Available online at www.sciencedirect.com

4 (2008) 19–30www.elsevier.com/locate/livsci

Livestock Science 11

Live weight, body size and carcass characteristics of youngbulls of fifteen European breeds

P. Albertí a,⁎, B. Panea b, C. Sañudo b, J.L. Olleta b, G. Ripoll a, P. Ertbjerg c,M. Christensen c, S. Gigli d, S. Failla d, S. Concetti d, J.F. Hocquette e, R. Jailler e,

S. Rudel e, G. Renand f, G.R. Nute g, R.I. Richardson g, J.L. Williams h

a Centro de Investigación y Tecnología Agroalimentaria, Gobierno de Aragón, 50080, Zaragoza, Spainb Departamento de Producción Animal y Ciencia de los Alimentos, Universidad de Zaragoza, 50013, Zaragoza, Spain

c Department of Food Science, University of Copenhagen, DK-1958 Frederiksberg C., Denmarkd CRA, Istituto Sperimentale per la Zootecnia, 00016 Monterotondo, Italy

e INRA, UR1213, Unité de Recherches sur les Herbivores, Centre de Clermont-Ferrand./Theix F-63122, Francef INRA, UR337, Station de Génétique Quantitative et Appliquée, F-78352 Jouy-en-Josas cedex, France

g Division of Farm Animal Science, University of Bristol, BS40 5DU, United Kingdomh Parco Tecnologico Padano, Via Einstein, Polo Universitario, Lodi, 26900, Italy

Received 16 May 2006; received in revised form 3 April 2007; accepted 6 April 2007

Abstract

A total of 436 young bulls from fifteen Western European breeds, including beef, dairy and local types from five countries, werestudied to assess variability in live weight, live weight gain, body measurements and carcass traits. Animals were logged indoors,and fed a diet based on concentrate and straw offered ad libitum from 9 months of age to slaughter at 15 months of age. The weight,body length, height at withers and pelvis width, of the animals were recorded at 9, 12 and 15 months of age. After slaughter, 15carcass variables were recorded, including carcass weight, EU classification scores, morphological measurements and dissectiondata. Data were analysed by GLM, regression and principal component analysis procedures.

Significant differences were found between breeds for all variables studied, however, the body size measurements and thecarcass traits were more useful to discriminate among cattle breeds, than either live weight or daily gain. With respect to the bodysize and carcass traits the studied breeds could be grouped as:

– Specialized beef breeds, comprising Piemontese, Asturiana de los Valles, Pirenaica, Limousin, South Devon, Charolais andAberdeen Angus, all of which were characterized by high muscularity, wide pelvis and medium height and a low to medium levelof fatness.

– Local and dairy breeds, comprising Jersey, Casina, Highland, Holstein and Danish Red, the latter two breeds were tall animals,while the former three breeds were small in size. In general the group was poorly muscled and tended to have a high or mediumlevel of fat.

– Intermediate group, Avileña, Marchigiana and Simmental: these breeds were characterized by an intermediate muscleconformation and fatness level and were relatively tall.

⁎ Corresponding author. Tel.: +34 976 716438; fax: +34 976 716335.E-mail address: palberti@aragon.es (P. Albertí).

1871-1413/$ - see front matter © 2007 Elsevier B.V. All rights reserved.doi:10.1016/j.livsci.2007.04.010

20 P. Albertí et al. / Livestock Science 114 (2008) 19–30

This study provides a detailed assessment or a wide range of variables in the major breeds, and several minor breeds, that areused in breeding programmes across Europe and elsewhere, and will provide information that will be of use to define breedingstrategies to meet the demands of the European beef market.© 2007 Elsevier B.V. All rights reserved.

Keywords: Beef cattle; Phenotypic relationships; Carcass morphology; Longissimus thoracis measures; Carcass traits

1. Introduction

A large number of breeds of cattle exist in WesternEurope many of which are well known and some arewide spread. These breeds have been a great source ofgenetic diversity and have been used for crossbreedingthroughout the world. Rigorous selection of specializedbreeds has produced extreme dairy and beef breeds thatare widely used, while many local and hardy breeds thatexist now have decreasing numbers. It is likely that thisgenetic diversity could produce meat with manydifferent qualities; however, this has not been studiedin detail.

It is well known that the optimal slaughter ages andweights vary widely among cattle breed types (Longet al., 1979) depending on how rapidly they mature,which is characterized by laying down of fat during a“finishing” period. In general, animals that producelonger carcasses tend to have a poorer conformation anda higher level of fat in their carcass (Piedrafita et al.,2003), however, for some small breeds this observationis not always the case. Variations in the yield of carcasscomponents are related to the genetic composition andbody size differences, which are inherent breed differ-ences (Jenkins et al., 1981). Albertí et al. (2005)suggested that diverse beef breeds could be consideredin three broad groups: high meat producing animals, thatcorrespond to late maturing animals with short carcasssize and high “blockiness”; medium meat producingbreeds, with intermediate characteristics, and low meatproducing breeds, that correspond to early maturinganimals with large carcass size and low “blockiness”.Different types of breed require different productionsystems and the combination of breed and productionsystem result in products differentiated by diverseattributes, such as meat colour or fat content (Gil et al.,2001). These attributes of the meat are very important forthe consumer.

