib) was trimmed free

of extraneous fat and muscle epimysial tissue, lyophilized (48 h, under vacuum at -20°C;

Lyophilisator Christ, model Delta 1-24K, Adolf Kühner AG, Birsfelden, Switzerland) and

homogenized. Intramuscular fat was defined as ether extractable fat (EEIMF) and determined by

the Soxhlet procedure using petroleum ether (SLB, 1969). Total collagen content and collagen

hydrothermal solubility were determined by the method described by Arneth and Hamm (1971)

and adapted for the Technicon Autoanalyser II analysis chain (Technicon, Plainfield, NJ, USA).

4.2.4. Statistical analysis

Means, SD, ranges, root mean square errors (RMSE) and correlations were determined and

stepwise regression analyses were performed (SAS, 1985). Statistics generated in the stepwise

regression analysis included R2, Cp, and SE (Mallows, 1973; MacNeil, 1983). The standard error

of prediction (SEP) was calculated according to Herring et al. (1998). The SEP measures the

ability of the technician to rank or predict differences between animals correctly, correcting for a

tendency to consistently over- or under-predict the true measurement. After completion of

regression analyses and generation of prediction equations from the first data set (n = 96), data

from the second set (n = 27) were used to validate the prediction equations.

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13

4.3. Results and discussion

4.3.1. Test and validation ofthe prediction equation

Table 1 presents the means, standard deviations, minimum and maximum values of all variables

that were used in modeling prediction equations to estimate intramuscular fat in live beef cattle.

There was no significant difference (P > 0.05) in means for the test and validation sets for any

variable listed.

Table 1

Descriptive statistics oflive animal linear and ultrasound as well as carcass traits by data set

Test Data Set (n = 96) Validation Data Set (n == 27)

Trait1 Mean SD Min. Max Mean SD Mm. Max.

EEIMF (%) 3.10 1.35 1.18 9.12 3.46 1.19 1.23 6.37

USIMF (%) 3.25 0.75 1.00 5.67 3.44 0.66 2.44 4.93

SCF9/10 (mm) 19.5 5.4 7.7 36.1 19.4 6.7 8.9 34.0

SCF12/13 (mm) 10.5 4.7 1.7 24.6 11.3 5.1 2.3 19.6

USSCF12/13 (mm) 7.2 2.4 2.3 13.5 7.4 2.2 3.8 11.4

Hide thickness (mm) 7.6 0.9 5.1 105 7.6 0.9 5.7 9.4

Days on feed (d) 331 116 117 497 315 124 117 497

Weight gain (kg/d) 1.09 0.20 0.61 1.58 1.10 0.22 0.63 1.55

Age at slaughter (mo) 18.9 3.9 11.8 24.5 18.2 4.2 11.7 24.0

Live weight (kg) 661 110 458 918 636 88 475 821

Hot carcass weight (kg) 392.5 79.0 251.6 566.2 376.7 69.9 263.8 529.4

Graded fatness (units 1-5) 3.9 0.8 2.0 5.0 40 0.8 3.0 5.0

EEIMF = ether extractable intramuscular fat; USIMF = ultrasound predicted intramuscular fat

content using current Pie QUIP technology; SCF9/10 = carcass subcutaneous fat thickness between 9th

and 10th rib; SCF12/13 = carcass subcutaneous fat thickness between 12th and 13th rib; USSCF12/13 =

ultrasound predicted subcutaneous fat thickness between 12th and 13th rib

When breed group was used as an independent variable in the prediction equations, no significant

breed group effect was observed (data not shown). Therefore, since it was determined that a

single equation could be used across breeds and individual breed equations were not merited or

required, data were pooled for all breeds. While some variables were significantly different

between breeds (hide thickness), this variation can be accounted for by using those variables as

independent vanables in the equations.

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14

Table 2 presents the simple linear correlations for the traits described in Table 1 for both the test

and validation data sets. Values above the diagonal are correlations for the test data set, while

values below the diagonal reflect correlations for the validation data set. In the test data set,

significant correlations are noted between EEIMF and USIMF (P < 0.001), SCF12/13 (P <

0.001), USSCF12/13 (P < 0.001), hide thickness (P < 0.05), daily weight gain (P < 0.01) and

graded carcass fatness (P < 0.001).

USSCF12/13 was identified on the same scan used for determining intramuscular fat and is thus

immediately available during the chute-side scanning session. Therefore this variable was then

selected as possible independent variable to evaluate the construction of a new equation to

enhance the accuracy and precision of the current QUIP index for estimating USIMF in live beef

cattle. A step-wise regression analysis was conducted using USSCF12/13 and the quadratic of

USSCF12/13 with the stepwise regression presented in Table 3. Therefore, the new prediction

equation including USSCF12/13 as a variable is:

USIMFnewl (%) = -4.211 + [0.899 x USIMF(%)] + [0.516 x USSCF12/13 (mm)

- (0.025 x USSCF12/132(mm2)] + [0.003 x live weight (kg)]

The prediction equation was tested against an independent set of animals (n = 27) described in

Tables 1 and 2 in order to validate it. Table 4 presents the statistics for the validation of the

equation (USIMFnewl). Statistically, an unbiased and valid equation should have an intercept of

zero with a slope of one. This equation meets this test since the slope and intercept values include

one and zero respectively within their standard error values.

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Table2

Simple

linearcorrelationsforliveanimallinearandultrasoundaswellascarcass

traitsbydata

set1

Trait2

12

34

56

78

910

11

12

EEIMF(%)

(1)

-

0.41

0.18

0.40

0.33

0.24

-0.18

0.26

-0.14

0.09

-0.06

0.52

USIMF(%)

(2)

0.66

-

0.07

0.33

0.38

-0.24

-0.51

0.32

-0.47

-0.22

-0.44

0.45

SCF9/10(mm)(3)

0.39

0.31

-

0.53

0.48

0.15

-0.19

0.26

-0.19

0.28

0.21

0.50

SCF12/13(mm)

(4)

0.65

0.65

0.40

-

0.61

0.32

-0.63

0.61

-0.60

-0.09

-0.24

0.57

USSCF12/13(mm)

(5)

0.39

0.29

0.43

0.60

-

0.31

-0.23

0.17

-0.25

-0.30

-0.22

0.58

Hidethickness(mm)

(6)

-0.10

-0.20

0.25

-0.08

0.41

-

-0.06

0.28

-0.08

0.17

0.03

0.27

Daysonfeed(d

)(7

)-0.17

-0.48

0.08

-0.44

-0.44

0.17

-

-0.79

0.98

0.72

0.85

-0.41

Weightga

in(k

g/d)

(8)

0.34

0.38

0.09

0.43

0.42

0.06

-0.69

-

-0.68

-0.25

-0.46

0.57

Age(mo)

(9)

-0.11

-0.39

0.10

-0.40

-0.49

0.09

0.98

-0.78

-

0.75

0.87

-0.39

Liveweight(k

g)(1

0)0.24

-0.36

-0.02

-0.29

-0.12

0.18

0.67

-0.01

0.69

-

0.95

0.04

Hotcarcasswt(k

g)(1

1)0.05

-0.34

-0.15

-0.46

-0.44

0.02

0.81

-0.22

0.82

0.94

-

-0.15

Graded

fatness(units)(1

2)0.55

0.27

0.35

0.64

0.47

0.09

-0.40

0.49

-0.36

-0.03

-0.17

-

1Valuesabovethedi

agon

alaresi

mple

linearcorrelationsofvariablesusedinthetestdata

set

(n=

96;P<

0.05

ifr>

0.20

;P<0.01

ifr>

0.25

;P<

0.001

ifr>

0.32),

variablesbelowthediagonal

aresimple

linearcorrelationsforthesame

variablesasusedinthevalidationdataset

(n=

27;P<

0.05

ifr>

0.38;P<

0.01

ifr>

0.56;P<0.001

ifr>

0.64

).

2EEIMF=

etherextractableintramuscular

fat;USIMF=ultrasoundpredictedintramuscularfatcontentusingcurrentPieQUIPtechnology;

SCF9/10=

carcasssubcutaneous

fatthicknessbetween

9thand

10th

rib;

SCF12/13=

carcasssubcutaneous

fatthicknessbetween

12th

and

13th

rib;

USSCF12/13=ultrasoundpredictedsubcutaneous

fatthicknessbetween

12thand13

*ri

b.

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Table

3

New

regressionequationforultrasoundpr

edic

tedpercentageofintramuscular

fat(USIMFnewl)from

live

animalvariablescollected

atharvest

(n=

96)'

Trait

Step

Rz

C„

Partialregressioncoefficients

RMSE(%)

Intercept

USIMF(%)

Liveweight

(kg)

USSCF12/13(mm)

USSCF12/13'(mmz)

USIMFnewl

10.19

1473

1.52

0.202

0.882

20.25

9.30

1.17

-2.471

1.048

0.003

30.30

5.25

1.36

-3.019

0.919

0.003

0.129

40.32

500

1.34

-4.211

0.899

0.003

0.516

-0.025

Cp=Mallow's

stat

isti

c,RMSE=rootmeansquare

error;USIMF=ultrasoundpr

edic

tedintramuscularfatcontentusingcurrentPieQUIP

tech

nolo

gy,

USSCF12/13=ultrasoundpredictedsubcutaneous

fatthicknessbetween

12th

and13*

rib;

USFATnewl=USIMF

equationin

clud

ingUSSCF12/13.

Table4

Regressionofetherextractableintramuscularfa

t(EEIMF)onnewequationsofultrasoundpr

edic

tedliveanimalintramuscular

fat

(n=

27)

Predictioneq

uati

on1

Intercept

+SE

Bi

±SE

R1RMSE

(%)

2

rPr/

SEP4(%)

USIMFnewl

USIMFnew2

-0.464

-0.176

0.625

0.875

1.259

1.099

0.195

0.259

0.63

0.42

0.75

0.93

0.79

0.65

0.78

0.62

1.03

0.96

1USIMFnewl

includesUSSCF12/13;USIMFnew2

includeshidethickness.

2

rp=Pearsoncorrelationcoefficient.

3

rr=Spearmanrankcorrelationcoefficient.

4SEP=standarderrorofpr

edic

iton

.

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17

In order to detect and evaluate new possible sources of variation in the precision of the

estimation of the USEVIF, a subsequent prospective step was conducted with the variables

available before slaughter in this study. All the variables shown to be significantly correlated

with EEIMF, i.e. USIMF, hide thickness, live weight, USSCF12/13 and daily weight gain,

were used in a stepwise regression analysis as independent variables to generate a new

ultrasound predicted intramuscular fat content value (Table 5). When both hide thickness and

USSCF12/13 are included in the equation, hide thickness becomes a significant variable and

both USSCF12/13 and daily weight gain are eliminated.

Table 5

Stepwise regression analysis for development of a new ultrasound prediction equation to

intramuscular fat (USIMFnew2) from live animal variables collected at harvest (n = 96)

Trait Partial Rz Model Rz cp P>F

USIMF (%) 0.1932 0.1932 19.71 0.0001

Hide thickness (mm) 0.1116 0.3048 6.67 0.0003

Live weight (kg) 0.0429 0.3477 2.88 0.0175

USSCF12/13 (mm) 0.0047 0.3524 4.26 0.4285

Weight gain (kg d"1) 0.0019 0.3543 6.00 0.6734

USIMFnew2 includes hide thickness; Cp = Mallow's statistic; USIMF = ultrasound predictedintramuscular fat content using current Pie QUIP technology; USSCF12/13 = ultrasound

predicted subcutaneous fat thickness between 12th and 13th rib.

Based on Cp statistic, the best model for predicting EEIMF is an equation to include USIMF,

hide thickness and live weight. Therefore, the new prediction equation for ultrasound

prediction of intramuscular fat using hide thickness (Table 6) is:

USIMFnew2 (%) = -5.534 + [0.994 x USIMF (%)] + [0.460 x hide thickness (mm)] + [0.003

x live weight (kg)]

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Table6

New

regression

equation

forultrasoundpredic

tedpercentageofintramuscular

fat(USIMFnew2)from

liveanimalvariablescollected

atharvest

(n=

96)1

Partialregression

coefficients

Trait

Step

R2

Cp

RMSE

(%)

Inte

rcep

tUSIMF(%)

Hidethickness(mm)

Liveweight(kg)

USIMFnew2

ÎÔ7Ï7

21.11

L24

"

0.715

0.734

20.28

8.27

1.17

-3.503

0.853

0.502

30.32

4.00

1.14

-5.534

0.994

0.460

0.003

USIMFnew2

includeshideth

ickn

ess;

Cp=Mallow's

statistic;RMSE=rootmeansquare

error;USIMF=ultrasoundpr

edic

tedintramuscular

fatcontent

usingcurrentPieQUIPtechnology.

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19

USIMF is the value produced by the current Pie QUIP technology for estimating EEEVIF in

live beef cattle (Gresham 1997). However, it appears that the current equation predicting

EEIMF can be increased in accuracy and precision by including hide thickness. The test of the

accuracy of the new equation developed (USIMFnew2) to predict EEIMF is presented in

Table 4 using steers from the validation data set. USIMFnew2 was validated because the

intercept contains 0 and the slope contains 1. It presented a lower SEP compared to

USIMFnewl. In addition, the authors have observed that it is more difficult to obtain quality

images with animals (breeds) that possess thick hides. Since the Pie QUIP technology is

based on attenuation and scattering of sound waves, hide thickness might be a contributing

factor to why some researchers have criticized the QUIP technology for not being as accurate

as other documented technologies (Herring et al., 1998; Wilson et al., 1998). For practical

reasons the measurement of hide thickness using biopsy is problematic during a chute-side

scanning session in the field. The hide thickness in the live animal can also be measured on

the same scan as for USIMF but no data about the precision of its estimation is available in

the literature. Nevertheless the very easy determination of upper and lower hide limits on the

scan would lead one to expect a similar high precision compared to the measurement of

USSCF12/13 (Houghton and Turlington, 1992).

4.3.2. Comparisons ofthe new equations developed with results ofthe literature

In order to validate technologies and certify scanning technicians the SEP has been most

commonly used to test models for predicting EEIMF (Kriese, 1996; Herring et al., 1998;

Hassen et al., 2001). The most recent study to evaluate ultrasound technology was reported by

Hassen et al. (2001) comparing the Aloka 500V and Classic 200 scanners using the USOFT

prediction technology (Izquierdo et al., 1996), which is the standard in the United States.

Table 7 compares the results of the data set II reported by Hassen et al. (2001) for the

accuracy of each machine on a test set of animals versus the results of the validation test of

the new Pie QUIP equations generated and validated in this study. In reviewing the data in

Table 7 it is apparent that the new adjusted Pie QUTP equations are quite comparable to the

models and machines reported by Hassen et al. (2001). In addition, the accuracy and precision

for the Pie QUIP technology as reported in this study is superior to the values reported by

Herring et al. (1998) and representative of the field results for the current Pie QUIP

technology summarized by Gresham (2001). The major differences observed in Table 7 is that

the USOFT technology reported by Hassen et al. (2001) has a slightly lower SEP and higher

correlation coefficients compared to the equation generated using hide thickness

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20

(USIMFnew2). However, the equation generated by using USSCF12/13 (USIMFnewl) is

quite comparable in both correlation coefficients. The difference in the correlation

coefficients might be explained in part by the lower EEIMF content in the loin eye of the

animals reported in this trial and especially a lower standard deviation. Therefore, this trial

presents two new prediction equations that can be used to improve the precision and accuracy

of the Pie QUIP technology while still affording the opportunity to provide instant chute-side

results. Nevertheless the prediction equation using the measurement of hide thickness from

biopsy (USFATnew2) can only be considered for research purposes. Operators using the

USOFT technology described by Amin et al. (1997) and participating in the Centralized

Ultrasound Processing program described by Hays et al. (1999) must capture their images on

a disk and then mail the images to a central laboratory for interpretation. This negates the

opportunity for instant, real-time chute-side evaluation of either breeding or feedlot animals.

Table 7

Comparison of prediction statistics for predicting intramuscular fat in live beef animals for Pie QUIP

technologies and USOFT

Technology

Machine

Equation2

Pie QUIP

Scanner 200

USIMFnewl

Scanner 200

USMFnew2

USOFT1

Aloka 500 Scanner 200

Trait

Observations (n) 27 27 /1 /1

EEIMF (+ SD) (%) 3.46(1.19) 3.46(1.19) 3.66(1.73) 3.66(1.73)

Difference to EEIMF (%) +0.34 +0.15 +0.42 +0.67

SEP (%) 1.03 0.96 0.84 0.81

rP 0.79 0.65 0.88 0.89

rr 0.78 0.62 0.88 0.91

1

Technology developed by the IOWA State University (see Amin et al. 1997); results of the data set II

reported by Hassen et al. (2001).

2USDVlFnewl includes USSCF12/13; USIMFnew2 includes hide thickness.

3EEIMF = ether extractable intramuscular fat; SEP = standard error ofprediction; rp

= productmoment correlation coefficient; rr

= rank correlation coefficient.

Table 8 compares results for the Pie QUIP technology utilized at the 1997 Beef Improvement

Federation (BIF) proficiency testing program for scanning technicians (Wilson et al., 1998)

with the same technology used by the authors in the current study (n = 123), plus a

comparison of the new formulas (USIMFnewl and USIMFnew2) when tested on the

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21

validation set of data (n = 27). It is apparent from reviewing the data in Table 8 that both the

USIMFnewl and USIMFnew2 equations have significantly reduced the SEP for predicting

intramuscular fat (measured by EEIMF) in live beef cattle with the Pie QUIP technology. In

addition, the SEP values of 1.03 and 0.96% reported here are in agreement with similar values

(0.98% and 1.03%, data set III) reported by Hassen et al. (2001).

Table 8

Comparison of trials using the Pie QUIP technology

Model USIMF1 USIMFnewl2 USIMFnew2z USIMF"

Traif

Observations (n) 123 27 27 43

EEIMF (± SD) (%) 3.19(1.33) 3.46(1.19) 3.46(1.19) 3.81 (1.54)

Difference to EEIMF (%) +0.11 +0.34 +0.15 not available

SEP (%) 1.18 1.03 0.96 1.32

rP 0.45 0.79 0.65 not available

rr 0.43 0.78 0.62 not available

Results for ultrasound predicted intramuscular fat in this trial on all animals using current Pie QUIP

technology (Gresham 1997).

