Meat Science 70 (2005) 597–603 www.elsevier.com/locate/meatsci

The eVect of production system and age on concentrations of fatty acids in intramuscular fat of the longissimus and triceps brachii muscles of Angus-cross heifers R.W. Purchas a

a,¤

, T.W. Knight b, J.R. Busboom

c

Institute of Food, Nutrition and Human Health, Massey University, Private Bag 11 222, Palmerston North, New Zealand b AgResearch Grasslands, Private Bag 11 008, Palmerston North, New Zealand c Department of Animal Sciences, Washington State University, Box 646310, Pullman, WA 99164-6310, USA Received 29 September 2004; received in revised form 22 December 2004; accepted 22 December 2004

Abstract The concentrations of fatty acids were measured in intramuscular fat from the longissimus lumborum (LL) and triceps brachii (TB) muscles of Angus-cross heifers Wnished either on a high-concentrate ration in Washington, USA, (US cattle, n D 15) or on pasture in New Zealand (NZ cattle, n D 16). Half of the NZ cattle were of a similar age to the US cattle (NZAge) and half were of a similar weight (NZWt). Intramuscular fat levels were higher for the LL muscle and for the US cattle but only within the LL muscle (P < 0.05). Aspects of the fatty-acid patterns that are of relevance to human nutrition tended to favour the pasture-Wnished NZ cattle with lower n ¡ 6/n ¡ 3 fatty acid ratios (P < 0.001), higher concentrations of an anticarcinogenic conjugated linoleic acid (C18:2 c9,t11) (P < 0.05) and its precursor (trans-vaccenic acid, TVA) (P < 0.01), and lower levels of the 18-carbon trans monounsaturated fatty acids other than TVA (P < 0.01). Concentrations of 20 of the 22 fatty acids analysed diVered signiWcantly between the two muscles. When values were adjusted to a common intramuscular fat level by covariance, most of the group diVerences remained, but a number of the muscle diVerences became non-signiWcant. For almost half the fatty acids considered, there was a signiWcant interaction between treatment group and muscle, which indicates that the results for one muscle do not necessarily apply to other muscles, although the ranking of the groups was usually the same for both muscles.  2005 Elsevier Ltd. All rights reserved. Keywords: Beef; Intramuscular fat; Pasture Wnished; Fatty acids; Conjugated linoleic acid; Omega-3 fatty acids

1. Introduction Although the amount of fat in the human diet is important because of its disproportionate contribution to dietary energy density, the nature of that fat also has important implications for well-being and health (Enser, 2001). Fat associated with beef has tended to be seen as having some undesirable characteristics, mainly because ¤ Corresponding author. Tel.: +64 6 350 4336x2536; fax: +64 6 350 5657. E-mail address: [email protected] (R.W. Purchas).

0309-1740/$ - see front matter  2005 Elsevier Ltd. All rights reserved. doi:10.1016/j.meatsci.2004.12.020

of the relatively high proportion of saturated fatty acids (SFA), some of which have been shown to be associated with higher levels of plasma cholesterol (Schaefer, 2002) and, as a result, an increased risk of cardio-vascular disease. More recently, however, it has become apparent that only some of the saturated fatty acids have hypercholesterolaemic eVects (Schaefer, 2002), that certain monounsaturated fatty acids (MUFAs) such as oleic acid (C18:1 c9) have a beneWcial rather than a neutral eVect (Moreno & Mitjavila, 2003), and that there are additional favourable characteristics of any fat source that should be taken into account when considering its overall nutritional value.

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R.W. Purchas et al. / Meat Science 70 (2005) 597–603

Examples of such favourable characteristics include lower ratios of n ¡ 6/n ¡ 3 polyunsaturated fatty acids (PUFAs) with particular emphasis on higher concentrations of the long-chain n ¡ 3 PUFA (Larsson, Kumlin, Ingelman-Sunberg, & Wolk, 2004), higher levels of certain conjugated linoleic acids (CLA) and their precursors (Pariza, 2004; Turpeinen et al., 2002), and lower levels of some trans-MUFAs such as elaidic acid (C18:1 t9) (Enser, 2001; Lock & Bauman, 2003). The objective in this study was to compare levels of important fatty acids in the intramuscular fat of two muscles from cattle Wnished within two contrasting Wnishing systems, and to assess the extent to which the levels of diVerent fatty acids were aVected by the amount of fat present within the muscle. The study diVered from other similar research in that production systems operating in diVerent countries were compared. This approach has the advantage of being a valid between-country and between-system comparison, but has the disadvantage of not being able to use a common population of animals at the outset, and of having several variables that diVer between the systems.