Current selection programmes are aimed at increas-ing adult weight and size of beef breeds. It is known thatgrowth rate is positively associated with adult bodyweight (Kempster et al., 1982) and that animals with ahigh weight gain produce more muscle fibres withgreater glycolytic activity, which is likely to favour meat

ageing and hence tenderness (Jurie et al., 1995), whichis a major factor in the meat preference of consumers.Selective breeding in cattle has been very successful inincreasing the quantity of beef production, but up to nowthere has been little attention paid to product quality. Ifthe genes controlling various aspects of meat quality areidentified, they could be included in selection pro-grammes to enhance meat quality that could, potentially,take into account regional variations in consumerpreferences. Hence, body size and carcass compositionshould be considered as well as beef quality traits, inorder to take the biological relationships between theseparameters into consideration. Understanding the rela-tionship between these different traits is being facilitatedby the recent rapid increase in information available onthe bovine genome and the tools to study geneexpression (Bouley et al., 2005; Sudre et al., 2005).Several studies to characterize production and carcasstraits of European beef breeds have been reported(Piedrafita et al., 2003; Chambaz et al., 2003; Keane,2003; Özlütürk et al., 2004; Albertí et al., 2005;Cuvelier et al., 2006), however, most of these examineda small number of breeds and so are not representative ofthe European cattle breeds.

The study presented here defines the variation in liveanimal, carcass and meat characteristics in 15 Europeanbreeds of cattle reared under comparable managementconditions, and so represents one of the most compre-hensive comparisons of variation among Europeanbreeds.

2. Material and methods

2.1. Animals

A total of 436 pure-breed bulls from the fifteenbreeds were reared in five experimental researchcentres: breeds were from the United Kingdom, 31Jersey, 27 South Devon, 30 Aberdeen Angus and 29Highland; from Denmark, 29 Holstein, 29 Danish RedCattle and 20 Simmental; from Spain, 30 Asturiana delos Valles, 31 Casina, 30 Avileña and 31 Pirenaica; fromItaly, 30 Piemontese and 28 Marchigiana and fromFrance, 31 Limousin and 30 Charolais.

21P. Albertí et al. / Livestock Science 114 (2008) 19–30

2.2. Rearing conditions

Animals were selected to be as unrelated as possibleto ensure that the full range of genetic diversity presentwithin breeds was included in the study. The bulls werepurchased between 6 and 9 months of age on average forbeef breeds, while for dairy breeds; the bulls werepurchased as young calves (on average 7 days for Jerseyand 1 to 1.5 months for Holstein and Danish Red). Dairycalves were fed with milk substitute and rolledconcentrate until 2 months of age and then pelletedconcentrate and hay ad libitum until 9 months of age.

At 9 months of age, all the animals were transferred tothe experimental farms, where they were weighed andquarantined for 15 to 30 days to adapt to their newenvironment. After this they were divided into groups of7 to 8 animals, and put onto the standardised diet, whichconsisted of a concentrate compounded from barleyflakes (80 to 84%), soya bean meal (7.5 to 11%) sodiumbicarbonate (0.6%) with suitable vitamin supplements(1.5%) and barley straw, all fed ad libitum. The energydensity ratio ranged from 12.9ME/kg DM for most of thebreeds, to 13.5 ME/kg DM for British breeds. Theprotein content was 160 g CP/kg DM up to 10 months ofage and then decreased to 150 g CP/kg DM to slaughter.

2.3. Weight and body size measurements

National welfare and animal experimental regulationswere observed when handling the animals. Animals wereweighed at the beginning of the experimental period andat 9, 12, 15 months of age and prior to slaughter. Averagedaily gain (ADG) was calculated between the initialweight and weight at slaughter and the 365-day weightcalculated by linear regression. Body measurementswere recorded, 9, 12 and 15 months of age. These were:height at withers (measured from the highest point of theshoulder blade to the ground), pelvis width (measured atthe trochanter major ossis femoris) and body length(measured from humerus breastbone articulation to theischium tuberosity).

2.4. Carcass evaluation

At75%mature bullweight,whichwasabout 15monthsof age, animals were slaughtered by captive bolt pistoland exsanguination in either commercial or experimentalslaughterhouses, depending on the experimental facilitiesof each country. Carcass dressing followed a standardisedproject protocol, without use of electrical stimulation, andwith the removal of the remaining subcutaneous fat coverand testicles. Carcasses were split into two sides with tail

on the right side of the carcass and chilled at 4±1 °C for24 h. Temperature in the centre ofM. longissimus thoracis,at the 10th thoracic rib, was not allowed to fall below10 °Cwithin the first 10 h.

Immediately after slaughter, the following measure-ments were recorded from the left half-side of carcass:hot carcass weight; hot kidney knob and channel fatweight (KKCW); dressing percentage, calculated as (hotcarcass weight/slaughter live weight)×100; conforma-tion score was graded according to the SEUROP classi-fication (EEC, 1991) with a scale ranging from 1 (verypoor conformation) to 18 (very good conformation); fat-ness score was measured by UE classification, witha 15-point scale (1, very low fat to 15, very high fat).Morphological measures were taken according to themethodology described by De Boer et al. (1974).Variables recorded were: carcass length, from the ante-rior edge of symphysis pubis to the middle of theanterior edge of the visible part of the first rib; internaldepth of breast, at the level of the fifth rib from theventral edge of the spinal canal of the posterior aspectof the body of the fifth thoracic vertebra to ventralaspect of the middle of the body of the sixth sternebra;limb length, from the medial malleolus of the tibia ina straight line to the anterior edge of the symphysispubis. From these measurements a carcass blockinessindex was calculated. This index expresses the rela-tionship between hot carcass weight (kg) and carcasslength (cm) as follows: Blockiness index=hot carcassweight⁎100 / carcass length. High values indicate highmuscular development (Albertí et al., 2001).