2

Equations from this study: USIMFnewl includes USSCF12/13; USIMFnew2 includes hide thickness.

3Results obtained at the 1997 Beef Improvement Federation (BIF) proficiency testing program for

scanning technicians (Wilson et al. 1998).

4EEIMF = ether extractable intramuscular fat; SEP = standard error of prediction; rp

= productmoment correlation coefficient; rr = rank correlation coefficient.

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4.3.3. Possible muscle characteristics influencing intramuscularfat estimation by ultrasound

Some reports have questioned the influence of collagen on accuracy and precision of using

ultrasound technology to predict EEEVIF in beef cattle (Brethour, 1990; Park et al., 1994).

Therefore, it would be of interest to investigate whether collagen content and collagen

solubility might be related to the accuracy or precision of estimating EEIMF content of live

beef muscle by real-time ultrasound. These two collagen traits were arbitrarily added as new

independent variables to the previously generated prediction equations (USIMFnewl) and

(USIMFnew2). If either of these variables did in fact influence the prediction of EEIMF, this

should be reflected in the test of the new equations. The following equations were then

generated by step-wise multiple regression analysis as previously described:

USIMFnew3 (%) = -6.785 + [0.830 x USIMF (%)]+ [0.062 x USSCF12/13 (mm)] + (0.010 x

collagen content (mg 100 g"1)] - [0.041 x collagen solubility (%)]+

[0.004 x live weight (kg)]

and

USIMFnew4 (%) = -7.084 + [0.767 x USIMF (%)]+ [0.231 x hide thickness (mm)] + [0.009

x collagen content (mg 100 g"1)] - [0.037 x collagen solubility (%)]+

[0.004 x live weight (kg)]

Table 9 compares the statistics of each of the four prediction equations when collagen content

and collagen solubility are added to the equations previously generated from live animal

measurements only (USIMFnewl and USIMFnew2). When the collagen traits were added as

variables together with USSCF12/13 to the equation (USIMFnew3), both correlation

coefficients and the R2 decreased while the SEP increased from 1.03 to 1.31%. Similar results

were obtained when the collagen traits were added to the USIMFnew2 equation yielding

equation USIMFnew4. Since adding the values to existing equation would not increase

precision or accuracy of the established statistical measures, it would indicate that the

collagen traits were not influencing the ability to estimate EEIMF by real-time ultrasound in

this study.

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Table 9

Comparison of the new equations for ultrasound prediction of intramuscular fat in live beef

cattle with and without inclusion of collagen content and collagen solubility

Prediction equation USIMFnewl USIMFnew2 USIMFnew3 USIMFnew4

Variable(s) added to USSCF12/13 Hide USSCF12/13& Hide thickness <&

USIMF equation thickness collagen traits collagen traits

Trait

Observations (n) 27 27 27 27

EEIMF (±SD) (%) 3.46(1.19) 3.46(1.19) 3.46(1.19) 3.46(1.19)

USIMFnew (±SD) (%) 3.12(0.75) 3.31 (0.70)3.03 (1.11) 3.42 (1.02)

Difference to EEIMF (%) 0.34 0.15 0.33 0.04

RMSE(%) 0.75 0.93 0.92 0.98

R2 0.63 0.42 0.43 0.35

SEP (%) 1.03 0.96 1.31 1.01

rP 0.79 0.65 0.66 0.59

rr 0.78 0.62 0.65 0.56

EEIMF = ether extractable intramuscular fat; USIMFnew = new ultrasound predicted intramuscular

fat content; RMSE = root mean square error; SEP = standard error of prediction; rp= product moment

correlation coefficient; rr = rank correlation coefficient.

4.4. Implications

At the present time, there are several technologies utilized in predicting intramuscular fat in

live beef cattle. It would appear that the Pie QUIP technology should be available as one

technology of choice for operators desiring to use chute-side scanning in order to produce

instant results that can be immediately utilized in selection of potential breeding seedstock, or

determining harvest date for finished cattle by the producer. Evidence is presented that hide

thickness may influence the precision and accuracy of this determination, therefore further

studies with larger numbers of animals are necessary to determine the relationship between

ultrasound and biopsy measures of hide thickness and so develop a new prediction equation

for field use. The results indicate that neither total collagen content nor collagen solubility

should influence the ability to estimate intramuscular fat by this technique. Evidence is

documented that the revised Pie QUIP technology is similar in accuracy and precision to other

reported technologies.

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5. Characteristics of steers of six beef breeds fattened from eight months of

age and slaughtered at a target level of intramuscular fat.

I. Growth performance and carcass quality

Based on:

A. Chambaz, I. Morel, M. R. L. Scheeder, M. Kreuzer and P.-A. Dufey, 2001,

Arch. Anim. Breed. 44:395-411

Abstract

Growth performance and carcass quality of 132 steers originating from six beef breeds, Angus (AN),

Simmental (SI), Charolais (CH), Limousin (LI), Blonde d'Aquitaine (BL), and Piedmontese (PI),

fattened under the same conditions on the same diet, were compared at a target level of 3.5 %

intramuscular fat (IMF) in the M longissimus dorsi. This target level was set on basis of the results of

a preliminary study investigating, with 784 persons, the visual preference of marbling using

photographs. The total mix ration, provided at ad libitum access, consisted of maize silage, grass

silage and concentrate (52 %, 26 % and 22 % of DM, resp.). Series 1 was performed m a tie-stall barn

while a loose-housing system with straw bedding was used in series 2. The animals were assigned to

slaughter either when the target IMF content was reached according to the estimation with a real-time

ultrasound system applied in the live animals or when 15 months of fattening had passed. AN, SI, CH

and LI reached 3.5 % IMF on average at final weights of 501 ± 43, 628 ± 60, 693 ± 117 and 668 ± 65

kg, respectively. BL and PI did not reach this target, although the average fattening period was about

three times longer for BL and PI than for AN and the final weights were 758 ± 93 and 647 ± 64 kg,

respectively. Under the conditions of this experimental approach, daily gains were highest in AN,

followed by CH, SI, LI and BL and lowest in PI. The daily feed intake was significantly lower for PI

than for CH, SI and AN. The AN expressed the best feed conversion efficiency m terms of DM

expenditure per kg gain over the complete fattening period while this efficiency was lowest m the PI

group followed by BL. Among the four breeds, which reached the target IMF content, LI steers

showed the greatest proportion ofpremium cuts and the highest lean to fat and lean to bone ratio in the

sirloin, followed, in descending order, by CH, SI and AN. However all four groups were graded

around 4 m fatness score (high to very high). The present results revealed for all breeds the difficulty

to reach the desired extent of marbling and at the same time favourable carcass conformation, carcass

size (except AN) and fat cover which meet market demands.

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5.1. Introduction

Over the last decades, consumers increasingly demand and purchase beef produced under

conditions as natural as possible (Badertscher Fawaz et al., 1998). This includes suckler cow

systems, forage-based diets and the ban of artificial growth enhancers (Harrington, 1994) and

encouraged the development of a multitude of beef label programs (Dufey and Chambaz,

1999a). Generally, one aim of these label programs is to achieve a clear differentiation from

the conventional production as for instance by prescribing a certain type of diet, fattening

steers instead of bulls and/or the use of beef breeds or crossbreds, the latter particularly in

countries where beef production is largely a by-product of dairy husbandry (Geay and Micol,

1988) like Germany (Augustini et al., 1990) and Switzerland (Dufey and Chambaz, 1999a).

Carcasses and meat of improved quality are expected from these labels when compared with

conventionally produced beef, particularly in sensory respect (Buttery et al., 1997). One major

visual component affecting purchase decision of the consumers is marbling, which describes

the visible proportion and distribution of intramuscular fat (IMF) as determined in the M.

longissimus dorsi (M.l.d.) (Savell and Cross, 1988). Marbling is often monitored in breed

comparisons but was rarely used as the decisive slaughter criterion. Typically, animals in such

studies were slaughtered at the same chronological age (Riley et al., 1986; Dikeman and

Crouse, 1975; Oldigs et al., 1989), physiological age (Geay and Robelin, 1979), or weight

(Dubeski et al., 1997a). Few investigations applied the same fatness score (Schläpfer, 1986;

Kaufmann and Chavaz, 1989) as slaughter criterion or used the same percentage of carcass

fat, applying a posteriori selection (Geay and Malterre, 1973). To our knowledge, only in one

study (Smith et al., 1976) growth and feed efficiency of different beef crosses were compared

at a distinct IMF content (5 %), which was performed by statistical adjustment of means and

the application of a regression for days on feed.

The objective of the present experiment was to compare growth performance, carcass traits

and meat quality of steers at the same target IMF content of 3.5 % in M.l.d. The steers

originated from six beef breeds and were fattened under the same conditions on an identical

forage-based diet for all animals. This first communication focuses on results concerning

growth and slaughter performance. Meat quality issues are described in a subsequent

communication (Chambaz et al., 2001b).

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5.2. Materials and methods

26

5.2.1. Animals and experimental design

A total of 132 steers originating from six beef breeds, Angus (AN), Simmental (SI; original

without Red Holstein blood), Charolais (CH; line 'maternal quality'), Limousin (LI), Blonde

d'Aquitaine (BL), and Piedmontese (PI), were used in this study. AN and SI were obtained

from Swiss farms, CH, BL and LI were imported from France and PI from Italy. All animals

were purebred, except AN (75% AN blood on average) which had been derived from grading

up native Swiss dairy breeds mainly by the use of American AN bulls. Two subsequent

fattening series were carried out, each including 11 steers per breed. Series 1 was performed

in a tie-stall barn (space allowance per animal: 0.95 x 1.86 m), while a loose-housing system

with straw bedding (10 m2 per animal including an outside area) was used in series 2 due to

the intermediately performed transformation of the barns. All imported animals had to pass a

3-week period of quarantine in the country of origin as well as in Switzerland. During the

quarantine in Switzerland the animals were already adapted to the experimental diet. All

steers were purchased from suckler herds at the same time and entered the trial at a similar

average age of 238 + 22, 236 ± 25, 235 ± 29, 249 + 19, 230 ± 21 and 240 ± 13 d (means ± SD)

for AN, SI, CH, LI, BL and PI (PI, only series 2), respectively. In series 1, the PI had to pass a

prolonged quarantine in Italy and started the experiment at an average age of 281 + 16 d. All

steers were weighed at the start and at the end of fattening (directly before transport to

slaughter) as well as periodically every two weeks during fattening.

5.2.2. Diet

A total mix ration consisting of maize silage, grass silage (early-cut grass-clover mixture) and

concentrate was provided. Table 1 gives the ingredient and nutrient composition of this diet

and its components. The concentrate consisted per kg of 400 g barley, 240 g soybean meal,

200 g triticale, 100 g wheat and 60 g mineral-vitamin premix. Maize silage and grass silage

were offered in a ratio of 2:1 in terms of dry matter based on 2-weekly determinations owing

to the fact that growing cattle have high requirements for energy relative to protein. In

addition, samples from all diet components were taken every two weeks for nutrient

compositional analysis. There were only small differences between the two series in the

nutrient content of forages and concentrate, while certain variations were found within series

particularly when using new batches of grass silage.

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Table 1

Composition and nutrient contents of the experimental diet (means ± SD)'

Ingredient Maize silage Grass silage Concentrate Total Mix Ration2

Proportion (g/kg total dietary dry matter) 520 260 220 1000

Dry matter (DM, g/kg) 320 ± 28 385 ±84 879 +07 460 ± 28

Organic matter (g/kg DM) 963.2 ± 3.2 870 7 ±16.7 915.4 ±2 1 928.6 1 4.4

Crude protein (g/kg DM) 73.7 ±34 195 8 ±26.0 207 5 ±5.0 135.0 1 7.8

Crude fibre (g/kg DM) 206.0 ±15.0 211.7 ±24.9 38.0 ±2.6 170.6 ±11.5

Net energy3 (MJ NEV/kg DM) 6.70+ 0.13 6.23+ 0.51 8.12±0.10 6.89 ± 0.14

Metabohzable energy4 (MJ ME/kg DM) 11.0 ± 0.1 10.3 ± 0.6 12.6 10.1 11.2 1 0.2

Absorbable protein3 (g APD/kg DM) 70.5 ± 1.4 80.7 ±48 133 7 ±6.6 87.1 ± 1 9

Average of 61 determinations2Minerals and vitamins content per kg DM: 7 g Ca, 3 g P, 2 g Mg, 1 g Na, 0.18 mg Se, 54 mg Zn, 9 mg Cu, 54

mg Mn,

2'900 IU vitamin A, 56 mg a-tocopherol3

According to RAP (1999)

"According to DLG (1997)

Concentrate proportion was calculated in a way that the complete ration, being provided ad

libitum, generally covered the requirements for both net energy for growth (NEV) and

absorbable protein at the duodenum (APD) at the start of fattening. These requirements were

calculated on the basis of the Swiss recommendations developed for dairy crossbred steers

expecting average daily gains of 1.2 kg/d (RAP, 1999). The mineral-vitamin premix was

designed to supplement deficiencies and imbalances of the ration. Breed specific differences

in nutrient requirements were deliberately not accounted for in this study in order to exclude

direct diet effects. The composition of the ration was not changed during fattening. This

procedure may have resulted in a certain excess of protein as the fattening period proceeded

but ensured that deficiencies were avoided. The increasing demand for energy with growing

live weight was covered by the associated increase of the intake capacity of the steers at a

constant medium energy density of the complete ration. Feed intake of each steer was

recorded daily in both series. In series 2 this was accomplished by the use of electronically

controlled doors (Insentec, NL-Marknesse).

5.2.3. Endpoint offattening

The aim of this investigation was to slaughter all animals when they reached a similar IMF

content. The target IMF level was determined in a preliminary study investigating the visual

preference of marbling. In that study, 784 persons (46 % female, 54% male; 11.3 % < 21,

29.3 % between 21 and 40, 45.1 % between 41 and 60, 14.3 % > 60 years of age) had to select

the most preferred out of six photographs of M.l.d. cuts taken between the 9th and 10th rib,

which represented six classes of IMF content covering the range from 1 to 6 %. The

photographs showed only the central part of the slice to exclude any influence of the shape of

the M.l.d. and were presented without noticeable colour difference to avoid potential bias due

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to colour. The most frequently selected marbling classes were either the ones with 3 % and

4 % IMF with 30 % and 17 %, respectively, or the class without any marbling (27 % of the

responses). The 2, 5 and 6 % IMF classes were only chosen by 14, 6 and 5 % of the

participants. A range of 3 to 4 % IMF is also often quoted in other studies as to be the

minimum level necessary for the subjective perception as 'good beef quality' (Campion and

Crouse, 1975; Goutefongea and Valin, 1978; Savell and Cross, 1988). For these reasons a

content of 3.5 % IMF in M.l.d. was chosen as the target value. The IMF content was

estimated with a real-time ultrasound system in the live animals. Images were captured with a

Pie Medical scanner 200 SLC (NL-Maastricht) equipped with a 3.5 MHz, 18-cm transducer

(Model ASP-18) and a computer program, which provides the opportunity for chute-side

evaluation of IMF (Classic ultrasound equipment, Tequesta, USA-Florida). This program was

developed for beef animals up to 24 months of age (Gresham, 1996). The hair was clipped

and linseed oil was applied to the skin before scanning to ensure sufficient acoustical contact

between probe and skin surface. The probe was centred directly above the 12th and 13th rib

and placed lateral from the backbone of the animal at a point situated approximately in the

middle of the M.l.d. Applying this technique finally yielded chemically analysed average IMF

contents of 3.35 ± 1.12, 3.47 ± 0.93, 3.49 ±1.11, 3.48 ± 1.08, 2.34 ± 0.64 and 2.40 + 0.63 for

AN, SI, CH, LI, BL and PI, respectively. This illustrates that the target value was reached on

average only in four breeds whereas this was impossible for BL and PI. However, BL and PI

were kept in the comparison in order to give an impression of their performance under the

specific conditions of this experimental approach. For further details on IMF content see

(Chambaz et al., 2001b).

5.2.4. Data and sample collection at slaughter

The animals were slaughtered without subsequent electrical stimulation of the carcasses at a

commercial slaughter plant. Hot carcass weights were determined at about 1 h post mortem.

Carcass grading was performed by experienced official staff according to the Swiss beef

classification grid (Proviande, 2001), which is widely equivalent to the EUROP grading

system. Subcutaneous fat thickness was measured 2 days post mortem between the 12* and

13th rib at 3/4 of the distance of the lateral length of the M.l.d. from the backbone with an

electronic calliper square. Carcass length was measured from the 1st rib to the head of the

Symphysis ossium pubis and the leg length from the head of the Symphysis ossium pubis up to

the Os malleolare. After chilling for 48 h, the left side of the carcass was divided into fore-

and hindquarter between the 9th and 10th rib. The flank was cut along the distal edge of the M.

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iliocostalis lumborum through the ribs until joining the cut along the 9l /10l ribs. This

procedure divided the carcass into pistola and forequarter with adhered flank. The pistola was

then separated 5 cm cranial of the head of the Symphysis ossium pubis into leg and sirloin

together with the rump. The separation between the sirloin and the rump was done behind the

6th lumbar vertebra. The sirloin was furthermore dissected into striploin (i.e., M.l.d.),

tenderloin (i.e., M. psoas major), bone and fat tissue. The proportion of the so-called 1st

category cuts comprised the trimmed striploin, tenderloin and rump (M. glutaeus médius).

The dressing procedure followed guidelines ofABZ (1997).

5.2.5. Statistical analysis

Data were statistically analysed with the NCSS program (version 1997, Hintze, Kaysville,

Utah, USA). In a first evaluation, data of both series were included in order to be able to

compare overall series differences, using a two-way ANOVA with breeds and series as fixed

effects and breed x series interactions. Because interactions between breed and series occurred

in most growth variables, the data were finally analysed separately for each series with one¬

way ANOVA considering breed as fixed effect in the model. The Tukey test was used for

multiple comparison among means considering p < 0.05 as significant.