The area of the M. longissimus thoracis (LT), at the5th to 6th rib level, was calculated by planimetry,recorded by tracing: an acetate sheet was placed on thecranial side of surface of the loin, and the border of themuscle marked on the sheet using glass marker pen.Medium-lateral and dorso-ventral diameters (maximumand minimum diameters, respectively) were alsomeasured (Hammond, 1936).

2.5. Tissue composition

The 6th thoracic rib joint was collected 24 h post-mortem, weighed and dissected into muscle, fat and bone,tendons and noticeable blood vessels according to themethod described by Robelin and Geay (1975). Resultsare expressed as percentage of the entire rib weight.

2.6. Statistical analysis

Differences among breeds for slaughter weight, ageat slaughter, average daily gains and carcass traits were

Table1

Age,slaug

hter

weightsandaveragedaily

gains(±ES)of

cattlebreeds

Breed

AA

ASV

AVI

CAS

CHA

HIG

HOL

JER

LIM

MAR

PIE

PIR

RED

SD

SIM

S.E.

Significance

n30

3030

3130

2929

3131

2830

3129

2720

Average

daily

gain

(kg/day)

1.97

a1.30

ef1.34

e1.12

ghi

1.53

c1.01

I1.18

gh

1.08

hi

1.46

cd1.22

fg1.22

fg1.37

de

1.09

hi

1.84

b1.49

c0.038

⁎

(±S.E.)

0.05

0.03

0.05

0.03

0.04

0.02

0.04

0.02

0.03

0.03

0.02

0.03

0.04

0.56

0.06

Slaug

hter

weigh

t(kg)

597.7b

cd55

7.7e

f55

0.9e

fg43

3.5h

634.0a

443.5h

596.3b

cd37

8.4i

565.4e

d52

3.5g

527.3fg

602.4a

bc

580.0c

de

591.7b

cd62

1.8a

b8.66

⁎

(±S.E.)

4.6

8.8

13.4

7.1

7.3

3.4

9.3

1.5

5.4

7.2

7.3

9.5

10.6

6.2

20.9

Slaug

hter

age(days)

428.6c

d46

0.6b

462.2b

461.4b

460.6b

510.6a

458.0b

414.7d

e42

8.0c

d45

9.2b

461.0b

c44

4.8b

c45

4.3b

398.5e

455.9b

5.74

⁎

(±S.E.)

8.8

5.5

6.3

4.7

3.9

8.9

1.0

6.4

4.1

3.7

3.5

5.8

2.8

9.0

2.4

AA,A

berdeenAngus;A

SV,A

sturiana

delosValles;AVI,Avileña;C

AS,

Casina;CHA,C

harolais;H

IG,H

ighland;HOL,H

olstein;JER,Jersey;LIM

,Lim

ousin;MAR,M

archigiana;P

IE,P

iemontese;

PIR,P

irenaica;RED,DanishRed;SIM

,Sim

mental;SD,South

Devon

.⁎ N

oteforallparameters,Pb0.00

1.Different

superscriptletters

inthesamerow

indicate

differencesbetweenbreeds

forat

least1%

.

22 P. Albertí et al. / Livestock Science 114 (2008) 19–30

determined by variance analysis, using the GLMprocedure, considering breed as an unique effect, andthe Duncan method at α=0.05 used to test differencesamong breeds. Weight and morphological measure-ments for body length, wither height and pelvis widthwere estimated at 365 days from the linear regression,and the Duncan method at α=0.01 used to testdifferences among breeds. For carcass traits a principalcomponents analysis procedure adjusted by mean ofquartimax rotation was performed to represent thediversity among the 15 cattle populations. All thesestatistical analyses were carried out using the SASstatistical package v. 8.02 (SAS, 1997).

3. Results

3.1. Live characteristics

The average daily gain (ADG) of young bulls from9 months of age to slaughter showed significant differ-ences among breeds (Table 1). Aberdeen Angus andSouth Devon had a high daily gain of 1.97 and 1.84 kg/day respectively. Holstein, Casina, Danish Red, Jerseyand Highland had the lowest ADG (1.1–1.0 kg/day),whereas Charolais, Simmental, Limousin, Pirenaica,Avileña, Asturiana de los Valles, Marchigiana andPiemontese, had an intermediate ADG (1.5–1.2 kg/day).

The mean slaughter age for all breeds was 450.6±39.0 days, but some breeds were slightly younger atslaughter (398–429 days): South Devon, Jersey, Limou-sin, and Aberdeen Angus, while Highland was older,with a slaughter age of 511 days. Therefore, breeds werecompared on the adjusted weights at 365 days calculatedby regression. Awithin breed linear regression was fitted,as the age scarce coefficients were not significant on thisage interval. The adjusted weights at 365 days of ageestimated from a linear regression are showed in Table 2.

The South Devon with and average weight of 510 kgwas the heaviest breed at 365 days, while theintermediate breeds included both beef and dairy breeds:Holstein, Aberdeen Angus, Danish Red, Pirenaica,Simmental, Charolais, and Limousin with an averageweight of 483 kg. The Asturiana de los Valles, Avileña,Piemontese and Marchigiana weighed between 422 kgand 395 kg. The lightest breeds were Highland, Jerseyand Casina. The Charolais was the heaviest breed atslaughter with an average weight of 634 kg at 461 daysand the lightest was the Jersey with a weight of 378 kg at415 days, but really breeds keep the same group ascommented previously at 365 days of age.

Differences between most breeds in body sizemeasurements, height at withers, pelvis width and

Table2

Liveweightandmorphological

traits,adjusted

atage365days

bylin

earregression

AA

ASV

AVI

CAS

CHA

HIG

HOL

JES

LIM

MAR

PIE

PIR

RED

SD

SIM

S.E.

Significance

n30

3030

3130

2929

3131

2830

3129

2720

Weight(kg)

483.7b

422.3c

418.9c

d32

1.4f

481.75

290.9g

483.8b

314.4f

467.0b

395.2e

399.4d

e48

3.1b

483.2b

509.7a

481.8b

5.84

⁎

(±S.E.)

7.8

5.2

6.9

3.9

5.0

5.4

4.5

4.1

3.9

4.4

4.0

5.5

5.9

9.5

11.2

With

erheight

(cm)

120.2d

119.0d

e12

2.9c

111.3f

120.6d

103.3g

133.1a

112.9f

118.7d

e12

4.7b

c118.8d

e117.6e

125.9b

119.5d

e12

6.1b

0.61

⁎

(±S.E.)

0.58

0.50

0.57

0.40

0.40

1.26

0.53

0.75

0.41

0.45

0.29

0.63

0.52

0.57

0.90

Pelviswidth

(cm)

48.6cd

49.0bc

45.4f

40.9g

52.8a

38.2h

46.7ef

38.5h

50.4b

47.9cde

49.0bc

49.2bc

46.4ef

49.3bc

47.0de

0.41

⁎

(±S.E.)

0.48

0.36

0.31

0.25

0.26

1.05

0.31

0.33

0.30

0.25

0.19

0.39

0.36

0.62

0.65

Bodylength

(cm)

142.7b

152.0a

149.1a

136.4c

d14

5.0b

121.6h

129.8e

f13

3.6d

e13

8.6c

133.9d

133.3d

e15

0.7a

128.6fg

144.1b

125.9g

1.02

⁎

(±S.E.)

0.87

0.95

1.16

0.88

0.58

1.67

0.64

0.96

0.73

0.49

0.45

1.42

0.69

0.93

1.07

AA,A

berdeenAngus;A

SV,A

sturiana

delosValles;AVI,Avileña;C

AS,

Casina;CHA,C

harolais;H

IG,H

ighland;HOL,H

olstein;JER,Jersey;LIM

,Lim

ousin;MAR,M

archigiana;P

IE,P

iemontese;

PIR,Pirenaica;RED,D

anishRed;SIM

,Sim

mental;SD,S

outh

Devon.

⁎ Noteforallparameters,Pb0.001.

Different

superscriptletters

inthesamerow

indicate

differencesbetweenbreeds

forat

least1%

.

23P. Albertí et al. / Livestock Science 114 (2008) 19–30

body length, adjusted to 365 days of age were highlysignificant for each morphological trait (see Table 2). At365 days, the pelvis width correlated with wither height(r=0.50, Pb0.001) and with body length (r=0.42,Pb0.001) while no correlation was found between bodylength and wither height. Jersey, Casina and Highlandhad a low withers height, between 113 cm and 103 cm,while Holstein had the greatest (133 cm). Simmental,Danish Red, Marchigiana and Avileña breeds rangedbetween 126 and 123 cm, whereas the others breeds hadintermediate values (120 cm to 118 cm). Three breeds,Casina, Jersey and Highland had a small pelvis widthwith values between 41 and 38 cm, Charolais had thewidest pelvis (53 cm). Highland had the shorter bodylength of 122 cm, whereas Asturiana de los Valles,Pirenaica and Avileña, had greatest body lengths (152 to149 cm).

When the relationship between breeds is exploredusing the morphological traits, wither height and pelviswidth, three main groups can be defined based on frameor mature size (see Fig. 1), these are: small breeds(Highland, Jersey and Casina); beef breeds, character-ized by their muscularity, wide pelvis andmedium height(Charolais, Limousin, South Devon, Pirenaica, Asturi-ana de los Valles, Piemontese and Aberdeen Angus), andtall breeds with average muscle development (Holstein,Simmental, Danish Red, Marchigiana and Avileña).