5.3. Results

5.3.1. Growth performance

The steers of the different breeds were of a similar age at the start of the experiment, except

for the PI in series 1 which were older by 40 d at the start as explained (2.1), but there were

already certain initial differences in live weight (Table 2). Weight was highest in the CH

steers (significant in series 2) and lowest in the PI steers (series 2) at the start of fattening. At

slaughter, when the target IMF content was reached by AN, SI, CH and LI steers, AN steers

were significantly lightest in both series. All other groups had a similar final live weight in

series 1. However, in series 2, CH were the heaviest group of those large-framed breeds

which reached the target IMF content. Compared with the weight at slaughter, group

differences in age were more pronounced and consequently the duration of fattening greatly

differed. BL and PI did not reach the target IMF content on average despite fattening period

was three times longer than for AN. BL and PI therefore were on average significantly older

at slaughter than all other breeds, and BL were also heaviest while PI remained in the same

weight range as SI, LI and CH (CH series 1 only).

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Table 2

Growth characteristics of the steers originating from different beef breeds (n=l 1 per series)1

AN SI CH LI BL PI Average SEM

Initial live weight (kg)Series 1 290ab 294ab 295ab 288ab 278b 321a 294z 8.9

Series 2 342" 343b 391a 323b 323b 268e 332y 9.3

Final live weight (kg)Series 1 478b 631a 642a 637a 695a 656a 623z 23.1

Series 2 524d 625c 744ab 698bc 820a 637° 675y 18.5

Age at slaughter (d)Series 1 368d 540c 517c ôos1* 696ab 705a 572 22.5

Series 2 393e 477d 540c 614b 683a 660ab 561 12.3

Days of fatteningSeries 1 146d 319c 304e 371bc 479a 424ab 340y 22.1

Series 2 140e 226d 283c 352b 442a 421a 311z 11.1

Daily gain (g/d)Series 1 1279a loss00 1170ab 951

cd 869d 796d 1025z 39.7

Series 2 1306a 1252ab 1274a 1069e 1123bc 876d 1150y 33.0

DM intake (kg/d)Series 1 7 79abc 8.21ab 8.26a 731e 7.15e 7.5l* 7.70z 0.178

Series 2 8.04b 8.11ab 8 96a 7 79b 8 44ab 6 79e 8.02y 0.213

Total energy intake (GJ NEV)Series 1 7.87c 17.89b 17 20" 18.64ab 23.50a 21.93ab 17 84y 1.240

Series 2 7.41d 12.34° 17.03b 18.51b 23.82a 19.01b 16.36z 0.872

Total protein intake (kg APD)Series 1 101e 226b 218b 236ab 296a 275ab 225 15.5

Series 2 96d 158c 217b 236b 312a 248b 211 11.1

Feed conversion efficiencyFeed DM/gain (kg/kg)Series 1 6.14d 7.66bc 7.1lcd 7 72bc 8.30b 9.63a 7.76y 0.265

Series 2 6.16c 6.5 lbc 7.11ab 7 32a 7.47a 7.79a 7.06z 0.193

NEV/gain (MJ/kg)Series 1 42.6d 52.6bc 49 0cd 53.0^ 57.1b 66.2a 53.4y 1.82

Series 2 40.7C 43.7bc 47.8ab 49.7a 49 3a 51.9a 47.2Z 1.29

APD/gain (g/kg)Series 1 545d ôôô^ 621cd 671bc 720b 830a 675y 22.8

Series 2 529c sôo"0 611ab 634a 646a 677a 610z 16.3

AN = Angus, SI = Simmental, CH = Charolais, LI = Limousin, BL = Blonde d'Aquitaine, PI = Piedmontese

'Means within one line without a common supersenpt differ significantly (P<0 05), series averages within the same variable with different

superscripts are significantly different (P<0 05)

As Figure 1 illustrates, the average live weight gain was quite linear at least in the first 5 to 6

months of fattening, but slightly lower from then on, particularly when regarding the average

final weight and fattening duration. The average daily gains were highest in AN, followed by

CH, SI, and LI. BL had average daily gains similar to LI while PI steers showed the lowest

growth rates from the beginning on. Overall, an inverse relationship between age at slaughter

respectively days of fattening and the average daily gains was observed. The assumed level of

daily gains of about 1.2 kg/d was reached on average only by AN and CH in both series and

SI in series 2 but not by LI, BL and PI.

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800

700

_600

f 500

5

400

300

200

-° *'''

-Jr

-A-AN

-«-SI

-e— CH

—f—LI

-•—BL

-H-PI

i i i i i l i

0 10 20 30 40

Fattening weeks

50 60 70

Fig. 1 : Evolution of the live weight of steers of different beef breeds (average of series 1 and

2). Curves end when the first animal of the respective breed was slaughtered. The last symbol,

connected with the solid line by a dotted line, indicates the average live-weight at slaughter of

all animals per group.

The average daily feed intake over the whole fattening period was, in part, significantly

lowest for LI compared with CH, SI and AN (Table 2). The evolution of feed intake during

fattening (Figure 2) showed a similar pattern for AN, CH and SI in one group and LI together

with BL in another one while PI where lowest over the whole period. The highest feed intake

was generally recorded approximately in the middle of the fattening period, shortly before the

first individuals of the breed groups reached the target IMF and were assigned to slaughter.

The periodically occurring changes in feed intake (Figure 2) could be mainly led back to

changes in batches of grass silage. Feed conversion efficiency in terms ofDM expenditure per

kg gain over the complete fattening period was highest for AN while this efficiency was

lowest for LI, significant against efficiency of SI in series 2 and AN when only comparing

those breeds which reached the target IMF. PI and BL performed even worse in this respect

(Table 2). As diet composition was not changed during fattening, intake and utilization of net

energy and absorbable protein showed the same relative group difference as DM intake.

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According to the days on feed required, the total energy and protein intake was by far lowest

for AN and highest for BL and PI.

10

ë, 8

a>

(04-1

S 7

O

'rä 6a

10 20 30 40 50

Fattening weeks

•x

-A--AN

—--SI

—e--CH

—I--LI

—•--BL

—X—-PI

60 70

Fig. 2: Evolution of daily dry matter intake of the steers of different beef breeds (average of

series 1 and 2). For explanations see Fig. 1.

In series 2, the steers started and finished fattening with a higher weight and had daily gains

higher by 125 g/d on average compared with those found in series 1. The age at slaughter,

however, was similar (Table 2). Differences between the two series in live weight at slaughter

were particularly high for BL, CH, LI and AN with +18, +16, +11 and +10 % in series 2,

respectively, but low in SI and PI. In line with the higher daily gains, feed consumption was

significantly elevated in series 2, but also feed conversion efficiency was improved relative to

series 1, particularly in PI (-19 % feed DM/kg gain), SI (-15 %) and BL (-10 %). Overall, the

steers required 7.4 (6.1-9.6) kg DM, 50 (41-66) MJ NEV and 643 (529-830) g APD per kg

of weight gain.

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5.3.2. Carcass quality

Dressing percentage was significantly different between breed groups being particularly high

in LI and also in BL and PI (Table 3). CH were intermediate and AN as well as SI were lower

in dressing percentage by approximately 8 units in relation to LI. This increased the relative

group differences from live weight to slaughter weight, particularly in series 1. Carcass length

increased with increasing slaughter weight but not generally at the same proportion in all

groups. LI and PI were similar in carcass length as SI despite clearly higher slaughter weights

which is reflected in a better conformation of LI and PI carcasses.

Table 3

Carcass quality traits of the steers originating from different beef breeds1

AN SI CH LI BL PI Average SEM

Hot carcass weight (kg)Series 1 261d 347e 375be 39gabc 447a 413ab 374z 15 3

Series 2 282d 333e 426b 424b 514a 40 lb 397y 10 7

Dressing percentage (%)Series 1 54 5e 55 0e 58 0b 62 5a 64 3a 63 r 59 6y 0 59

Series 2 53 9d 53 4d 57 4° 60 8b 62 8a 62 9a 58 5Z 0 39

Carcass length (cm)Series 1 129e 136ab 134abe 134be 140a 137ab 135 16

Series 2 130e 135bc 140ab 136be 145a ns1* 137 15

Carcass gradingConformation score2

Series 1 2 6a 2 3a 12b llb 12b 14b 1 6 0 14

Series 2 2 5a 2 4a 10b 10b 10b 14b 1 5 0 09

Fatness score3

Series 1 4 4a 4 0ab 3 7b 3 9* 3 0e 3 1e 3 7z 0 14

Series 2 4 8a 42ab 4 4ab 4 5ab 3 7b 2 9e 4 1y 0 19

Subcutaneous fat layer (mm)Series 1 11 8a 12 6a 110ab 11 9a ?0bc 5 7e 10 0 109

Series 2 16 2a 12 5ab 13 lab 12 2b 7 3e 3 4d 10 8 0 90

Means within one line without a common superscript differ significantly (P<0 05), series averages within the same variable with different

superscripts are significantly different (P<0 05)Conformation score C=l H = 2, T = 3, A = 4, X = 5 (widely equivalent to EUROP grading with C = E)3Fatness score 1 (low) to 5 (high fatness) equivalent to EUROP grading

Carcass grading led to a differentiation into two distinct groups of breeds. LI, CH, BL and PI

were generally present in the best conformation class while AN and SI were graded

significantly lower. In contrast, fatness score and thickness of the subcutaneous fat layer only

slightly differed among the four breeds reaching the target IMF content although AN showed

a trend to the highest fatness. Generally these breeds were graded above 4. BL and PI, which

did not reach the target IMF, provided significantly leaner carcasses with a far lower

subcutaneous fat thickness, particularly for PI. The within breed correlations between

thickness of the subcutaneous fat layer and IMF content were significant in BL and CH with

0.50 and 0.46 (p < 0.05), respectively, but not significant in LI (0.27), PI (0.18) and SI (0.03)

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and even negative in AN (-0.30) due to two AN steers with simultaneously high IMF content

and few subcutaneous fat.

AN carcasses showed the lowest pistola proportion (significant against CH and LI in the first

and against BL and PI in both series) but the differences between breed groups were low

compared with those found in other carcass variables (Table 4). However, AN were also

lowest in percentage of 1st category cuts, significantly so against LI and partly also CH. BL

and PI carcasses were high in percentage of 1st category cuts, striploin proportion of sirloin,

lean to bone ratio and particularly lean to fat ratio. In descending order, these variables were

less favourable in LI, CH, SI and AN. Tenderloin proportion of sirloin was highest in BL

while PI, LI, CH and SI expressed intermediate levels and AN carcasses were lowest in this

respect. Most of these group differences were observed in both series although the level of

differences varied to a certain extent. Average values calculated over all groups were also

similar in both series, except the slightly but significantly lower proportion of 1st category

cuts in series 2 relative to series 1.

Table 4

Properties of the valuable cuts in the carcass of the steers originating from different beef breeds1

AN SI CH LI BL PI Average SEM

Pistola (% of carcass weight)Series 1 413° 41 6bc 43 5a 42 9ab 43 2a 43 3a 42 6 0 34

Series 2 41 3b 42 0ab 42 0ab 42 1ab 42 9a 43 3a 42 3 0 34

1st category cuts2

(% of carcass weight)Series 1 6 8e 7]bc 7 3b 7 9a 8 2a 8 2a 7 6y 011

Series 2 6 6e 7 0be 70bc 7 4b 8 0a 8 1a 7 4z 0 10

Sirlom composition

Striploin3 (% of sirlom)Series 1 36 6b 35 7b 36 6b 40 4a 39 6a 39 4a 38 1 0 60

Series 2 34 2b 35 3b 35 9b 38 6a 40 5a 40 5a 37 5 0 50

Tenderloin4 (% of sirloin)Series 1 12 6e 15 6b 15 5b 16 0b 18 5a 16 5b 15 8 0 40

Series 2 12 8e 14 4b 15 lb 14 6b 17 3a 18 5a 15 5 0 35

Lean / fat ratio

Series 1 4 3e 5 2be 57bc 7 0b 12 0a 10 6a 75 0 62

Series 2 3 5d 5 4e 5 2ed 5 5e 9 0b 13 9a 71 0 43

Lean / bone ratio

Series 1 3 9b 3 9b 4 0b 5 3a 5 0a 4 9a 45 0 13

Series 2 3 8d 3 9d 4 2ed 4?bc 5 6a 5jab 45 0 13

Means within one line without a common superscript differ significantly (P<0 05), series averages within the same variable with different superscripts

are significantly different (P<0 05)2Stnploin, tenderloin and rump'Defined as the part between the 9th rib and the last lumbar vertebra of the M longissimus dorsi

4M psoas major

5.4. Discussion

This study compared steers of different breeds deliberately not slaughtered at the same age or

weight but at a target IMF. This implies that differences in growth rates and carcass size-

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related properties between steers of different breed are not reflecting those usually found. The

targeted IMF level was reached on average only in the AN, CH, LI and SI steers, whereas in

BL and PI not a single individual animal was able to reach this level although the fattening

period was largely extended. For this reason, the present results include both a comparison

among those breeds which were potentially able to meet the target level and a comparative

description of properties found in two breeds which were unable to meet this target but were

fattened up to the maximum IMF content possible under the feeding and housing conditions

described.

5.4.1. Growth development ofsteers ofdifferent breedsfattened to a similar IMF content

Steers were already 8 months of age when entering the fattening period because they had been

reared under the relatively extensive conditions of suckler systems. Differences in genetic

growth potential and, possibly, rearing environment led to certain initial weight differences

between breed groups. Using a medium-energy density ration (6.9 MJ NEV/kg DM,

equivalent to 11.2 MJ ME/kg) from then on was found to yield the typical, widely linear

growth development in all breeds with daily gains gradually declining not before several

months of fattening had passed, whereas very intensive feeding gives a clear peak in growth

rate as shown earlier in bulls (Gerhardy et al., 1995). Although the ration was calculated to

provide sufficient energy and protein for average daily gains of 1.2 kg (RAP, 1999), this level

was only reached by the groups fattened for comparably short periods of time (AN, SI and

CH, particularly in series 2). It has to be kept in mind that this assumed level was derived

from data on dairy crossbred steers fattened up to 550 kg. A prolonged period of fattening is

very likely to decrease overall daily gains and, particularly, feed conversion efficiency due to

the increasing proportion of nutrients and energy required to cover demands for maintenance

relative to growth and fat retention (Crouse et al., 1985a).

Accordingly, very low average daily gains and a particularly poor feed conversion efficiency

were found in PI compared with all other breeds. The lower feed conversion efficiency

observed in PI of series 1 is partially due to the age difference at the start of the trial as

animals were older by 41 d in series 1. The BL being slaughtered approximately at the same

age nevertheless performed significantly better than PI although also ranked behind the other

breeds in most fattening performance traits particularly in series 1. This additional difference

to BL could have been at least partly due to the low feed intake capacity of the PI steers which

was obvious throughout the whole fattening period (Figure 2). These findings are in

agreement with the study of Tartari et al. (1988), who compared PI, CH and LI bulls reared

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for 251 days under identical conditions kept individually in a tied stall. Accordingly, PI bulls

had the lowest stomach and gut proportions of empty body weight. This anatomical

disadvantage of PI is enlarged by feeding diets of low energy density (Geay and Micol, 1988;

Tartari et al., 1988). Another component particularly affecting the performance of PI and BL

steers were the housing conditions in series 1, which were unsuitable for the long duration of

fattening and the very high live weights reached. Accordingly, in series 1 eight out of eleven

BL steers and five out of eleven PI steers expressed joint and leg problems during the last

fattening weeks, due to restrictions in movement in the tie-stall barn. This explains the

particularly high difference between series in average daily gains of BL steers whereas series

differences were lower in all other groups, especially in AN with the shortest fattening period.

Furthermore, this illustrates that an unfavourable housing system may affect the expression of

breed differences in performance of steers of suckler origin. Other factors such as diet and

season within year did not differ between series and so presumably were without major effect

in the present study. The importance of individual vs group housing of cattle for the result of a

breed comparison has also been reported earlier (Cole et al., 1964; Mir et al., 1999).

Among the breeds reaching the target IMF content, LI required the most extended fattening

period and had lower growth rates than SI and particularly AN and CH steers. LI are known

for their high feed conversion efficiency compared to other breeds at the same slaughter age

(Geay, 1982; Geay and Micol, 1988), but this was obviously more than compensated by the

adverse effect of prolonged fattening in the present study. This is confirmed by the longest

fattening period and the lowest feed conversion efficiency found by Smith et al. (1976) in LI

crosses when comparing carcasses of purebred AN to AN crosses with SI, CH and LI at the

same IMF content of 5%. LI have a lower digestive tract weight percentage of empty body

weight than CH though still far higher than PI (Tartari et al., 1988). Compared to CH,

voluntary intake of bulky diets in LI therefore seems to be more restricted by the volume of

the abdominal cavity, especially that of the rumen, than by energy requirements (Geay and

Micol, 1988) also suggesting the use of diet of a higher energy content for LI. In line with

this, Geay and Robelin (1979) demonstrated that, for a given body weight, a certain

combination of energy intake and growth rate is ideal in terms of feed conversion efficiency.

Typically, in an advanced growth stage late-maturing cattle show higher weight gains at the

same age than early-maturing animals, provided feed quantity and quality are not limiting

factors (Lehmann, 1979; Jenkins and Ferrel, 1984; Shorthose and Harris, 1991) as possibly

was the case in our study.