3.2. Carcass characteristics

Carcass characteristics for each breed showed highlysignificant differences among breeds for all variablesstudied (see Table 3). Carcass weights ranged from189.7 kg for Jersey to 386.6 kg for Charolais. In general,dairy breeds and local breeds produced lighter weightcarcasses than specialized beef breeds. In addition, thedairy and local breeds had low dressing percentage; thelowest for Jersey (50.1%), while the specialized beefbreeds (Charolais, Pirenaica, Asturiana de los Valles,Piemontese and Limousin) had values over 60%. Thebreed type was also reflected in conformation andfatness. The highest conformation score was 14.6 for thedouble muscled Piemontese and the lowest 4.4 forJersey, whereas fatness scores ranged from 3.6 for thePiemontese to 11.0 in the Aberdeen Angus. Similarly,kidney knob and channel fat weight (KKCW) rangedfrom 0.8 kg in Piemontese to 8.4 kg in Highland.Comparing the morphological traits, carcass lengthvaried from 119.1 cm in Highland to 135.1 cm inHolstein, and hind-limb length varied from 64.4 cm inHighland to 86.0 in Holstein and internal depth of breastvaried from 33.4 cm in Asturiana de los Valles to

Fig. 1. Cattle breeds means for body size measurements, adjusted at365 days of age by linear regression.

24 P. Albertí et al. / Livestock Science 114 (2008) 19–30

43.0 cm in Holstein. The blockiness index, ranged from1.6 kg/cm for Jersey to 2.9 kg/cm for Charolais.

Muscle depth measured in the loin area was highestfor Asturiana de los Valles and Piemontese (more than52 cm2) and lowest for Jersey (24.9 cm2), the greaterdifferences among breeds were due to the differences inthe dorso-ventral diameter (minimum diameter) ratherthan in the medium-lateral dimension (maximumdiameter). Muscle percentage measured from thedissection of the 6th rib dissection ranged from 58.9%in Holstein to 79.9% in Piemontese, whereas fatpercentage varied from 3.2% in Piemontese to 21.7%in Aberdeen Angus. Again the dairy and the local breedshad the highest values for fat and the lowest values formuscle percentages, as would be expected. Limousinhad the lowest bone percentage (15.0%).

3.3. Principal components analysis

The first two dimensions of the PCA analysis (factor1, 48.8%, factor 2, 24.6%) explained 73.4% of the vari-ation among breeds (see Fig. 2). The first dimension wasrelated to the blockiness index, carcass weight, loin area,minimum diameter of the loin, dressing percentage andconformation score, while the second dimension waspositively influenced by fat percentage measured at dis-section, fatness score and KKCW, and was negativelyrelated to muscle percentage at dissection. Therefore, abreed with high muscularity and low fatness, e.g., thePiemontese, appears in the right-bottom square of the plot.

4. Discussion

4.1. Weight and growth rate

The fifteen cattle breeds analysed here originate inEurope where, currently, animals are slaughtered in theearly stage of maturity to produce beef. Commercialrequirements for each country and the localmarkets differ,

so beef is produced fromboth bulls and steers. In the studyreported here, only entire bulls were studied. Consider-able variation was observed among the 15 breeds for liveweight gain, weight at all ages and morphological traits.This study underlines the large differences seen betweendairy breeds and specialized beef breeds as well amonglocal breeds, which were reflected in all the traitsrecorded. Previous research has shown that when cattleof diverse breeds are compared at a similar age, withsimilar management, some variation will be found incarcass weight, while carcass composition will depend onthe range of target weights and differences in growthcurves of each breed (Kempster et al., 1982). The dietoffered during the fattening period will influence thegrowth rate and carcass characteristics, especially fatness.In a study where breeds were compared at a constant fatthickness, breed group differences in composition werereduced by more than 50% relative to differences at aconstant age (Koch et al., 1979). Therefore, in this studyfat cover, which is a parameter used commercially toselect bulls for slaughter was not used, instead slaughterwas at a fixed weight within a target slaughter age. A highenergy/protein diet was offered ad libitum through thefattening period, and the bulls studied exhibited an almostlinear average daily gain. Two of the breeds, the AberdeenAngus and South Devon showed a very high live weightgain during the experimental period (9 to 12 months ofage, data not shown). This may have been a result ofcompensatory growth, following period on a restricteddiet prior to the animals entering the study. The highaverage daily gain in these breedswas unexpected, but hasalso been observed in another study. Chambaz et al.(2003) found that Angus steers, reared until they reachedthe same intramuscular fat content, had a 6 to 26% highergrowth rate than Limousin, Simmental or Charolaisbreeds, supporting the higher growth rate of this breed asobserved in our animals. Although this was not observedby others (e.g., Cuvelier et al., 2006). Asturiana de losValles, Piemontese and Marchigiana showed a moderateaverage daily gain compared to other beef breeds. It isknown, that for some beef breeds, gut and digestivesystem capacity, can limit intake and hence reduce liveweight gain. Cima (1996) observed a lower weight of therumen (−30%) and of the whole gut (−13%), forAsturiana de los Valles breed, than would be expectedof animals of this size, this limits their feed intake capacityand hence is likely to impact on growth rate comparedwith other breeds.

If the breeds used in this study were raised underdifferent management systems or diets, they could ex-hibit higher live gain: Asturiana de los Valles hasachieved 1.5 kg/day, Avileña 1.6 kg/day, Pirenaica

Table 3Means for carcass traits of 15 European cattle breeds

Breed AA ASV AVI CAS CHA HIG HOL JER LIM MAR PIE PIR RED SD SIM SE Significance

n 30 30 30 31 30 29 29 31 31 28 30 31 29 27 20Carcass weight (kg) 335.7de 348.7cd 324.7ef 244.7h 386.6a 245.1h 320.0efg 189.7h 360.0bc 307.5g 335.9de 371.5ab 318.7fg 346.9cd 344.4cd 29.46 ⁎

Dressing percentage 56.2ef 62.6ab 58.8d 56.2e 61.0c 55.2f 55.7g 50.1h 63.7a 58.8d 63.7a 61.1bc 54.9f 58.7d 55.5ef 2.29 ⁎

Kidney fat weight (kg) 7.1bc 2.3g 3.9f 3.9f 5.8 de 8.4a 6.1cd 7.5b 5.1e 1.4h 0.8h 3.2fg 6.2cd 5.4de 2.4g 1.79 ⁎

Conformation score (1–18) 9.8d 12.1b 8.0e 7.6e 9.9d 8.2e 4.6f 4.4f 10.2d 11.1c 14.6a 11.5bc 5.1f 10.2d 8.4e 1.55 ⁎

Fatness score (1–15) 11.0a 4.1gh 5.8e 5.9e 8.9bc 6.8d 8.2c 4.5fg 8.4c 5.0f 3.6h 4.9f 9.1b 6.0e 7.2d 1.40 ⁎

Carcass length (cm) 130.3c 127.1d 133.1ab 121.1f 133.0ab 119.1f 135.1a 119.8f 126.6 d 123.9e 123.3e 132.4bc 132.1bc 130.7bc 130.4c 4.30 ⁎

Internal depth of breast (cm) 40.3c 33.4f 37.0d 33.8ef 34.4ef 37.4d 43.0a 39.3c 34.7e 39.6c 37.3d 34.7e 41.7b 37.3d 41.7b 1.99 ⁎

Hind-limb length (cm) 70.8h 81.0e 83.4b 75.3f 83.2bc 64.4j 86.0a 67.3i 81.5de 72.8g 70.4h 82.1bcde 81.8cde 72.5g 83.0bcd 2.61 ⁎

Blockiness index (kg/cm) 2.6fg 2.7bcd 2.4ih 2.0j 2.9a 2.1j 2.4i 1.6k 2.8ab 2.5gh 2.7cde 2.8abc 2.4hi 2.7def 2.6ef 0.19 ⁎

Maximum diameter (cm) y 10.1bc 9.3ef 9.1f 8.6g 10.2bc 9.4ef 9.6de 8.1h 9.6de 10.2bc 10.3b 9.2ef 9.9cd 11.0a 9.6 def 0.78 ⁎

Minimum diameter (cm) y 5.1f 6.7a 5.3ef 5.2ef 5.8bcd 4.6g 4.4g 3.7h 6.1bc 5.5de 6.1bc 6.2b 4.6g 5.7cd 5.5 de 0.70 ⁎

LT area, (cm2) y 41.8ef 52.7a 40.3fg 35.8h 43.9e 35.0h 35.6h 24.9i 44.7de 47.8cd 52.2a 48.7bc 37.8gh 51.6ab 42.1ef 6.17 ⁎

Fat (%) z 21.7a 7.8g 12.6ef 14.7cd 15.4c 19.6b 19.3b 13.0def 13.2def 8.9g 3.2h 9.7g 19.5b 14.2cde 11.7f 3.53 ⁎

Bone and others (%) z 16.9f 17.2ef 18.3de 18.9d 16.9f 17.6ef 21.8a 20.4bc 15.0g 21.0ab 16.9f 17.5ef 19.4cd 17.3ef 20.5bc 2.35 ⁎

Muscle (%) z 61.4h 75.0b 69.1de 66.3g 67.7efg 62.8h 58.9i 66.5fg 71.9c 70.0d 79.9a 72.9c 61.1h 68.5def 67.8efg 3.49 ⁎

AA, Aberdeen Angus; ASV, Asturiana de los Valles; AVI, Avileña; CAS, Casina; CHA, Charolais; HIG, Highland; HOL, Holstein; JER, Jersey; LIM, Limousin; MAR, Marchigiana; PIE, Piemontese;PIR, Pirenaica; RED, Danish Red; SIM, Simmental; SD, South Devon.⁎Note for all parameters, Pb0.001. Different superscript letters in the same row indicate differences between breeds for at least 5%.y Measured on longissimus thoracis (LT) at 6th rib level.z Percentages from the 6th rib dissection.

25P.

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/Livestock

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(2008)19–30

Fig. 2. Principal component analysis of 15 European beef breeds in function of their carcass characteristics. CW—Carcass weight; DP—Dressingpercentage; KW–Kidney knob and channel fat weight; CS—Conformation score; FS—Fatness score; CL—Carcass length; BD—Internal depth ofbreast; LL—Limb length; BI–Blockiness index; MAD—Maximum diameter of the LTat 6th rib level; MID—Minimum diameter of the LT muscle at6th rib level; RA—LT area at the 6th rib level; FP—Fat percentage (6th rib dissection); BP—Bone and others percentage (6th rib dissection); MP—Muscle percentage (6th rib dissection).

26 P. Albertí et al. / Livestock Science 114 (2008) 19–30

1.7 kg/day (Albertí et al., 2001; Piedrafita et al., 2003);Limousin 1.62 kg/day, Holstein 1.23 kg/day (Andersenet al., 2001) and Danish Red 1.14 kg/day (Andersen,1974). They could also exhibit lower live gain, AberdeenAngus only achieved 1.66 kg/day when fattened with apulp or a cereal-based diet (Cuvelier et al., 2006), whileCharolais had an average daily gain of 1.3 kg/day and

Limousin 1.2 kg/day when they were fed with sugar beetpulp silage plus hay complemented with concentrate(Jurie et al., 2005).