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5.4.2. Carcass characteristics ofsteers ofdifferent breedsfattened to a similar IMF content

Large-framed late-maturing cattle are typically physiologically younger at the same

chronological age than small-framed early-maturing cattle and have a higher priority for

protein accretion, especially at limited growth rates (Byers et al., 1988) as was the case in our

study in contrast to typical feedlot conditions (Micol et al., 1993). These prerequisites made

extended fattening periods inevitable to reach or try to approach the target level in IMF. As a

consequence carcasses were much heavier than in the early-maturing AN steers. This

provided best graded carcasses in conformation scores and high dressing percentages in the

late-maturing breeds. Robelin (1986) and Szücs et al. (2001a) showed that the dressing

percentage increases with age and/or weight. Thus, the great differences in slaughter ages

between groups enlarged the differences in dressing percentage. The ratio of leg length to

carcass weight is an indicator of compactness (Keane and Allen, 1998) and well reflected the

better conformation of the later maturing cattle, i.e. BL, PI and LI (data not shown). Among

the four breeds reaching the target IMF content, LI steers showed the best carcass value in

terms of the lean tissue-related properties. The superiority of LI compared to CH, SI and AN

is in agreement with other studies comparing LI and SI (Byers et al., 1988), LI and CH (Geay

and Malterre, 1973; Geay, 1982) as well as LI, SI and AN (Geay and Malterre, 1973; Geay,

1982; Geay and Micol, 1988). Also dressing percentage of LI steers was clearly higher than

that of AN, SI and CH, an effect which was even obvious in crossbred bulls slaughtered at

similar age or weight when comparing LI and SI (with Red Holstein blood) sired bulls

(Gerhardy, 1994).

However, one unfavourable consequence of slaughter according to a high target IMF was that

the 1st category cuts were too big in size for the common retail market, particularly with the

large-framed breeds. Furthermore, the carcasses were excessively fat. It is well-known that

deposition of fat in the body cavity, between the muscles, and in the subcutaneous site occurs

earlier than the deposition of fat within muscle to be noted as marbling (Smith, 1988). This

explains why carcasses in terms of European continental demand were extraordinarily fat

even before a sufficiently high IMF content was reached. Robelin (1978) found no significant

differences in IMF content among Holstein, Salers, Charolais and Limousins bulls, i.e. four

breeds differing in maturity, when their fatness scores were identical, which supports the

assumption that fat deposition in different body sites is related. However, in our study,

correlations between the thickness of the subcutaneous fat layer and the intramuscular fat

were rather low, if significant at all, or even negative (AN). This can be explained by the

deliberately reduced variation in IMF content by applying a target value but possibly also

from the difference in fat retention during growth phases between adipose tissues and within

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muscle. It should be noted that the genetic correlation reported by the American Angus

Association between marbling score and external backfat thickness at the 12th rib is nearly

zero (AAA, 2001). Similar to the results in the other breeds, Campion and Crouse (1975)

found correlations between the same variables in the range of 0.41 to 0.46 depending on sire

and dam breed, whereas the genetic correlation reported by Geay and Renand (1994) was

even higher with 0.64. Regarding the very similar fatness of the large framed breeds which

reached the same IMF content under identical feeding conditions in our study, it may be

assumed that the ratios of undesired subcutaneous fat and intermuscular fat to IMF is more

variable within than between similarly maturing breeds. The AN, by contrast, already

developed significantly more depot fat in relation to IMF and were not able to develop a

highly favourable conformation or muscling within the period required to reach 3.5 % IMF. In

the later maturing breeds, however, the target IMF content together with a good conformation

was only achieved at a rather late growth stage resulting in very heavy carcasses. Overall, the

very good conformation scores can be assumed to be not sufficient to economically

compensate the unfavourably big-sized cuts and the excessive fatness.

5.4.3. Dietary energy concentration requiredfor steers ofdifferent breedsfattened to a

similar IMF content

A limitation in dietary energy density could have been beneficial in the case of the early-

maturing AN steers in terms of feed conversion efficiency while the contrary is true for the

two late-maturing breeds, PI and BL, but also for LI. When using purebred AN instead of

75 % AN steers, as in the present study, a limitation in dietary energy concentration and/or

restricted feeding would have been even more advisable. Also from the carcass quality aspect

the present study provides indirect evidence regarding the search for the ideal energy density

of the diet. Although energy content of the diet was high enough to reach 3.5 % IMF at least

in four breeds, the fattening period presumably could have been reduced by a more

concentrated ration. This would be particularly advantageous for those breeds whose feed

intake may be a limiting factor, i.e. LI and also BL and of course PI, but is highly

recommendable in all breeds, except AN, in order to obtain smaller-sized valuable cuts. In

contrast, for the AN a slightly extended fattening period would also be tolerable from the

viewpoint of slaughter weight which would increase due to the longer time span required to

reach 3.5 % IMF. The final extent of fat deposition in AN steers, however, would probably

not be reduced under these conditions. From the present results it remains open whether or not

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the genetic limitations in BL and PI would still have prevented to reach the level of 3.5 %

IMF when offering a ration of higher energy density.

5.5. Conclusions

Out of the six breeds investigated only AN, SI, CH and LI, turned out to be able to reach the

target IMF content of 3.5 % in a fattening system based on a forage-based diet of medium-

level energy concentration following an 8 months rearing period as suckler beef. BL and PI

did not retain the desired amount of IMF, obviously due to the limited feed intake and the

genetic predisposition for a very high protein accretion. All breeds except AN, however, had

to be fattened for unusually long periods and provided carcasses too heavy to be acceptable

for the common beef market as well as for existing label programs. Although the carcasses of

the AN were of common size, the ratio of carcass fatness to IMF was the most undesirable of

all breeds compared. Generally, the carcasses of all those breeds which reached the desired

IMF content were unacceptably fat. Thus, the present results revealed the difficulty to find an

acceptable compromise between a desired extent of marbling as well as favourable carcass

conformation on one hand and carcass size as well as fat cover which meet market demands

on the other hand.

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6. Characteristics of steers of six beef breeds fattened from eight months of

age and slaughtered at a target level of intramuscular fat.

II. Meat quality

Based on:

A. Chambaz, M. R. L. Scheeder, M. Kreuzer and P.-A. Dufey, 2001,

Arch. Anim. Breed. 44:473-488

Abstract

Meat quality of Angus (AN), Simmental (SI), Charolais (CH), Limousin (LI), Blonde

d'Aquitaine (BL), and Piedmontese (PI) steers (n=22 per breed group) was measured in the

M. longissimus dorsi (M.l.d.) and the M. biceps femoris, regio glutea (M.b.f.). Animals were

fattened in two subsequent series on a forage-based diet until a target level of 3.5%

intramuscular fat (IMF) was reached according to real-time ultrasound assessments in the live

animals or until 15 months of fattening had passed. Series 1 was performed in a tie-stall barn

while a loose-housing system with straw bedding was applied in series 2. The actually

measured IMF contents in M.l.d. were 3.35, 3.47, 3.49, 3.48, 2.34 and 2.40 % for AN, SI, CH,

LI, BL and PI, respectively. Breed group differences in IMF content were mostly

accompanied by a contrary variation either in muscle water or protein content. Muscle

cholesterol levels were similar for all breeds amounting to 47 and 51 mg/100 g on average in

M.l.d. and M.b.f, respectively. Early and late postmortem muscle pH was relatively similar

among breeds, but water-holding capacity, measured as losses due to drip, ageing, thawing

and cooking, was unfavourably high in AN (drip loss excepted) in both muscles. Cooking loss

tended to be lowest in PI, drip loss in SI. The AN showed the palest meat. In line with

lightness, heme iron contents were clearly lowest in both muscles in the AN steers. There was

no relationship found between IMF and shear force among breed groups. No significant

differences between breed groups occurred in M.l.d. collagen solubility and shear force. Apart

from breed differences, there were several differences noted between fattening series, namely

clearly better water-holding capacity and lower shear force of the meat from series 2 (group

housing) than from series 1 (tied system). The results indicate that in steers of similar IMF

content and raised under the same feeding and management conditions, differences in most

M.l.d. and M.b.f. quality traits were apparent, with the exception of shear force and M.l.d.

collagen solubility.

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6.1. Introduction

Interest in label beef production is consistently increasing in Europe (Neumann and Martin,

1991; Gerhardy, 1994; Roche et al., 2000) in a situation of declining consumption of red meat

and a general decrease in acceptance of conventionally produced and marketed beef. Labels

intend to provide a quality which is or can be especially oriented towards consumer demands

including controlled beef quality and a greater consideration of the welfare of the animals

(Boissy et al., 2000). Although pricing systems still heavily rely on carcass rather than on

actual meat quality traits, labels are expected to guarantee a certain level and, particularly,

homogeneity of beef quality (Branscheid and Claus, 1989) in order to fulfil the consumer's

implicit expectations (Temisan, 1990). Current Swiss beef labels encourage both the use of

purebred beef breed cattle in order to profit from a higher carcass quality and, as category,

steers to improve meat quality. This should provide products which can be distinguished from

conventional continental European beef which is based on calves largely representing a by¬

product of dairy husbandry. Visual perception of the products is often associated with beef

quality and can affect purchase decision of the consumers. One main component of visual

quality, together with colour, is marbling which expresses proportion and distribution of

intramuscular fat (IMF) in the M. longissimus dorsi (M.l.d.). It is still unclear in how far

known breed differences in meat quality traits such as water-holding capacity and tenderness

(Dufey, 1987, 1988a; Tatum et al., 1990; Wheeler et al., 1996) can be recovered under the

condition of a similar extent of marbling. Furthermore, the repeatedly postulated low

cholesterol content of Piedmontese beef (Montana Range, 2001) might get lost if really

existing when the cattle is fed to high IMF contents. The objective of the present investigation

was to compare the meat quality of six beef breeds fattened under the same conditions. The

animals of this study were slaughtered at a target level of 3.5 % IMF. This level is higher than

usually found in Switzerland (Dufey and Chambaz, 1999b). As reported in a first

communication (Chambaz et al., 2001a), clear differences in growth and carcass quality

developed between the breeds. Two breeds were unable to reach the target value in IMF, but

were kept in the comparison in order to give information of their meat characteristics under

the conditions applied.

6.2. Materials and methods

A total of 132 steers of six beef breeds, namely Angus (AN), Simmental (SI), Charolais (CH),

Limousin (LI), Blonde d'Aquitaine (BL), and Piedmontese (PI), selected at 8 months of age

from suckler beef rearing systems, were fattened in two subsequent series (11

animals/breed/series). The first series was carried out in a tie-stall barn and series 2 in a loose

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housing system with straw bedding. The animals had ad libitum access to a diet consisting of

maize silage, 520 g, grass silage, 260 g, and concentrate, 220 g per kg of dry matter. Steers

either were slaughtered when the ultrasonically assessed IMF reached 3.5 % in the Musculus

longissimus dorsi (M.l.d.) between the 12th and 13th rib or after a maximum of 15 months of

fattening (Chambaz et al., 2001a). This approach resulted in large differences in slaughter age.

Members of the AN, SI, CH, LI, PI and BL groups finished the experiment at an average age

of 381, 509, 529, 610, 683 and 690 days, respectively. The coefficient of variation in

slaughter age was lowest for AN with 6.6 % and highest for CH with 19.6 %. Further details

on the experimental procedure are given in (Chambaz et al., 2001a).

The animals were slaughtered in a commercial slaughter plant after approximately 1 h of

transport by bleeding after captive-bolt stunning. Measurements of pH were performed 1 h

and 48 h post mortem (p.m.) in the M.l.d. at the 10th rib and in the Musculus biceps femoris,

regio glutea (M.b.f.) with a portable pH meter (WTW 197S, Wissenschaftlich-Technische

Werkstätten GmbH, D-Weilheim) equipped with a Sensor EB4 probe (Wintion, CH-

Gerzensee). Samples of the M.l.d. and M.b.f. were taken from the left carcass side 48 h and

14 d p.m. Meat colour was determined with a Chroma-Meter (CR-300, Minolta, CH-

Dietikon-Ziirich) applying the light source D65, which yielded data for L* (lightness; 0-100 =

dark-light), a* (red-green index) and b* (yellow-blue index) when directly put onto fresh cuts

of M.l.d. and M.b.f. 48 h p.m. These samples were sealed under vacuum as slices of 2 cm

thickness. Separate samples of approximately 300 g were frozen at -30 °C until analysed for

chemical composition, heme iron and cholesterol content. Two slices, also vacuum-packed,

were stored for 12 days at +2 °C for measurements in the aged meat. Weight loss during this

period was determined as ageing loss. Afterwards the samples were stored frozen (-30 °C)

until being thawed at 2-A °C over 24 h and then broiled for 5 min on a grill (type BP-50,

Beergrill AG, CH-Zürich) at 195 ± 5 °C by direct radiant heat with the samples repeatedly

turned during heating. The apparatus was connected to an external electronical

thermoregulator (Ematherm A, Trafag AG, CH-Männedorf) and a thermoprobe (Pt 100,

Moser AG, CH-Hombrechtikon) to control temperature. According to preliminary

assessments, this procedure resulted in a meat core temperature of approximately 68 °C.

Within these procedures, losses due to thawing and cooking (directly after cooking) were

recorded. In independent samples, drip losses were quantified as described by Honikel (1998)

storing fresh 2 cm thick slices of each muscle for 48 h at 2 °C. Cooked samples cooled to

ambient temperature were sheared by the original Warner-Bratzler device (model 3000, G-R

Electric MFG Co, Manhattan, Kansas, USA). Ten cores per sample of 1.27 cm diameter were

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obtained parallel to fibre orientation from the cooked slices according to Kastner and

Henrickson (1969) with an electrical drill at a speed providing uniform samples.

In the raw, homogenised muscle samples, heme iron contents were determined as pigments

according to Barton (1967) by extracting pigments with acetone and spectral photometric

determination at a wavelength of 640 nm on a Lambda 2 photometer (Perkin-Elmer, D-

Überlingen). Heme iron was calculated assuming the conventionally applied value of 9.06 %

heme iron in pigment. Cholesterol was enzymatically determined by a colorimetric method

(Boehringer Mannheim, 1994) with the same spectrophotometer as described for heme iron

analysis, but at a wavelength of 405 nm. As described in the guidelines (Boehringer

Mannheim, 1994), homogenized muscle samples were hot saponified during 30 min in

advance of the cholesterol determination. Lyophilized samples of M.l.d. and M.b.f. were

analysed for their contents of dry matter (3 h, 105 °C) as well as ash (total ash; 4 h, 550 °C;

Naumann and Bassler, 1997), fat (petrol ether extract; SLB, 1969) and protein (crude protein;

KJELDAHL method; AOAC, 1995). Furthermore the lyophilized samples were analyzed for

collagen content (hydroxyproline x 8) as described by Arneth and Hamm (1971) adapted to

the Technicon (Plainfield, New Jersey, USA) analyse chain. Collagen hydrothermal solubility

at 90 °C was determined as outlined by Kopp et al. (1977). All chemical analyses were carried

out in two replicates.

Data were statistically analysed with the NCSS program (version 1997, Hintze, Kaysville,

Utah, USA). In a first evaluation, data of both series were included in order to be able to

compare overall series differences, using a two-way ANOVA with breed and series as fixed

effects and breed x series interactions. Because interactions between breed and series

frequently occurred in meat quality traits, data were finally analysed separately for each series

by one-way ANOVA with breed as fixed effect in the model. The Tukey test was used for

multiple comparison among means regarding P < 0.05 as significant.

6.3. Results

On average of both experimental series the actually measured IMF contents in M.l.d. (means

± SD) were 3.35 ± 1.12, 3.47 ± 0.93, 3.49 ± 1.11, 3.48 ± 1.08, 2.34 ± 0.64 and 2.40 ± 0.63 %

for AN, SI, CH, LI, BL and PI, respectively (Table 1). The corresponding levels found in

M.b.f. were 3.35 ± 0.98, 3.92 ± 0.94, 3.02 ± 1.14, 2.69 ± 0.93, 2.01 ± 0.75 and 1.71 ± 0.65 %.

Therefore, at an overall slightly lower average IMF content of M.b.f. of 2.78 % compared to

3.09 % in M.l.d., the breed differences were roughly also reflected in M.b.f, particularly with

again the lowest contents being found in BL and PI (partially significant against other breeds).

Part of the within-breed variation in IMF content resulted from significant series differences

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44

as the average IMF content achieved was lower in series 1 than in series 2. The M.l.d and

M.b.f. of BL and PI had the highest protein content of all groups on average of both series,

particularly when compared with AN, SI and CH, whereas LI held an intermediate position.

Breed group differences in IMF content were mostly associated by contrary variations in

muscle water and protein content. In line with protein and dry matter content there was a

small but partially significant variation in ash content of the two muscles. In contrast, no

significant differences were observed between breed groups in cholesterol content of both

muscles ranging at a similar level of around 47 and 51 mg/100 g in M.l.d. and M.b.f.,

respectively.