The high rates of growth for Charolais and Simmentalseen in our study are in agreement with the work ofAndersen et al. (1977) and Cundiff et al. (1986) who usedthese sire breeds in a cross breeding study. The difference

27P. Albertí et al. / Livestock Science 114 (2008) 19–30

between the lighter weight breeds (Jersey, Highland andCasina), which had the lowest ADG (about 1.1 kg/day),and the heavy-weight breeds (Aberdeen Angus, SouthDevon, Charolais, Simmental and Limousin) which hadthe highest ADG (2.0 to 1.5 kg/day) are in agreement withthe positive correlation with adult body weight observedby Klosterman (1972). However, the use of ADG rate andlive weight is not a consistent parameter to sort breeds bytype (dairy, beef, or local) as dairy breeds (Holstein andDanish Red) grew more slowly and fattened later thanwould be expected from their target weights, whencompared with others breeds (Kempster et al., 1982).

4.2. Morphological measurements

The significant differences found among breeds for themorphological measurements regressed to a standardisedage of 365 days, confirmed the large phenotypicvariability between breeds and provided the basis forobjective classification in different breed types or groups.Jersey, Casina and Highland represent the small maturesize breeds, this group typically has narrow pelvisdevelopment, which indicates slow skeletal developmentand low muscling. Buchanan and Dolezal (1999)classified Jersey and Highland breeds as small by sizeand growth. In this respect, the Jersey breed would havethe typical shape of dairy breeds, but its narrow pelvis isemphasized by its small mature size. As would bepredicted from their small size, the slaughter weights werelower for this group than for other breeds, in addition, theaverage daily gain was also very low. In this study, thedairy breeds and local breeds such as Holstein, Sim-mental, Danish Red, Avileña and Marchigiana areclassified as tall animals with an intermediate pelviswidth. The Holstein breed was also classified as a tall andheavy breed by Long et al. (1979) in comparison withAngus, Brahman, Hereford and Jersey. The other breeds(Charolais, Aberdeen Angus, South Devon, Limousin,Piemontese, Asturiana de los Valles and Pirenaica)represent the specialized beef breeds, which werecharacterized by their wider pelvis, and medium witherheight. Overall, the grouping of breeds presented in thispaper is in agreement with other studies (Buchanan andDolezal, 1999).

4.3. Carcass traits

The differences found between breeds in carcassweight were expected. In the same way, the differencesfound among breeds for dressing percentage are inagreement with those given by several authors for thesame breeds: Wulf et al. (1996) for Limousin; Barton

and Pleasants (1997) for Angus and Jersey, Piedrafitaet al. (1999) and Albertí et al. (2000) for Asturiana de losValles, Avileña, Casina and Pirenaica, Cepin et al.(2000) for Holstein; Short et al. (2002) for Piemontese;Trombetta et al. (2005) for Marchigiana; Subrt and Divis(2002) for Charolais; Sami et al. (2004) for Simmental.Hence, with in limits, the carcass weight and dressingpercentage are more strongly influenced by the genetictype of the animal, reflected in the breed type, than bythe environment. The present study is the only one tohave compared such a wide range of pure breeds anddifferent genetic types and aptitudes. Additionally, it isthe only work in which some local or hardy breeds fromdifferent countries were compared with each other, i.e.,Highland (UK) and Casina (Spain).

In the present study, Piemontese and Asturiana de losValles breeds (also some of the South Devon) displaythe double muscling phenotype, which is more markedin hind-limbs than in forelimbs (Arthur, 1995). Thisexplains why conformation scores for these breeds werehigher than other breeds, as muscling and conformationscore is assessed from the roundness of limbs andshoulders. This also explains why conformation score isnot exactly reflected in blockiness index. In this study,Avileña, and Marchigiana showed similar blockinessindexes and the same percentage of muscle in dissec-tion, therefore it may be expected that conformationscores would be the same, however, the conformationscorers of these breeds differed by more than 3 points, asthe Avileña had longer hind-limbs than Marchigiana.

If the breeds are ranked by morphological measures,four breeds, Highland, Casina, Jersey and Piemontese,are found to have short carcasses and hind-limbs, how-ever, Jersey, Casina and Highland had a low blockinessindex, because of their lower slaughter weight, Piemon-tese had a high value for this trait, which was alsoreflected in the conformation scores. Similarly, Holstein,Danish Red, Avileña, Charolais and Pirenaica, did notchange their relative positions in the ranking of bothcarcass length and hind-limb length, but dairy and localbreeds had low blockiness indexes and poor conforma-tion scores, while specialized meat breeds remained atthe top for the latter two traits. In the European market,carcasses are valued by conformation (Drennan et al.,2002), which suggests that the blockiness index wouldbe a complementary tool to classify carcasses, especiallywhen very different types of breeds are compared in thesame market Díez et al. (2006).