Table 1

Chemical composition of the M 1 d and the M b f (wet weight) of the steers originating from different beef

breeds (n=l 1 per series)'

AN SI CH LI BL PI Average SEM

M longissimus dorsi

Water (g/100 g)Series 1 74 74ab

Series 2 74 30a

Ash (g/100 g)Series 1 0 99

Series 2 0 99d

Protein (g/100 g)Series 1 2129

Series 2 2106d

Fat(g/100g)Series 1 2 99ab

Series 2 3 70a

Cholesterol (mg/100 g)Series 1 47 5

Series 2 47 1

M bicepsfemonsWater (g/100 g)

Series 1 76 29a

Series 2 74 78

Ash (g/100 g)Series 1 0 96d

Series 2 0 98e

Protein (g/100 g)Series 1 19 78e

Series 2 19 58e

Fat (g/100 g)Series 1 2 80b

Series 2 3 89a

Cholesterol (mg/100 g)Series 1 52 0

Series 2 52 0

AN = Angus, SI = Simmental, CH = Charolais, LI = Limousin, BL - Blonde d'Aquitaine, PI = Piedmontese

'Means within one line without a common superscript differ significantly (P<0 05), series averages within the same vanable with different

superscripts are significantly different (P<0 05)

The differences in early postmortem muscle pH were small and mostly not significant among

groups for both muscles (Table 2). Moreover none of the animals presented a pHi-value

below 6.2, indicating that no PSE (pale, soft, exsudative) meat was found. Ultimate muscle

74 03b

73 64at

101

103ec

2147

21 82bt

3 50a

3 43*

47 5

46 4

74 40e

75 02

0 97e<

1 llal

19 80e

19 57e

4 00a

3 83a

51 9

51 8

74 55al

73 69al

102

106"

2126

21 43el

3 30al

3 69a

48 4

45 8

75 83al

75 33

0 99e<

1 04"

19 88e

19 55e

2 57tx

3 46a

50 2

50 8

74 41at

72 89b

101

106ot

2168

22 24at

2 88at

4 07a

48 8

45 6

7531atx

74 61

101"0

1 18a

20 66b

20 65b

2 28be

3 10ab

51 1

50 4

75 29a

73 86*

0 98

1 10b

2155

22 92a

2 25b

2 44b

48 7

46 4

75 66al

74 98

104al

1 06*

21 18a'

21 16al

173°

2 29"

50 6

50 2

75 17a

74 14a

0 97

1 18a

2149

22 57al

2 49at

2 30b

48 0

47 8

74 98"'

75 21

106a

109al

2160a

2170a

188*

154e

49 7

50 5

74 70y

73 75z

100z

107y

21 46z

22 01y

2 90z

3 27y

48 ly

46 5Z

75 41y

74 99z

101z

1 08y

20 49

20 37

2 54z

3 02y

50 9

510

0 273

0 294

0 015

0 017

0218

0219

0 268

0 280

0 98

0 68

0 239

0 245

0 010

0 032

0 163

0 154

0 254

0 262

146

081

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45

pH, measured at 48 h p.m., was in the desired range with an average of 5.54 (max: 5.82) in the

M.l.d. and of 5.50 (max: 5.79) in the M.b.f., when pooled over both series (slightly but

significantly higher in series 2). None of the animals expressed an ultimate muscle pH above

6.0, regarded as the threshold level for DFD (dark, firm, dry) meat or DCB (dark cutting

beef). In series 1, LI muscles had a significantly lower pH4g in the M.l.d. compared to PI, with

the other groups remaining intermediate.

Table 2

Development ofpH ofthe M 1 d and the M b f of the steers originating from different beefbreeds (n=l 1 per

senes)1

AN SI CH LI BL PI Average SEM

M longissimus dorsi

pHihSeries 1 6 54 6 57 6 54 6 48 6 67 6 60 6 57 0 054

Series 2 6 61ab 6 62ab 6 65a 6 63ab 6 63ab 651b 661 0 034

pH48hSeries 1 555ab 5 56ab 5 49ab 5 45b 5 53ab 5 58a 5 53z 0 027

Series 2 5 55ab 5 56ab 5 52b 5 56ab 5 58a 5 59a 5 56y 0 014

M bicepsfemorispHlh

Series 1 6 49 6 53 6 56 651 6 58 6 57 6 55z 0 078

Series 2 6 64 6 66 6 66 6 67 6 60 6 64 6 64y 0 039

pH48hSeries 1 5 52 5 49 5 46 5 43 5 52 5 47 5 48z 0 022

Series 2 5 49b 5 52ab 5 51ab 5 52ab 5 54a 5 55a 5 52y 0 012

Means withm one line without a common superscript differ significantly (P<0 05), series averages within the same vanable with different

superscripts are significantly different (P<0 05)

Table 3 describes traits of water-holding capacity of the meat. The LI showed the highest

M.l.d. drip loss on average of both series. In M.b.f. the pattern was not the same, with the BL

having the highest and AN and SI showing the lowest drip loss. Drip loss was higher in both

muscles in senes 2 especially in BL, whereas AN meat showed an inverse pattern. PI had the

lowest ageing losses in both series for M.l.d. and M.b.f, whereas M.l.d. of BL and SI (only

series 2) and M.b.f. of AN showed significantly higher ageing losses. On average of both

series, the AN expressed the highest thawing and cooking losses for both muscles whereas the

losses were lowest in BL and PI. This illustrates that, within the different losses measured for

the same muscle, only thawing and cooking losses, making up the major proportion of total

losses, expressed a similar pattern over breeds. Accordingly, the overall water-holding

capacity was low in AN and high in PI. Muscles of animals in series 2 expressed a lower

cooking loss than those in series 1. Water-holding capacity of M.b.f. was on average better

than that of M.l.d. in all types of losses measured.

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46

Table 3

Water holding capacity of the M 1 d and the M b f of the steers originating from different beef breeds (n=l 1 per

senes)'

AN SI CH LI BL PI Average SEM

M longissimus dorsi

Drip loss (%)Series 1 3 09b 2 57b 3 54ab 4 30a 2 92b 2 54b 3 16 0 271

Series 2 1 76d 3 23"° 372abc 4 26ab 4 29a 2 89c 3 36 0 253

Ageing loss (%)Senes 1 3 94 3 66 3 86 3 47 4 23 3 44 3 77y 0 214

Series 2 3 40ab 3 78a 3 44ab 3 08ab 3 60a 2 72b 3 33z 0 180

Thawing loss (%)Series 1 7 61a 6 66ab 5 97b 5 77b 5 36b 5 18b 6 09z 0 379

Series 2 8 79a 8 14ab 752abc ôsi* 6 30e 6 72"= 7 33y 0 429

Cooking loss (%)Series 1 22 18a 18 55ab 17 00b 15 37b 15 12b 15 58b 17 3(F 0 903

Series 2 17 30a 14 82ab 14 76ab 12 99b 11 70b 12 03b 13 93z 0 824

M bicepsfemorisDrip loss (%)

Series 1 224ab 144b 2 56a 2 62a 2 77a 2 72a 2 39 0 201

Series 2 1 83e 193c 2 75b 2 91ab 3 45a 2 71b 2 59 0 156

Ageing loss (%)Series 1 3 17a 2 58ab 3 08ab 2 63ab 2 96ab 2 39b 2 80 0 180

Series 2 3 16a 2 70ab 2 61ab 2 92ab 2 85ab 2 54b 2 80 0 138

Thawing loss (%)Series 1 7 05a 5 65ab 6 33ab 5 31ab 5]5ab 4 43b 5 65 0 464

Series 2 7 71a 5 81b 6 51ab 5 29b 5 28b 5 14b 5 96 0 351

Cooking loss (%)Series 1 18 67a 13 32b 15 18ab 12 53b 12 37b 1155b 13 94y 0 896

Series 2 13 58a 11 22ab 13 32ab 12 70ab 10 68b 10 72ab 12 04z 0 692

'Means within one line without a common superscript differ significantly (P<0 05), senes averages within the same variable with different

superscripts are significantly different (P<0 05)

Meat-colour related traits are given in Table 4. The M.l.d. and M.b.f. of the AN and CH steers

were lighter than those of the other groups, and PI steers had the darkest M.l.d. and one of the

darkest M.b.f. In both muscles, BL and PI showed a trend to less reddish and yellowish meat

than that of the other breed groups. As expected, during ageing for 12 days colour turned

towards higher lightness and redness in both muscles (data of non-aged meat not shown). The

variation in M.l.d. lightness during ageing was significantly higher in AN than in SI with

intermediate values in the other breed groups (data not shown). Heme iron content was lowest

for AN and CH steers in both muscles and was highest for PI and SI in M.l.d. as well as for SI

and LI in M.b.f. The M.b.f. heme iron content was higher by 44 % compared with that of the

M.l.d. There was a significant negative correlation between heme iron content and L* in both

muscles (r = -0.77 in M.l.d. and r = - 0.65 in M.b.f., P < 0.001).

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Table 4

Colour after 14 d of ageing and heme iron content of the M 1 d and the M b f of the steers onginating from

different beef breeds (n=l 1 per series)'

AN SI CH LI BL PI Average SEM

M longissimus dorsi

L*

Series 1 39 8a

Series 2 40 2a

Series 1 14 1

Series 2 14 3ab

b*

Series 1 4 0ab

Series 2 4 5a

Heme iron

(mg/100 g wet weight)Series 1 1 1

Series 2 12

M bicepsfemonsL*

Series 1 38 3

Series 2 38 6

Series 1 16 6

Series 2 17 0

b*

Series 1 47

Series 2 61

Heme iron

(mg/100 g wet weight)Series 1 1 65b

Series 2 1 87b

'Means within one line without a common superscript differ significantly (P<0 05), series averages within the same variable with different

superscripts are significantly different (P<0 05)

Shear forces were not significantly different in M.l.d. among groups, but AN M.l.d. tended to

have low values (Table 5). There was a significant decrease in shear values in the second

senes and especially in SI and PI. Unlike as in the M.l.d., there were significant differences

among groups in shear force of M.b.f., but the trend remained similar. As expected, shear

force and collagen contents were higher in the M.b.f. compared to M.l.d. although at similar

collagen solubility. Muscle collagen content was higher in AN, SI and CH than in LI (only

M.l.d.), BL and PI. There were no significant differences in M.l.d. collagen solubility among

breed groups, but AN tended to have the highest solubility in both series, following the same

trend as in shear force. Collagen solubility in M.l.d. increased in series 2 (+22 %) especially

in the PI, LI and SI.

36 T

2,1 T1

40 r

39 0al38 3al

37 4b36 2"

37 7al

34 5e

36 3e

37 6

38 0

0 73

0 60

14 3

14 7a

13 9

14 5a

14 8

14 6a

13 8

13 3b134

13 7al14 1

14 2

0 39

0 25

4 1al

4 3al4 7al

4 5a

5 2a

4 2al3 8al

3 6b3 5°

2 9e

42

40

0 42

021

150a

1 52ab

1 13"

131a

128ab

138ab

146a

136ab

159a

1 61a

135

140

0 075

0 082

36 4"

36 6b38 9a

36 8al374ab

36 2b36 9ab

37 lab36 8ab

36 7ab37 5

37 0

0 58

0 48

17 0

17 3a

169

17 3a

16 7

171ai16 3

16 7al16 3

16 3"16 6Z

17 0y

031

0 22

54 62

5 8al57

5 3al50

5 0al50

4 8"53

55

0 42

0 28

2 22a

217ab

175"

2 12al

2 06al

2 27a

194al

2 03al

199al

2 or1

194z

2 08y

0 101

0 080

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48

Table 5

Collagen properties and shear force of the M 1 d and the M b f of the steers originating from different beef

breeds (n=l 1 per series)'

AN SI CH LI BL PI Average SEM

M longissimus dorsi

Collagen (mg/100 g wet weight)Senes 1 537a 536a 515ab 470ab 481ab 438b 496 197

Senes 2 549a 541a 560a 496ab 426e 476bc 508 154

Collagen solubility (%)Senes 1 30 5 28 8 28 5 23 1 28 4 27 0 21T 1 89

Senes 2 35 6 34 8 31 9 33 0 30 6 36 1 33 7y 136

Warner-Bratzler shear force (N)Senes 1 30 8 34 8 32 4 31 2 33 1 37 3 33 y 1 81

Senes 2 28 5 27 9 30 5 28 8 27 3 26 1 28 2Z 1 28

M bicepsfemorisCollagen (mg/100 g wet weight)

Senes 1 594abc 695a 615ab 539^ sii* 488e 574z 26 9

Series 2 677a 604ab 680a 638a 495e 529be 604y 22 4

Collagen solubility (%)Series 1 35 0a 26 5b 32 8ab 31 lab 28 lb 29 0ab 30 4 1 54

Senes 2 33 8a 33 2a 33 4a 31 4ab 26 5be 25 3° 30 6 143

Warner-Bratzler shear force (N)Senes 1 33 2b 46 0a 38 0ab 40 8ab 37 6ab 34 4ab 38 4y 2 88

Senes 2 28 8be 32 1abc 38 0a 34 4ab 35 0ab 25 8C 32 4Z 1 91

Means within one line without a common superscript differ significantly (P<0 05), series averages within the same variable with different

superscripts are significantly different (P<0 05)

6.4. Discussion

6.4.1. Realized contents ofintramuscularfat in steersfed on aforage-based diet

Steers of four out of the six beef breeds evaluated were able to reach on average a target level

of 3.5 % IMF in the M.l.d. which was desired in order to ensure the favourable impression of

marbling (Chambaz et al., 2001a). However, no single PI and BL steer was able to reach this

target level although age was almost twice as high as in AN when fattening was finally

terminated (Chambaz et al., 2001a). This illustrates that some cattle breeds obviously do not

have the inherent ability to deposit increasing amounts of IMF regardless of the length of the

fattening period, although carcass fatness is still increasing in this period (Smith, 1988). Most

other breeds show a widely linear increase not only in adipose tissue but also in IMF up to

high slaughter ages as described for instance by Szücs et al. (2001a; 2001b) for German SI

bulls. The high variability between animals of the same breed group in IMF content probably

was mainly due to the restricted accuracy of the ultrasound method of IMF determination in

live animals. However previous attempts analysing biopsies, as another potential way of

estimation, proved to be far less precise because of the inhomogeneity of the distribution of

fat in the muscle.

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49

6.4.2. Differences in meat quality ofsteers ofdifferent breedfattened to a target IMF level

As expected there was an antiparallel relationship of the breed group differences in IMF and

other compositional traits. The differences in chemical composition of muscle between BL

and PI on one hand with low IMF and AN, SI, CH and LI on the other hand are in accordance

with the results of Browning et al. (1990) who reported that muscles of leaner carcasses were

higher in water and protein content. Nevertheless, the close inverse relationship between

moisture and fat content described by these authors and others (Van Koevering et al., 1995)

was less clear in the present study, especially in the M.b.f. where variations in IMF content

were more associated with variations in protein content. This is probably due to the

deliberately low differences in IMF. Accordingly, breed group differences in muscle protein

and fat content were of very low importance in a dietetic sense. This is also true for muscle

cholesterol content which was similar in all groups. Even muscles differed only slightly but

equally by about 2-3 mg/100 g muscle tissue. This is in accordance with the results found by

Eichorn et al. (1986), Baker and Lunt (1990) as well as Gariepy et al. (1999). Rhee et al.

(1982) observed only a significant difference when comparing seven marbling-score

categories when muscles "practically devoid" of marbling were compared with higher

marbling scores which had higher cholesterol levels. However this contradicts findings of

Slover et al. (1987) and Van Koevering et al. (1995) reporting a positive relationship between

fat and cholesterol content of raw beef. Browning et al. (1990) also noted a trend towards

higher cholesterol levels from lean to fatter carcasses, but the difference of only 1.8 mg

cholesterol/100g raw meat found is of little practical importance. Probably based on its low

IMF content, there is a frequently cited opinion that meat of PI cattle has a considerably lower

cholesterol content than meat of other breeds (Montana Range, 2001). However, neither the

present data, where PI steers had been fattened to untypically high IMF contents for this

breed, nor other scientific studies (Baker and Lunt, 1990; Gariepy et al., 1999; Rule et al.,

1999) found any clear differences in muscle cholesterol between PI or PI crosses and other

breeds.

Although PSE meat is not a major problem in beef, Dufey (1987; 1988b; 1989) occasionally

recorded too low pHi-values in meat of bulls fattened in a tied stall, whereas no PSE meat

was found here even in series 1 where the animals were kept under the same housing

conditions as in the studies cited. The absence of DCB in this study is not surprising. DCB is

a result of a reduced glycogen content in the muscle prior to slaughter and is often associated

with stress caused for instance by mixing cattle or shipping fatigue. The duration of transport

to the slaughter plant did not exceed 1 hour. Moreover the use of steers instead of bulls

reduced the probability of DCB as steers have lower stress susceptibility (Warriss, 1990). In

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50

normal beef, p.m. glycolysis reduces pH to 5.8 or lower within 48 h (Kreikemeier et al., 1998;

Immonen and Puolanne, 2000). In an ultimate pH range of 5.8-6.0, defined as borderline

DCB, meat already tends to have an abnormal colour and an increased risk of spoilage,

particularly when vacuum-packed (Warriss, 1990), although quality is still intermediate

between true DCB and normal beef (Voisinet et al., 1997). This type of borderline dark

cutters was found here in two out of 132 carcasses. Despite the low variation in pH, breed

groups differed to a certain degree in water-holding capacity, meat colour and tenderness

related items. The range in drip loss found in the present study was mostly favourable as a

level of up to 4.5 % is still considered as acceptable (Ender and Augustini, 1998) whereas

higher rates are undesired in retail packaging and so impair the appearance of the product at

sale. Muscle tissue of M.l.d. but not of M.b.f. from LI steers showed drip losses on average

just at the acceptable limit, and several members of this group had too high drip losses from

M.l.d. according to the threshold level given above. Cooking losses were highest in both

muscles in AN steers. Accordingly, differences between breeds were not related between drip

loss and cooking loss, except in PL This can be explained by the different compartments of

bound water being stressed by the procedures. Drip losses are passive losses which strongly

depend on loosely-bound water only affected by gravity, whereas meat was subjected to

mechanical and thermal stressors along with ageing (vacuum), freezing/thawing and cooking.

Accordingly, no correlation between drip loss and cooking loss was noted earlier (Honikel,

1986). Cross et al. (1984) and Crouse et al. (1985b) did not find a difference in cooking losses

between AN, SI, CH and Hereford as well as between AN and SI, respectively. The level of

cooking losses were about two-fold lower in our study compared to those found by Gerhardy

et al. (1995) with meat from 16 and 20 months old bulls and still lower than those found in

young heifers by Scheeder et al. (1996) of 24 % and 25 % for M.l.d. and M.b.f, respectively.

However, variation in cooking methods might have played an important role in this respect.

Apart from marbling, meat colour influences the visual appeal of meat to retail purchasers

(Shorthose and Harris, 1991) and thus is an important criterion for purchase decision of the

consumer (French et al., 2000). Consumers appear to prefer beef which is neither extremely

pale nor dark, with the range of L* values between 34 and 40 being considered as normal

(Ender and Augustini, 1998). On this basis, the meat of some AN and CH steers can be

judged as to be too pale. An increasing IMF content tends to increase meat lightness

(Shorthose and Harris, 1991; Frencia and Monvoisin, 1993), but the differences between BL

and PI on one hand and the four other breeds on the other hand were not consistent.

Obviously some independent breed differences in lightness exist. Similar to the present study,

Liboriussen et al. (1977) found a higher heme iron content of the M.l.d. of SI sires compared

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51

with LI, CH and BL. However, the LI meat was lighter than that of CH which was not the

case in our study. Accordingly, the heme iron content, the effective part of the pigments of

bovine meat (Renerre, 1982), similarly varied between breeds as the L* value. Heme iron is

also of interest for human nutrition because of its high bioavailability (Varnam and

Sutherland, 1995). In this view, AN and, partially, CH beef is inferior to beef of the other

breeds.

Overall, the animals of the present study provided a very tender M.l.d. applying the thresholds

of 38 N in Warner-Bratzler shear force for very tender meat and of 45 N for tender meat as

given by Shackelford et al. (1991) and Van Koevering et al. (1995) which were obtained

under similar measurement conditions. In accordance with Scheeder et al. (1996), the

differences in peak shear force found between M.l.d. and M.b.f. were low. Thus even the

M.b.f, where shear forces were extremely low for PI and AN in series 2, could be used as

steak instead of using it as lower priced cuts, particularly when size of this cut is not limiting

due to big sized carcasses. Only the M.b.f. of the SI in the first series had to be judged on

average as too tough to be marketed as steak. However, Scheeder et al. (1996) noticed that

peak shear force data provide little explanation for the collagen component of tenderness thus

indicating that there might nevertheless exist clear differences in tenderness ofboth muscles.

There was no clear relationship between IMF content and shear force among breed groups.

The residual variation in IMF content among the breeds after fattening to a similar IMF level

was contrary to the repeatedly stated inverse relationship between IMF and shear force

(Augustini and Liidden, 1992; Shackelford et al., 1994a). For instance PI meat in series 2 had

the lowest IMF content and the lowest shear forces in both muscles. This is in accordance

with Shackelford et al. (1994b) who found that meat from PI crosses with AN or Hereford

dams, despite having a lower IMF content as the steers in our study, showed the lowest shear

force among eleven genetic groups including AN and CH. This finding is also well supported

by Tatum et al. (1990) who compared Gelbvieh, Red AN and PI sires. Blumer (1963) and

Dikeman (1996) reported that IMF content is a poor predictor of tenderness and only accounts

for 5 to 10 % of the variability in tenderness. Wheeler et al. (1996) also found an increase in

tenderness at a common marbling end point when using PI crosses with various dams instead

of twelve other breed sires. Koch et al. (1976) observed a difference of less than 3 N in shear

force among AN and CH on one hand and LI and SI on the other hand when data were

adjusted to a similar slaughter age. Though statistically significant, the differences were small

and the values remained in the acceptable range of tenderness. Dufey (1987) compared

tenderness of M.l.d. samples from bulls of purebred SI, SI with 75 % Red Holstein blood,

pure-bred Swiss Braunvieh and its crossing with 75 % Brown Swiss blood. The sensory

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52

analysis revealed that the SI bulls had the toughest meat apart from the Brown Swiss crosses,

and an improvement in meat quality took place with a high Red Holstein blood proportion. In

another study, Dufey (1988a) compared SI and Brown steers, their crosses with AN, and

Holstein in taste panel evaluations of tenderness for M.l.d. and M. semimembranosus samples

with the result that meat from SI steers was rated low in tenderness. Crossbreeding with AN

positively influenced tenderness. In a third study, Dufey (1989) compared tenderness of

M.l.d. samples from Fl bulls sired by SI and BL with various dams (SI, Brown Swiss,

Holstein). No significant differences were found in tenderness between the sire groups.

Branscheid and Herzog (1996) noted that crossbreeding of German SI with CH and PI

improved tenderness (shear force and panel tenderness). Accoridng to this trend described in

literature, the present data also showed a higher shear force of meat of SI steers compared to

the other breed groups but this tendency was only apparent in the first series.

6.4.3. Fattening series differences in meat quality ofsteers (tied housing vs loose housing)

Steers of the present study originated from two different fattening series, and housing system

was changed from a tied system to group housing on straw beds without change in the diet.

The statistical evaluation of the data revealed several effects of fattening series on meat

quality. In detail, the IMF content was higher, the cooking losses were clearly lower, collagen

solubility in M.l.d. was higher and, finally, shear force was lower in series 2. From the design

chosen, no clear attribution of the effects to housing system is possible and time-dependent

effects as well as random trends in the animals selected from the breeds cannot be excluded.

However, based on the results of other studies, some of the effects might nevertheless be

explained by the variation in the housing situation. Using the tie-stall barn in series 1 resulted

in a restricted movement and subsequent joint and leg problems which reduced growth rate at

the end of fattening for the oldest animals (Chambaz et al., 2001a). Management practices

which alter growth and muscle accretion rates can have a profound effect on muscle

proteinases and collagen characteristics.An improved growth rate may result in a decreased

calpastatin activity at 24-31 h p.m. and consequently in an improved tenderness (Shackelford

et al., 1994b). Thomson et al. (1996) found that meat from steers growing fastest prior to

slaughter was more tender and had higher /i-calpain and calpastatin activities 2 h p.m. They

concluded that a high /x-calpain activity is likely to be related to a high rate of protein

degradation and this should result in an increase of the myofibrillar fragmentation index and,

therefore, tenderness. From their study they presumed that the level of/i-calpain activity close

to slaughter had been more important in determining tenderness than the calpastatin activity at

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2 h p.m.. Fast-growing cattle would also have a more intense protein turnover and therefore

fewer heat-stable intermolecular collagen crosslinks (Fishell et al., 1985; McCormick, 1994).

This could partly explain the higher collagen solubility and the lower shear force in series 2 in

all breed groups. As collagen solubility was higher in M.l.d. in series 2, but not in M.b.f.,

certain additional series differences in the animals selected can be assumed. SI and PI had the

highest series difference in slaughter age (63 days and 45 days younger in series 2 than in

series 1, respectively; Chambaz et al., 2001a) and simultaneously showed the highest

improvement of shear forces in series 2. Although the increase in the toughness of meat with

animal age is well known (Shorthose and Harris, 1990) it can be assumed that these effects

were not determined by age alone. Firstly, all breed groups showed a decrease in shear force

even those which were older in series 2 than in series 1 such as AN, CH and LI (25, 23 and 9

days older in series 2). Secondly, there were no significant correlations between age and shear

forces within breed groups and series (data not shown). However from the age-related

changes in collagen properties, nevertheless a certain decrease in tenderness with age can be

expected. It is difficult to explain the better water-holding capacity found in series 2 by a

single factor since animal genetics, age and slaughter conditions are known to significantly

contribute to the expression of variations in these traits.

6.5. Conclusions

The overall goal of the present study was to compare a wide range in beef breeds in their

differences in meat quality when a constant IMF content of 3.5 % is achieved. The target level

in IMF, however, excluded two breeds, namely PI and BL, as not appropriate for this attempt.

In the other four breeds, the target was reached on average but individual variation in IMF

was still high within breed due to the restricted accuracy of the in vivo ultrasound method

applied for determination. From the present material, without further attempt to reduce

variability in IMF content, certain breed differences in important quality traits such as colour,

water-holding capacity and tenderness-related variables were found. Differences were

particularly high between AN and the other suitable breeds (SI, CH, LI) with unfavourable

properties ofAN beef in total water-holding capacity and favourable estimates for tenderness-

related data. However, differences in shear force among breed groups were small, with all

breed groups being well above minimum levels of acceptance, even in a basically tougher

muscle such as the M.b.f, except in SI. The significantly better growth characteristics

obtained in the steers of the loose housing system compared with those in the tie-stall barn

seemed to be the major cause of the improvement in meat quality in the second series. Apart

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from the inability of the BL and PI to reach a high IMF content under the present feeding and

housing conditions, all breeds were suitable for premium beef quality marketing.

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55

7. Meat quality ofAngus, Simmental, Charolais and Limousin steers

compared at the same intramuscular fat content

Based on:

A. Chambaz, M. R. L. Scheeder, M. Kreuzer and P.-A. Dufey, 2001

(submitted to Meat Science)

Abstract

Meat quality and marbling properties of Angus, Simmental, Charolais and Limousin steers

(4 x 16) were compared at an average intramuscular fat content (IMF) of 3.25% in the M.

longissimus dorsi. The steers were fattened on a forage-based diet until the desired,

ultrasonically estimated IMF content was reached and therefore differed considerably in

growth and carcass characteristics. Extent and distribution of marbling was equal for all

breeds. Angus and Charolais provided bright meat with low heme iron content. Angus and

Limousin beef was found to be more tender in a sensory assessment than Simmental beef,

corresponding to the differences found in shear force (non-significant) and myofibrillar

fragmentation index. Flavour was similar among breed groups. Juiciness was highly

correlated with cooking loss, with Limousin and Charolais beef being juicier than Simmental

and Angus beef. In conclusion, clear differences in meat quality were observed between

breeds despite a similar IMF content.

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7.1. Introduction

Meat quality of beef breeds has been measured and compared in numerous studies. These

comparisons, however, mostly concentrated on potential breed differences at a similar age,

fattening period, weight, or fatness score of the animals. Only few studies so far investigated

meat quality of different breeds at the same intramuscular fat (IMF) content or marbling score

(Tatum et al., 1996). However, this was done after statistical adjustment of the means and not

by really fattening cattle until a similar IMF content was achieved, probably also because of

the difficulty to accurately estimate IMF in live animals. Cattle breeds clearly differ in their

capacity for IMF retention and, consequently, breed differences in age and weight are

expected to be large when animals are fed on a similar energy level and slaughtered at the

same IMF content (Koch et al., 1976; 1979; 1982). The influence of IMF content on beef

palatability has often been described but its quantitative importance is controversially

discussed (Dikeman, 1996; Lusk et al., 1999). Nevertheless, the visual appearance of IMF,

commonly called marbling, is the primary criterion for quality grading of beef carcass quality

in the United States and Canada (Dubeski et al., 1997). Moreover marbling is often linked

with beef palatability by consumers and can therefore play an important role on purchase

decision. Focus on marbling is especially decisive in branded beef programs where

occasionally a minimum amount and a fine texture of IMF is defined or demanded (AAA,

2001).

The objective of the present study was to compare the meat quality of one early- and three

later-maturing breeds fattened to the same IMF content under controlled conditions in order to

quantify the residual differences in marbling and meat quality between these breeds.

7.2. Materials and methods

7.2.7. Experimental design

The experiment was carried out with Angus, Simmental, Charolais and Limousin steers

originating from suckler herds. They entered the trial at an average age (± SD) of 238 ± 25 d

and were fattened in two subsequent series. The first series was carried out in a tie-stall barn

and the second series in a loose housing system with straw bedding. During the whole

fattening period the animals had ad libitum access to the same diet consisting of maize silage,

520 g, grass silage, 260 g, and concentrate, 220 g per kg of dry matter with a medium energy

density diet representing a typical semi-intensive European-type of fattening. The animals

were assigned to slaughter when the IMF content, estimated by a real-time ultrasound scanner

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200 (Pie Medical, Maastricht, Netherlands), reached the target level of between 3 and 4 %

(Chambaz et al., 2001). However, as a certain inaccuracy of this assessment had to be taken

into account, twelve animals per breed and series were fattened in order to be able to select

the eight animals per breed and series, which matched the requirements in IMF content best

(target content and similar variation within breed).

7.2.2. Experimentalprocedures performed at slaughter

On the day of slaughter, animals were weighed and transported 60 km to a commercial

slaughter plant and slaughtered within 4 h of departure from the research station. Hot carcass

weight was recorded at about 1 h post-mortem (p.m.). Carcasses were chilled for 48 h at 2 °C.

Measurements of pH and temperature were performed at 1, 3 and 48 h p.m. in the M.

longissimus dorsi (LD) at the 10th rib with a portable pH meter (WTW 197S,

Wissenschaftlich-Technische Werkstätten GmbH, Weilheim, Germany) equipped with a

Sensor EB4 pH probe (Wintion, Gerzensee, Switzerland) and temperature probe (TFK 150/E,

Wissenschaftlich-Technische Werkstätten GmbH, Weilheim, Germany). At 48 h p.m.

subcutaneous fat thickness was measured as described by Boggs et al. (1998), i.e. at 3A of the

lateral length of the LD from the backbone between the 12th and the 13th rib.

The left carcass side was cut between the 9th and the 10th rib, and the so-called 1st category

cuts, i.e. striploin, tenderloin and rump were prepared. Starting at the cranial part, the LD was

cut into slices first removing approximately 300 g for chemical analyses (fat and collagen)

after lyophilisation and homogenisation, followed by three 2 cm thick slices for later

determination of drip loss, cooking loss, Warner-Bratzler shear force (WBSF) and sensory

evaluation. Finally, three 1 cm thick slices were obtained for measurements of marbling traits,

heme iron, sarcomere length and myofibrillar fragmentation index (MFI). Samples were

vacuum-packaged and frozen at -28 °C either directly or after an additional ageing period of

12 days at 2 °C which was applied for a second determination of MFI and for analysis of meat

colour, WBSF and sensory evaluation.

7.2.3. Determination ofintramuscularfat content and marbling

The IMF content was analysed using petrol ether extraction (SLB, 1969). In order to

objectively quantify marbling traits, intramuscular fat of the intact slice was chemically

stained according to Albrecht et al. (1996). Briefly, the LD was soaked for 7 d in a formol

calcium solution, then in a solution of oil red (0.5 g oil red dissolved in 100 g isopropanol) for

7 h, followed by 4 h of washing in a 70 % isopropanol solution with agitation. With this

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method it was possible to distinguish IMF from collagen and to intensify the contrast between

IMF and other components of the LD. A digital photo image of the stained slice was then

captured by a computer assisted video camera. Marbling traits were evaluated by an image

analysis software (analySIS, Soft Imaging System GmbH, version 1999, Münster, Germany).

Variables obtained in each sample were: visible fat as a percentage of total muscle area,

average fat particle size, number of fat particles per cm2, the proportion of the area of the

three largest fat particles, and the number of particles > 30 mm2. All measurements were

carried out separately in each quarter of the cross-section of the LD. The coefficient of

variation of visible fat proportion among quarters within steers was calculated.

7.2.4. Analysis ofmeat quality

Meat colour traits (L*, a*, b*) were determined without blooming in the raw LD with the

Chroma-Meter CR-300 (Minolta, Osaka, Japan) applying the light source D65. Heme iron

was analysed as pigments (assuming 9.06 % heme iron in pigment) according to Barton

(1967). Pigments were extracted with acetone and measured photometrically (640 nm,

Lambda 2 spectral photometer, Perkin-Elmer, Überlingen, Germany).

Drip loss was quantified as described by Honikel (1998). Cooking loss was determined by

weighing the samples before and directly after cooking. For this purpose frozen vacuum-

packaged LD slices were thawed during 24 h at 2 °C and were stored for 1 h at room

temperature before cooking. The slices were then broiled for 5 min on a grill (type BF-50,

Beergrill AG, Zurich, Switzerland) at 195 ± 5 °C by direct radiant heat with the samples

turned two times. The grill plate was directly connected to an external electronical

thermoregulator (Ematherm A, Trafag AG, Männedorf, Switzerland) to minimize temperature

variations. According to preliminary assessments, this procedure resulted in a meat core

temperature of approximately 68 °C.

For shear force determinations on the original Warner-Bratzler device (model 3000, G-R

Electric MFG Co, Manhattan, Kansas, USA), the cooked samples were cooled to ambient

temperature. Ten cores of 1.27 cm diameter per sample were obtained parallel to fibre

orientation with an electrical drill to get uniform samples (Kastner and Henrickson, 1969).

The cores were obtained and sheared always in the same order beginning with the five first

dorsal cores followed by five ventral cores starting from the medial to the lateral end of the

slice. The mean WBSF values of the cores 1 and 2, 4 and 5, 6 and 7, and 9 and 10,

respectively, were combined to describe the positions dorsal-medial, dorsal-lateral, ventral-

medial and ventral-lateral.

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Two replicates of 0.5 g, taken from the centre of the raw LD, were analysed for sarcomere

length (Pospiech and Honikel, 1987). The samples were homogenized for 30 s with an Ultra-

Turrax T25 (Janke & Kunkel, IKA Labortechnik, Staufen, Germany) at 9500 rpm in 5 ml

borate buffer (0.1 M KCl, 0.039 M sodium tetraborat decahydrat and 5 mM EDTA, pH 7.1).

The length of five consecutive sarcomeres was measured 12 times per replicate (in total

equivalent to 120 determinations per animal) using an optical microscope (Olympus BX50,

Olympus Optical Co, Tokyo, Japan) and the same image analyzing software as for marbling.

The MFI was analysed in raw LD as described by Culler et al. (1978). The protein

concentration of the suspension produced with this method was determined by the biuret

method (Gornall et al., 1949). After dilution of the myofibril suspension to a concentration of

0.5 ± 0.05 mg/ml, the final protein concentration was controlled using the micro-biuret

method (Bailey, 1967). The MFI is equivalent to the absorption value of the myofibril

suspension, measured at 540 nm and multiplied by 200.

The collagen content (hydroxyproline x 8) was measured in lyophilised LD as described by

Arneth and Hamm (1971) and adapted to the Autoanalyser II chain (Technicon, Plainfield,

New Jersey, USA). Collagen hydrothermal solubility at 90°C was determined as outlined by

Koppetal. (1977).

7.2.5. Sensory evaluation

Sensory analysis was performed by an eight member, in-house trained panel. Panelists

simultaneously assessed four samples, cooked as described for cooking loss, which were

served hot on pre-warmed plates. Samples originated from one representative of each breed

provided in an arrangement which minimised the variation of IMF content within each

session. Panelists were asked to judge the samples on 8-cm unstructured line scales anchored

at each end with the descriptors very tough/tender, very slight/strong, very dry/juicy, very

much disliked/liked for tenderness, flavour intensity, juiciness and preference, respectively.

The marks of the panelists on the line scales were then converted to numbers by measuring

the position of each mark with a computer-assisted FIZZ digitiser (version 1.30, Biosystemes,

Couternon, France).

7.2.6. Statistical analysis

Data was statistically analysed with the NCSS program (version 1997, Hintze, Kaysville,

Utah, USA) by analysis of variance. The model included breed and series as fixed effects, the

interaction of these effects, and IMF as a covariate. The analysis of the sensory traits

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additionally considered the effects of panel session, panelists and IMF group as block nested

within series. The Tukey method was applied for multiple comparison among breed group

means considering P < 0.05 as significant. The tables give the least square means, the

standard ercor of the mean (SEM) and the level of significance of the effects and interactions.