Although variables related to muscular developmentwere good discriminators of breed type, variables de-fining fatness were not. Hence, if breeds are rankedby their fatness score, Jersey would be placed among

28 P. Albertí et al. / Livestock Science 114 (2008) 19–30

Piemontese, Asturiana de los Valles and Pirenaica.Conversely, Limousine and Charolais would be placedbetween Holstein and Danish Red. Ranking breeds byfat percentage in the dissection gave a group thatconsisted of beef breeds (Piemontese, Asturiana de losValles, Marchigiana and Pirenaica) but Limousin wasunexpectedly positioned between Jersey and SouthDevon, and Charolais was placed between Casina andHolstein. The same pattern was observed when rankingbreeds by KKCW. This indicates that fat variability washigher than muscle variability. From the dissection datathe coefficient of variation of fat percentage was nearlyfive times higher than that of muscle percentage.

Kempster et al. (1982) demonstrated that there is animportant genetic variation in partitioning of fat betweendifferent depots, and Casasús et al. (2000) reported thatin cattle this occurs even under similar nutritionalconditions. Beef cattle are characterized by their abilityto transform the nutrients mainly into proteins, whereasdairy breeds, which have a different hormonal andmetabolic status, tend to deposit more intra-abdominalfat (Kempster et al., 1982). Nevertheless, the variabilityof fat tissue could also depend on the differences in thephysiological ages of breeds at the same chronologicalage (Kempster et al., 1982; Micol et al., 1993), or on thedifferences in the growth potential between breeds(Jenkins et al., 1991; Arthur, 1995; Arthur et al., 1995;Albertí et al., 2000; Subrt and Divis, 2002; and Oliván etal., 2004). Therefore, since fat tissue development isaffected by multiple factors and there is not a clearrelationship between fatness and breed type.

4.4. Principal components analysis of breeds

The distribution of breeds plotted by PCA analysisreflected their production aptitude. Muscularity vari-ables, which influenced the first dimension, wereindependent of fatness measures, which stronglyinfluenced the second dimension. Therefore, breedsnaturally fell into three groups: the specialized beefbreeds are clustered spotted on the right side of the PCAplot, and included Piemontese, Asturiana de los Valles,Pirenaica, Limousin, South Devon, Charolais andAberdeen Angus. All these breeds had a high musclescore as expected for the beef breeds, however, theydiffer in fatness level. Aberdeen Angus was the fattest,in spite of its good conformation; and Piemontese wasthe leanest breed. The local and dairy breeds are placedon the left side of the plot and are represented by Jersey,Casina, Holstein, Highland and Danish Red. They hadlow muscling and medium (Casina) or highly fattened(Holstein, Simmental, Danish Red) carcases. Jersey had

low muscling. A third group of breeds fell betweenthe two previous groups, as an “intermediate” group(Avileña, Marchigiana, Simmental). These are meatproducing but not highly specialized breeds. Althoughoriginally considered as local, hardy or dairy breeds,they have undergone some selection and have improvedconformation characteristics. Therefore, they haveintermediate features: medium conformation and med-ium fatness level.

These results are consistent with those of Barton andPleasants (1997) who reported the same pattern of breeddistribution related to function. They found that themain variables explaining breed differences were boneor lean weights for the first axis and KKCW or rib eye(LT) for the second axis. Similarly, Albertí et al. (2005)in a study of seven Spanish breeds reported that the mostinfluential variables for classifying breeds were shapeand conformation for the first axis and measurements oflength and fat cover in the second axis.

Therefore, our results show that carcass traits andbody size measurements are useful to discriminatebetween breeds, probably because the muscle to boneratio is little affected by differences in degree of maturity,and each breed has a distinctive value over quite a widerange of fatness (Kempster et al., 1982). However, theuse of ADG rate and live weight is not a consistentparameter to group breeds by type (dairy, beef, or local)as within a short interval of weight it was possible to finddairy breeds (Holstein and Danish Red) that grow slowlyand fatten later than expected from theirs target weight incomparison with others breeds (Kempster et al., 1982).

For beef cattle breeds, the genetic selection for largesize, growth, or muscle development in Europeancountries is likely to have affected muscle biochemicalcharacteristics (Sudre et al., 2005), which could havesome impact on meat quality. This important question,which has been examined in the animals used in thisstudy, will be reported elsewhere.

5. Conclusions

The present study reports an interesting approach toobjectively describe morphological differences among arepresentative sample of cattle breeds from WesternEurope. The traditional types of cattle grouped as dairybreeds, specialized beef breeds and local breeds, can becomplemented by grouping the breeds according to theirslaughter weight (i.e., heavy, medium, and light) andanimal size (i.e., small/short/narrow, medium, and tall/large/wide). Indeed, these measurements together withthe carcass traits have provided a more useful method toclassify cattle breeds, than live weight or daily gain.

29P. Albertí et al. / Livestock Science 114 (2008) 19–30

The present study has demonstrated that body di-mensions and carcass traits differ between breeds, andthat animal types are clearly defined by these characters.These results provide valuable information when de-vising breeding strategies to meet the demands of theEuropean markets.

Acknowledgments

This work was funded in part by the EU Sixthframework programme project Assessment of GeneticVariation in Meat Quality and Evaluation of the Role ofCandidate Genes in Beef Characteristics with a View toBreeding for Improved Product Quality (GEMQUAL )QLRT-1999-30147.

The authors acknowledge the staff of the differentexperimental stations and abattoirs that were involved inrearing and slaughtering of the animals.

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