7.3. Results

Fattening the steers of the different breeds under the same conditions and to a similar IMF

content resulted in significant differences between breed groups in most of the growth and

carcass traits (Table 1). The fattening period of the Angus group was only 0.53, 0.50 and 0.41

of that of the Simmental, Charolais and Limousin groups, respectively. The Limousin steers

had a 13 to 21 % lower growth rate compared to the other breed groups. Carcasses of

Charolais and Limousin were significantly heavier than those of the Simmental and

particularly of the Angus steers. Simmental and Angus had the lowest dressing percentage

and proportion of 1st category cuts, which both were highest in the Limousin. However, breed

groups did not significantly differ in the thickness of the subcutaneous fat layer. There were

also effects of fattening series in some growth and carcass traits, but these did not

significantly interact with the breed group effects.

As intended, the IMF content was almost identical for all breeds (Table 2) with an average of

3.25 %, and also the coefficients of variation in IMF content were quite comparable,

accounting for 27, 21, 25 and 21 % in the Angus, Simmental, Charolais and Limousin groups,

respectively. Similarly, for none of the marbling traits measured significant differences

between breed groups were found. However, there was a trend (P = 0.1) in the LD of the

Angus steers to accumulate the highest proportion of visible fat within the three largest fat

particles. There were significant positional effects on visible fat proportion of total muscle

area (Fig. 1) which varied from 5.1 % on average at the dorsal-lateral position to 9.3 % at the

dorsal-medial position. Steers of the two fattening series differed in IMF content and some

marbling traits, but a significant interaction with breed group was only found for average

particle size which remained unaffected by breed group and fattening series alone.

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O)c

Dorsal-medial Dorsal-lateral Ventral-medial Ventral-lateral

Fig. 1. Area of visible fat as a proportion of total muscle area () and Warner-Bratzler shear

force (WBSF, o) measured at different positions within the LD (average of all breed groups).

Means within variable from different positions lacking a common letter are significantly

different (Tukey, P < 0.05).

There was no significant difference between breed groups in pH and temperature of the LD

measured 1 h p.m. (Table 3). The temperature decline from 1 h to 3 h p.m. was significantly

slowest for the Limousin, the group with the heaviest carcasses and the most pronounced

conformation. This was accompanied by a more rapid decrease of pH from 1 h to 3 h p.m.

compared to all other breed groups (significant against Angus). Breed group differences in

ultimate pH (48 h p.m.) were small but significant between Limousin and Simmental. Drip

loss increased in the order of Angus, Simmental, Charolais and Limousin, while the opposite

trend was apparent for cooking loss. When comparing Limousin and Angus, this resulted on

average in differences of 2.0 and 6.5 percentage units in drip loss and cooking loss,

respectively. The LD of the Angus and the Charolais steers was significantly brighter (high

L*) than that of the Simmental steers, with the Limousin taking an intermediate position. This

is in line with the correspondingly lower heme iron content of the LD in Angus and Charolais

relative to Simmental. Accordingly, there was a significant and negative correlation between

heme iron content and lightness (r = -0.68, P < 0.001). Breed groups did not differ

significantly in redness and yellowness of LD.

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Table

1

Growthandcarcass

traitsoftheselectedsteersfattenedtoasimilarintramuscular

fatco

nten

t1

Variable

Angus

Simmental

Charolais

Limousin

SEM

P-values

Breed

Series

BreedxSenes

Age

(d)

Fatt

enin

gperiod

(d)

Averagedailygains(k

g)Hotcarcassweight(k

g)

Dres

sing

percentage(%)

1stcategorycuts2(%ofcarcassweight)

Subcutaneous

fatlayer3

(mm)

16

16

16

16

381c

499b

513b

594a

16.6

0.000

0.604

0.479

141c

267b

281b

346a

16.3

0.000

0.186

0.474

1.30a

1.18a

1.22a

1.03b

0.325

0.000

0.017

0.741

275c

339b

395a

405a

11.9

0.000

0.099

0.349

54.3c

54.1c

57.9b

61.5a

0.46

0.000

0.011

0.479

6.76c

7.08bc

7.13b

7.62a

0.091

0.000

0.008

0.681

14

12

12

13

0.9

0.330

0.078

0.379

1Leastsquaremeanswithinthesamerowlacking

acommon

lett

eraresignificantlydifferent(Tukey,P<

0.05).

2Sumofthetrimmed

stri

ploi

n,tenderloinandrump

3Measuredbetweenthe

12th

andthe

13th

rib.

Table2

Intramuscular

fatcontentandvideoimagean

alysed

marbling

traitsofM

longissimusdorsi(LD)obtainedfromsteersofdifferentbreeds

Variables

Angus

Simmental

Charolais

Limousin

SEM

P-values

Breed

Senes

BreedxSenes

Intramuscularfat(%)

3.23

3.25

3.25

3.27

0.178

0.999

0.001

0.972

Visible

fat(%ofLD)

CV2ofvisible

fat(%)

Average

fatpa

rtic

lesize(mm2)

Fatpa

rticle

dens

ity(n/cm2)

Three

largest

fatpa

rtic

les(%ofvisible

fat)

Numberoffa

tpa

rtic

les>30mm2

8.0

7.1

7.1

7.2

0.47

0.437

0.051

0.192

34.4

32.0

29.5

28.9

2.74

0.483

0.484

0.153

1.7

1.5

1.7

1.5

0.10

0.216

0.864

0.019

4.6

4.8

4.4

5.0

0.27

0.528

0.039

0.548

31.2

27.2

22.9

25.9

2.32

0.100

0.006

0.064

2.6

2.4

2.8

3.7

0.42

0.176

0.776

0.576

1Leastsquaremeanswithinthesamerowlacking

acommon

lett

eraresi

gnif

ican

tlydifferent(Tukey,P<

0.05).

2Coefficientofvanationbetweenquarters

oftheLD

cut.

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Table3

Meat

quality

traitsm

theM

longissimusdorsiofsteersofdifferentbreedssl

augh

tere

datasimilarintramuscular

fatco

nten

t1

Variables

Angus

Simmental

Charolais

Limousin

SEM

P-values

Breed

Senes

BreedxSeries

pHih

pH3h

pH48h

Temperaturen,(°C)

Temperature3h(°C)

Drip

loss

48h

I

Cooking

loss14li(%)

Lightness

na(L

*)Redn

essi

4d(a

*)

Yellownessi4d(b*)

Heme

iron«,,(mg/kg)

Warner-Bratzlershearfo

rce1

4d(N

)Sarcomere

lengthigh(imi)

CV

ofsarcomerele

ngth

(%)3

Prop

orti

onofsarcomere=16am(%)

Prop

orti

onofsarcomere>2

3|im(%)

Myof

ibri

llar

frag

mentationin

dex4

8h

Myof

ibri

llar

frag

mentationmdex14d

Coll

agen

48h(mg/100

g)

Coll

agen

solu

biht

y48h

(%)

659

661

659

656

0031

622a

607ab

613ab

602b

0040

554ab

557a

551ab

550b

0018

385

379

379

389

031

314b

313b

323b

344a

040

25c

30bc

36b

45a

021

206a

171b

158bc

14

1c

071

400a

373b

395a

38

lab

054

142

143

142

147

025

43

41

47

49

030

117b

140a

121b

127ab

0050

29

33

32

29

14

185a

178b

177b

176b

0019

67

77

85

88

057

20b

60ab

9lab

122a

188

18

08

15

28

107

110a

88b

107a

Ilia

41

143

131

125

129

52

549a

536ab

525ab

482b

15

5

343a

316ab

296ab

286b

136

0691

0019

0006

0740

0047

0154

0094

0575

0000

0005

0000

0323

0000

0001

0001

0888

0327

0992

0250

0246

0011

0235

0181

0122

0005

0851

0060

0725

0002

0989

0597

0745

0001

0016

0098

0486

0020

0060

0024

0000

0947

0571

0033

0311

0536

0052

0476

0224

0358

0113

0080

0267

0954

0913

0393

0680

0023

0073

0713

0332

Leastsquaremeanswithinthesamerowla

ckin

gacommon

lett

eraresignificantlydifferent(T

ukey

,P<005

)Indicesrepresentthelength

oftheperiod

post-mortem

Coefficientofvariation

(n=

120)

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WBSF was not significantly different between breed groups (Table 3). However, significant

positional differences within LD were found (Fig. 1), with lower shear force in the medial

positions (28 N on average for the dorsal and ventral position) compared to the lateral

positions (34 N). The average sarcomere length was significantly greatest in the Angus meat,

which also had the smallest proportion of sarcomeres equal to or shorter than 1.6 /im

(significant against Limousin). The proportion of sarcomeres exceeding 2.3 /xm and the

variation in sarcomere length did not differ significantly. The MFI was initially (2 d p.m.)

lowest in the Simmental steers compared to all other groups, but differences were no longer

significant after ageing for further 12 days. Shear forces although not significantly different

between groups followed a trend inverse to the MFI at 2 d p.m., i.e. the highest shear forces

conesponded to the lowest MFI values. Content and solubility of collagen were significantly

higher in the Angus compared to the Limousin steers, with the Simmental and Charolais

groups taking intermediate positions. Only few meat quality traits were affected by series of

fattening and only in pH4gh and MFLjgh significant breed group x series interactions occurred.

The meat of the Angus and the Limousin steers was judged significantly more tender than that

of the Simmental steers (Table 4); the Charolais meat was scored intermediate. Conelations

between panel tenderness and various other traits are listed in Table 5. The correlations

calculated within breeds differed to some extent. However, in those variables, which were

significant in the complete dataset, also correlations within breeds mostly were oriented

towards the same direction. In detail, negative conelations of tenderness scores with WBSF

(except Angus) and positive conelations with MFI, inespective of the time p.m. measured

were found. Although pH3h measurements ranged from 5.56 to 6.47 and temperature at 3 h

p.m. from 27.4 to 36.6 °C no significant linear or curvilinear relationship was found among

these variables and both WBSF and tenderness scores within or over all breed groups.

Significant correlations between collagen related traits and tenderness were only observed

within the Charolais group. It has to be mentioned that the variation of slaughter age was

larger in this group with 20.2 % compared to the Limousin, Simmental and Angus with

coefficients of variation of 10.4, 10.2 and 5.3 % respectively. Breed groups did not

significantly differ in flavour scores, but differences between breed groups were found in

juiciness, with the meat of the Limousin and, to a lower extent, of the Charolais steers being

juicier than that of the Simmental and particularly the Angus steers. Juiciness scores were

negatively conelated with cooking losses (r = -0.75, P < 0.001). There was a tendency in the

order of preference (P < 0.06) in the order of Limousin, Charolais, Angus and Simmental. In

the sensory evaluation no breed x series interactions occuned.

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Table4

Resultsofthesensoryevaluationof14dag

edM.

longissimusdorsiobtainedfromsteersofdifferentbreedsslaughte

red

atasimilarintramuscular

fatcontent1'2

Vanables

Angus

Simmental

Charolais

Limousin

SEM

P-values

Breed

Senes

BreedxSenes

Tenderness

4.80a

3.98b

4.59ab

4.77a

0.178

Flavourintensit

y4.45

4.11

4.35

4.43

0.129

Juiciness

3.62c

3.85bc

4.55ab

4.68a

0.208

Preference

4.61

4.36

4.84

4.95

0.157

Means

withinthesamerow

lack

ing

acommon

letter

aresi

gnif

ican

tlydifferent(T

ukey

,P<

0.05

).2

Sensoryattributeswerescoredusingthefo

llow

ingscalesfortenderness,flavorin

tens

ity,

juicinessandpreference:1=veryto

ugh,

slight,dr

yandmuch

disliked;

8=

very

0.008

0.053

0.472

0.237

0.959

0.706

0.001

0.344

0.850

0.056

0.035

0.744

tend

er,strong,juicyandmuch

liked.

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66

Table 5

Conelations between selected traits and panel tenderness scores in steers of different breeds slaughtered at a

similar intramuscular fat content1

Panel tenderness score

Angus Simmental Charolais Limousin All data

Observations 16 16 16 16 64

Age (d) 0.07 0.01 0.24 -0.11 0.03

Hot carcass weight (kg) 0.28 -0.32 0.28 -0.23 0.05

Fat thickness 12/13th nb (mm) -0.57* -0.22 0.01 0.45 0.04

pH3h -0.39 -0.38 0.26 -0.42 -0.11

Temperature31l (°C) 0.05 0.03 -0.21 -0.25 0.04

Cooking loss14d (%) 0.10 -0.23 -0.05 -0.13 -0.06

WBSF214d (kg) 0.28 -0.53* -0.53* -0.27 -0.43***

Sarcomere length48h (pm) -0.22 0.09 0.16 0.29 0.15

MFI348h 0.26 0 34 0.31 0.19 0.42***

MFI314d 0.59* 0.61* 0.40 0.05 0 4o***

Collagen«,, (mg/100 g) -0.03 0.27 -0.57* -0.09 -0.16

Collagen solubihty48h (%) -0.32 0.31 -0.57* -0 15 -0 19

1*P< 0.05, *** P< 0 001

2Warner-Bratzler shear force

3

Myofibnllar fragmentation index.

7.4. Discussion

In the present study, steers of four common beef breeds, greatly varying in development

of maturity, were fattened under identical conditions until a similar IMF content was

reached. This procedure resulted in major differences between breed groups in age at

slaughter and carcass size. Together with other genetic differences between breeds, this

was likely to result in major differences also in meat quality and, possibly, in the

distribution of visible IMF in the LD, both of which may be decisive for the purchase

decision of the consumers. The chosen level of IMF of approximately 3.25 %

conesponds to 'slight degree of marbling', a grade, which was found to be prefened in

a study on visual quality and degree of marbling, involving US consumers (Killinger et

al., 2000). Even in pork, a certain consumer segment (21 %) obviously prefered pork

with that level of IMF content although most US consumers refused this degree of

marbling in pork (Brewer et al., 2001). In beef, 47 % Swiss consumers also were found

to prefer IMF contents of 3 to 4 %, whereas 27 % selected beef without any visible

marbling (Chambaz et al., 2001). However, in Europe, carcasses like those investigated

need a specific marketing since the common grading would classify them almost

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67

without exception as excessively fat and therefore of low value in the prizing system.

As can be seen from the similarly thick subcutaneous fat layer, this restriction cannot be

overcome by the use of a certain breed.

7.4.1. Marblingproperties

A high degree of marbling is often associated with a good meat quality in the trade and

by the consumers and can therefore play an important role for purchase decision and

price. The actual sensory impression during consumption of the meat, however, seems

to be rather independent from the marbling score. This was reported as a result from

surveys on meat quality, periodically carried out in the USA (Brooks et al., 2000;

Morgan et al., 1991). Nevertheless, marbling is still an important component of e.g. the

US quality grading system and is also considered in the regulations of some US branded

beef programs. These programs not only demand a minimum extent but also a fine

texture of marbling (e.g., AAA, 2001), obviously taking into account a potentially

negative perception when the marbling is too coarse. However, none of the breed

groups investigated in the present study was clearly superior in the traits describing

marbling characteristics. Nevertheless, within LD there were positional differences in

both visible fat proportion and shear force, which appeared to be inversely related (Fig.

1). The latter might reflect a partial substitution of proteinous structures by fat, however

as the relation was not really linear, this also could have been an artefact.

Since the four breed groups were equal in carcass fatness, IMF and marbling it was

possible to compare the sensory and apparatively measured meat quality of the different

breed groups almost independently from the fat-related properties ofthe meat.

7.4.2. Meat colour

Meat colour is a further important determinant of the visual appearance of meat.

Carpenter et al. (2001) showed that consumer preferences for beef colour influenced the

likelihood of purchase but - similarly to marbling - colour did not conespond with

differences in eating satisfaction. As expected, in the present study lightness was

inversely conelated to heme iron content, the effective part of the pigments of bovine

meat. Heme iron content of muscle increases with animal age especially up to 24

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months of age and then remains quite stable (Renene, 1982). This might explain the low

heme iron content of the much younger Angus but not the differences between the other

groups. Thus, the high heme iron content of Simmental beef may be attributed to a

genuine breed characteristic, either resulting from a more rapid increase or a generally

higher total iron content of the muscle.

7.4.3. Meat texture

Tenderness is one of the most important criteria for beef quality and it has been shown

experimentally that a lot of consumers are ready to pay a higher price once they can be

sure that the beef is more tender (Dransfield, 1998). The sensory scoring revealed that

Angus and Limousin beef was most tender, followed by Charolais and Simmental beef.

Although not significantly different between breed groups, the WBSF followed the

same trend and was significantly, however not closely, conelated with sensory

tenderness. Since meat was obtained from steers and was aged for 14 d, the tenderness

level was generally high and the variation in tenderness was relatively low. This

probably contributed to the rather loose relationship between tenderness score and

WBSF. The differences in tenderness between breed groups were basically in agreement

with other studies, which however were not carried out under the precondition of a

similar IMF content: Simmental beef was found to be less tender than that of their

crosses with Angus (Dufey, 1988). Also crossbreeding with Red Holstein (Dufey, 1987)

and Charolais (Branscheid and Herzog, 1996) improved the tenderness of Simmental

beef. When comparing animals of the same age, Angus crosses with Charolais were

found to yield the same tenderness as purebred Angus while crosses with Simmental

and Limousin had less tender meat (Koch et al., 1976). In some contrast, no differences

in tenderness were found between Simmental and Angus crossbred steers when

slaughtered at an equal backfat thickness (Laborde et al., 2001).

Various factors may be responsible for the breed group differences in tenderness found

in the present study. Early post-mortem pH has been proposed as one factor affecting

meat tenderness from the myofibrillar side as it influences the activity of endogenous

enzyme systems (O'Halloran et al., 1997). From several investigations (French et al.,

2000; Marsh et al., 1987; Pike et al., 1993) it appears that glycolysis and the resulting

meat tenderness is highest at a pHßh of 6.0 to 6.1. This can be either achieved by

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69

electrical stimulation of carcasses (Marsh et al., 1987) or producing heavy carcasses

with a conespondingly slow temperature decline (Pike et al., 1993) as was the case with

the Limousin group in the present study. However, ageing of the meat for 14 d

decreased initial tenderness differences in other studies (O'Halloran et al., 1997; French

et al, 2000) and can also be seen from the development of MFI in the present

investigation. Accordingly no significant relationship between pH3h and sensory

tenderness of meat aged for 14 d was found in our and also other studies (Shackelford et

al., 1994). Both Limousin and Simmental beef showed an ideal pH3h, but Simmental

beefwas graded significantly lower in tenderness.

Another important factor of the myofibrillar compound of tenderness could have been

sarcomere length, particularly when pH3h is greater than 6.3 (Smulders et al., 1990),

although not always a clear relationship between sarcomere length and tenderness is

found (O'Halloran et al., 1997). One precondition for a close conelation seems to be the

occunence of cold or heat shortening (Shorthose and Harris, 1991). Some contraction

obviously occuned in Limousin and, to a smaller extent, in Charolais and Simmental

probably not due to cold but heat shortening. Lochner et al. (1980) have shown that

carcass size and fatness have a great influence on cooling rate and can be at least as

important in determining muscle cooling rate as the temperature and velocity of the

chiller air. Moreover it is unlikely that cold shortening occuned in Limousin because

they had the lowest pH and the highest muscle temperature at 3 h p.m. and so should

have the lowest risk of cold shortened muscle from the four breed groups. Lee and

Ashmore (1985) observed contracted mean sarcomeres length (1.66 pm) and increased

toughness due to heat shortening in carcasses with similar backfat thickness and

temperature at 3 h p.m. although carcasses were about 100 kg lighter as in the Limousin

group of our study. In cold-shortened fibres often both contracted and stretched

sarcomeres are present resulting in a greater variation of sarcomere length (Locker et

al., 1975). This was not the case in this study and it can be presumed that the 14 d-

ageing period was sufficient to offset or overcome the effect of shortening as Limousin

panel tenderness scores were as good as that of the Angus which had almost no

sarcomeres contracted to less than 1.6 /im. Accordingly, the weak relationship between

the sarcomere properties investigated and tenderness scores as well as shear force

remained unclear in terms of both breed group differences and conelations.

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In the present study MFI turned out to be a suitable indicator for tenderness and breed

group differences in tenderness of the aged LD, which was also observed in other

studies (Culler et al., 1978; Vestergaard et al., 2000). MFI particularly reflects changes

occurring during ageing of meat. In the present experiment, MFI was measured at 48 h

and at 14 d p.m. In this period MFI increased by 30, 49, 17 and 16 % in the Angus,

Simmental, Charolais and Limousin groups, respectively, thus decreasing the initial

variation among breeds (particularly Simmental vs others). Therefore it can be assumed

that Simmental beef would have been even clearer inferior in tenderness against all

other groups without or with a shorter period of ageing.

The connective tissue-related traits were described by collagen content and collagen

solubility. As expected, steers from breeds which were slaughtered at a higher age

expressed a lower collagen solubility reflecting the increasing formation of mature or

heat-stable crosslinks (Purslow, 1994). In the Limousin group, this was obviously

compensated by a lower collagen content which actually could reflect a kind of dilution

of connective tissue by other muscle tissues. The only significant conelations between

collagen related traits and panel tenderness were found in the Charolais group, which

also showed the greatest variation in age. Nevertheless the negative relationship

between collagen solubility and tenderness in the Charolais is contradictory to the

expectations.

7.4.4. Flavour, juiciness and water-holding capacity ofmeat

Studies comparing beef breeds in meat flavour mostly reported no substantial

differences (Koch et al., 1976, 1979, 1982; Wheeler et al., 1996, 2001). In contrast,

Laborde et al. (2001) noted higher flavour scores in Simmental crossbred steers than in

Angus steers slaughtered at a similar backfat thickness, but the Simmental crossbreds

were 73 d older and their beef had a 31 % higher IMF content. Since in the present

study clear breed group differences in slaughter age also occuned and intensity of

flavour is known to develop with increasing animal age (Lawrie, 1991), differences in

flavour were expected but not found.

In contrast, the present study revealed clear breed group differences in water-holding

capacity and juiciness of the meat, traits which closely conelated. It seems that, due to

the great differences in age at slaughter which represent indirect effects of breed

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through the breed-specific rate of IMF accretion, breed group differences in juiciness

were far more pronounced than in other studies comparing beef not at the same IMF

content but at the same age or weight of the animals (e.g., Crouse et al., 1985). This

age-dependance may be at least partly a result of the decline in muscle water content

with age, shown to be closely associated with cooking loss in Simmental cattle by

Lüdden (1991). Another factor pronouncing beef group differences in cooking loss

could have been the differences in size of the LD slices (small in Angus). However, no

direct relationship between surface, weight and surface to weight ratio of the LD slice

and the level of cooking loss was found. Ranking in drip loss was opposite to that in

cooking loss and juiciness. Drip loss stresses other compartments of bound water than

cooking loss and therefore does not conelate with cooking loss (Honikel, 1986). In the

present study, overall group differences in cooking loss were of far greater importance

than those in drip loss.

7.4.5. Overall sensory preference ofmeat

Panelists tended (P < 0.06) to prefer the LD of Limousin and Charolais steers. This can

be explained by the lower juiciness of the Angus beef and, additionally, the lower

tenderness of the Simmental beef. Generally, the results of other studies, although not

carried out under the precondition of a similar IMF content, indicate that differences in

meat palatability are small between Bos taurus breed groups (Koch et al., 1979; Monin

and Ouali, 1991) and not very consistent due to the large variability within relative to

that between breeds (Wheeler et al., 1996).

7.5. Conclusions

The present study illustrated that beef from different breeds may differ in quality, when

the animals are fattened under the same conditions until they reach a similar IMF

content, particularly when early and late maturing animals are compared. These

differences might be less pronounced when age differences at slaughter would be

reduced by a variation of the feeding intensity. However, semi-intensive feeding

systems could be very competitive by reducing feeding costs particularly in grassland

regions. Under these conditions, Charolais and Limousin could have a certain advantage

which is expressed in favourable juiciness and tenderness, low cooking losses and

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certain cost-effective carcass traits. On the other hand, these breeds will provide very

heavy carcasses and an unusually long fattening period is required when an IMF content

of more than 3 % is demanded. For these reasons intensive fattening systems would be

advisable. Angus, with also very tender beef but inferior water-holding capacity, would

have the advantage of a much shorter fattening period at simultaneously higher average

daily gains providing the best choice for extensive systems. Simmental beef was ranked

inferior; therefore, crossbreeding might improve the Simmental offspring's beef quality.

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8. General discussion

The choice of an intramuscular fat content of approximately 3.5 % as slaughter criterion

was higher than the one commonly found in beef cattle in Switzerland as this ranges

between 1 and 2.5 % (Dufey and Chambaz, 1999b). The determination of this content in

living animals is not easy. The ultrasound technology permits to select animals for

constituting fattening groups conesponding to distinct markets but this technology is

still in development and therefore too preliminary to determine accurately enough the

slaughter end point when determined at a fixed level or a nanow range of intramuscular

fat.

The condition of a similar intramuscular fat content as slaughter criterion had a great

influence on growth characteristics of the different breed groups of this study, as well as

the use of steers instead of bulls and beef breeds instead of crossbreds with dairy cattle.

Since these last few years there is a tendency of fattening pure beef breeds and supply is

cunently lower than demand. These type of cattle convey a positive image at the same

time to the consumers, butchers and producers also. However most of the pure breeds

used in this study, Angus excepted, are later maturing than animals in conventional

fattening coming from dairy husbandry. Fattening duration, again Angus excepted, was

therefore longer than usual in conventional fattening with, in increasing order of

duration, the Simmental and Charolais, followed by the Limousin and lastly the Blonde

d'Aquitaine and Piedmontese. The slaughter order reflected the differences in maturity

rate of the different breeds. These results are in agreement with those found in the

literature and allowed to refute the widespread opinion that Limousin is an early

maturing breed. Using late-maturing cattle and slaughtering the animals at 3.5%

intramuscular fat prolonged the fattening duration which raised considerably the live

slaughter weights and the feeding duration and therefore feeding costs. Growth

performance in the conditions of this study was generally better for the earlier maturing

breeds or respectively for the youngest animals.

The finding concerning carcass quality was inverse with the later maturing animals

presenting the best carcass quality if the problems of high carcass weight are excluded.

The high dressing percentages added to the high slaughter weights in the later maturing

animals indeed increased the difficulty or made it impossible not to exceed the limits of

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carcass weights in force in Switzerland. It means that on one side the animals of the

later maturing breeds such as Blonde d'Aquitaine and Piedmontese should be fattened

longer to obtain a sufficient fat cover and on the other side they should be slaughtered

earlier than the earlier maturing breeds such as Angus to avoid that carcass weight limits

are surpassed. For example in our study no single animal succeeded to obtain

simultaneously 3.5% intramuscular fat content, optimal carcass fatness (conesponding

to the score 3 ofthe CH-TAX) and a carcass weight below 350 kg. It is concluded that it

is very difficult to reconcile the interest of the different contributors of the meat market

(producers, butchers and consumers) in any case in the conventional market.

Piedmontese meat is often vaunted for its low cholesterol content. It is probably due to a

subjective association with the low fat content of this later maturing breed. However the

present results as well as those of the literature showed that cholesterol content was not

conelated with fat content firstly and secondly that Piedmontese had not less cholesterol

than the others breeds.

Meat quality obtained in this study was judged good to very good from the panelists

which confirms that this type of production could meet the consumers' expectations

regarding sensory quality of labeled meat. In spite of a similar intramuscular fat content

important differences in tenderness, juiciness and flavor were observed between breed

groups (see appendix). Intramuscular fat is therefore not the basic cause for the

differences observed. Piedmontese meat was finally prefened to Simmental meat

whereas the four other breed groups were not different in preference.

In conclusion, no breed group of this study excels in all economically important traits.

The Angus for example were the most rapid to reach 3.5% intramuscular fat with the

least feeding costs. On the other hand carcass quality was the lowest of the six breed

groups of this study whereas sensory quality was judged good. On the other extreme,

Piedmontese had the worst fattening performance at an excellent carcass and meat

quality in accordance with results of the literature. The fattening of steers of this breed

is certainly economically not profitable for the producers who sells animals through the

cunent conventional Swiss market. Namely Piedmontese would need a feedlot-type

ration to reach a sufficient carcass fatness in reasonable fattening duration as well as

carcass weight conesponding to the cunent conventional Swiss market. The still high

cost of concentrate in Switzerland as well as the limited added value through the CH-

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TAX of the very good lean to bone ratio are unfavorable. However, as part of the

marketing through direct selling for example, the producer could compensate the higher

fattening costs by direct benefit from the higher lean yield of the carcass and the high

sensory quality. This last aspect is essential in direct selling knowing that consumers are

ready to pay more for this type of meat but their expectations are also higher. The ideal

breed combining all qualities in growth performance, carcass and sensory quality does

not exist but choice of one breed instead of another one must be determined by the

characteristics specific to the farm, i.e., quantity, type and cost of feed available as well

as economical environment and type of marketing. It remains open whether

crossbreeding of dairy dams with beef cattle sires, which is at present more usual in

Switzerland as the fattening of pure beef breeds, would have better reconciled the

complete range of objectives followed in this study.

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10.Appendix

Resultsofthesensoryevaluationof14dagedM

longissimusdorsi

obtainedfrom

steersofdifferentbreedsslaughtered

atasimilarintramuscular

fatcontent

Variables

Angus

Simmental

Charolais

Limousin

Blonde

d'Aquitaine

Piedmontese

SEM

P-values

Breed

Series

BreedxSeries

n16

16

16

16

16

16

Tenderness

3

Series

1513a

3.95c

488ab

4.94ab

4.15bc

4.18bc

0.314

0.046

Series2

448b

4.02b

430b

4.60b

4.42b

5.61a

0.232

0.001

Flavourintensity

445ab

4.11b

434ab

4.43ab

4.59ab

4.67a

0.120

0.027

0.494

0.095

Juiciness

362c

3.85bc

455abc

4.68ab

4.70ab

4.88a

0.219

0.000

0.059

0907

Preference

4.6lab

436b

484ab

495ab

4.61ab

5.22b

0.165

0.008

0.264

0390

1Meanswithm

thesamerow

lacking

acommon

letter

aresi

gnif

ican

tlydifferent(Tukey,

P<

0.05).

2

Sensory

attributeswerescoredusingthefollowingscalesforte

nder

ness

,flavorintensity,

juicinessandpr

efer

ence-1=verytough,

slight,dr

yandmuch

disliked,

8=

very

tend

er,strong,juicyandmuch

liked

3

Separa

tepresentationoftendernessscores

forseries

1and2becauseofasi

gnif

ican

tinteractionbetweenbreedand

series.

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11. Remerciements

J'aimerais remercier en tout premier lieu Piene-Alain Dufey, l'instigateur de ce projet

qui a suscité beaucoup d'intérêt dans les milieux agricoles et de la boucherie. Piene-

Alain a été à la fois un conseiller et un ami. Le travail à ses côtés restera pour moi le

meilleur souvenir de la thèse.

Je suis reconnaissant au Professeur Michael Kreuzer pour la lecture attentive et très

rapide des manuscrits ainsi qu'au Dr. Martin Scheeder pour les discussions à chaque fois

passionnées et stimulantes.

Merci au Prof. Nikiaus Künzi pour avoir accepté de fonctionner en tant que

corapporteur.

Je suis particulièrement reconnaissant à la station fédérale de recherches en production

animale à Posieux (RAP) et à sa directrice Mme Danielle Gagnaux de m'avoir donné

l'occasion de faire ce doctorat dans d'aussi bonnes conditions. Merci spécialement à

Isabelle Morel pour la planification et la conduite de la partie engraissement des

animaux et pour avoir participé comme coauteur à la rédaction d'une des publications de

la thèse. Merci à l'équipe du laboratoire des viandes pour les nombreuses analyses

effectuées dans le cadre de ce projet. La compétence et la disponibilité de Claudine

Biolley et Giuseppina Bächler furent très précieuses, merci aussi à Aude Auriou,

Ghislaine Kutnar, Jessica Messadène, Georges Guex, François Buchs et Denis Robatel.

Un merci particulier à Yves Arrigo, "Monsieur système D", pour pratiquement tous mes

problèmes informatiques et à Edi Lehmann pour sa gentillesse, sa compétence et sa

patience légendaire.

J'aimerais remercier pour leurs conseils précieux et la collaboration agréable André

Chassot tout d'abord, Monique Delacombaz, Gerhard Mangold, Peter Stoll, Josef

Sturny, Roger Daccord, Andreas Gutzwiller et Jean-Yves Deru. La période passée à la

RAP m'a permis aussi de nouer des contacts chaleureux non seulement avec les

personnes ayant directement collaboré au projet mais aussi avec d'autres collègues que

je profite de saluer ici: Bernard Papaux, Claude Chaubert, Franz Jans, Marco Meisser,

Roland Cotting, Walter Stoll, Giuseppe Bee, Ueli Wyss, Andreas Münger, Fredy

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Castella, Marcel Barman, Jean-Louis Gafner, René Vogel, Maria Rodrigues, Léon

Grand, Juliette Sciboz, Guy Maikoff et Hans Schnyder.

Pour terminer les remerciements aux collègues de la station, j'aimerais encore féliciter

l'équipe de l'atelier, Jean-Piene Mettraux, Jean-François Bise, Robert Chassot et Peter

Schäfer, pour leur ingéniosité et remercier les responsables des animaux, Michel Folly

et Gérard Brodard en particulier.

Je tiens à remercier le Prof. Jeny Gresham de l'université de Martin au Tennessee chez

qui je suis allé apprendre la technique ultrasons de détermination in vivo de la graisse

intramusculaire. Il a su par sa motivation me redonner l'énergie nécessaire quand les

résultats obtenus ne conespondaient pas à mes attentes. Je le remercie aussi vivement de

sa contribution active dans la rédaction du papier concernant la technique aux ultrasons.

Merci à la maison SUTER SA et à son directeur Ueli Gerber pour avoir accepté de

participer à ce projet de recherche et surtout pour avoir persévéré malgré la taille et l'état

d'engraissement non conventionnels des carcasses de cet essai. Merci à Hans-Ruedi et

Peter Gerber, Kurt Zenger, Joël Suter, Yvan Mercanton, Max Knecht, Stefan Seiler,

François Caula et Radosavljevic Ratomir pour l'excellente collaboration, leur flexibilité

et les bons moments passés ensemble.

Je remercie Piene Berchier pour les conseils statistiques de même qu'André Chassot.

Mes remerciements vont pour terminer à l'ASVNM (Association suisse des détenteurs

de vache nourrices et mères) et à PROVIANDE pour leur intérêt dans le projet et leur

contribution financière.

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12. Curriculum Vitae

Name:

Born:

Citizen of:

Alain Chambaz

3 March 1971

Arzier, VD

1977-1983

1983-1986

1986-1989

1989

Primary school in Vevey

Secondary school in Vevey

High school in La Tour-de-Peilz (CESSEV)

High school graduation (matura Type B)

1989-1991 Farm apprenticeship at Hans Dumermuth, Belp, Bern and

at Jean-Claude Joliquin, Combremont-le-Grand, Vaud

Agricultural school at Marcelin-sur-Morges

1991-1996 Study of Agricultural Science at the Swiss Federal Institute of

Technology, ETH Zurich, Departement of Agriculture and Food

Science including traineeship in human nutrition at the University

of Lausanne, Faculty of Medicine

1996 Graduation as Dipl. Ing.-Agr. ETH in Animal Sciences

1996-1997 Research assistant in human nutrition at the Nestle Research Center

in Vers-chez-les-Blancs, Lausanne

1997-2001 Doctoral Student at the Swiss Federal Institute of Technology

(ETH) and the Swiss Federal Research Station for Animal

Production, Posieux (RAP)


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