2. Materials and methods

et al. (2003). BrieXy, total lipid was measured by solvent extraction based on the method of Folch, Lee, and Sloane-Stanley (1957) and fatty acids as their methyl esters were quantiWed by gas-liquid chromatography (SGE BPX70 column 120 m length, 0.25 mm i.d.). Fatty acids are expressed here as percentages of the sum of all fatty acids measured. Fatty acids not listed in the tables due to low levels but included in the sum of all fatty acids were 15:1, 18:1 c, 18:2 tt, 18:2 ct, 18:3, 20:0, 21:0, 20:2, 24:0, and 24:1. The MUFA, C18:1 c/t is an unresolved mixture of cis and trans C18:1 fatty acids. 2.3. Statistical analysis Data analysis employed a block design within the GLM Procedure of SAS (SAS Institute Inc., Cary, NC), with individual animals as the blocks and the two muscles as samples within each block. Thus, the group eVect was tested against the animal-within-group term, and the eVects of animal, muscle, and group-by-muscle interaction were tested against the overall error term. Multiple comparisons between groups were tested using the least-signiWcant-diVerence method. Analyses of covariance within the GLM procedure were conducted in order to make comparisons with and without adjustment to a common level of intramuscular fat.

2.1. Animals and samples Longissimus lumborum (LL) and triceps brachii (TB) samples were obtained from the 31 Angus-cross heifers described in detail by Purchas and Busboom (2005). BrieXy, there were 15 heifers aged from 16–17 months that had been Wnished for 98 days on a high-concentrate diet (79% of the dry matter as concentrates with approximately equal amounts of corn, barley, and potato products) on a feedlot in Washington state, USA (US cattle) with a mean carcass weight of 322 kg, and 16 heifers Wnished on pasture in New Zealand (NZ cattle). The NZ cattle were in two groups, with half matching the US cattle for age (16–17 months), but were lighter (mean carcass weight of 260 kg) (NZAge group), and half that were older (27–28 months), but of a more similar carcass weight (294 kg) to the US cattle (NZWt group). The cattle were slaughtered and dressed according to normal practice in commercial meat plants in the respective countries. Samples were taken from the cranial end of the (LL) muscle (700–900 g), and from the central portion of the long head of the (TB) muscle (200–300 g). A sub-sample of approximately 20 g was taken from the internal part of each of these main samples within 36 hours of slaughter and frozen for subsequent analysis. 2.2. Analytical measurements The total lipid content and the concentrations of individual fatty acids were measured as described by Knight

3. Results and discussion Results are presented initially for fatty-acid concentrations relative to total fatty acid that have not been adjusted for the level of intramuscular fat (Tables 1–3), and then the eVects of adjusting to a common level of intramuscular fat are presented (Table 4). Generally, diVerences between the two pasture-Wnished NZ groups (NZAge and NZWt) were small and inconsistent compared with the diVerence between those groups and the US cattle. Therefore, the discussion below focuses on the latter comparison. 3.1. Unadjusted levels of fatty acids 3.1.1. Saturated and monounsaturated fatty acids Results for levels of saturated and monounsaturated fatty acids in LL and TB muscles (Table 1) are shown in a single row when the interaction between group and muscle was not signiWcant so that overall muscle means could be given, but in separate rows for the two muscles when the interaction was signiWcant (P < 0.05). The concentrations of short chain saturated fatty acids tended to be lower in the pasture-Wnished groups and to be higher for the LL muscle, with signiWcant interactions for some acids (Table 1). The exceptions for the group eVect were C16:0, where no group diVerence was apparent, and C18:0 where levels were lower for the US cattle.

R.W. Purchas et al. / Meat Science 70 (2005) 597–603

599

Table 1 Concentrations of saturated and monounsaturated fatty-acids in intramuscular fat from two muscles of feedlot-Wnished Angus-cross heifers from the USA (US cattle) and New Zealand pasture-Wnished Angus-cross heifers either of a similar age (NZAge) or a similar weight (NZWt) to the US cattle Fatty acid (% total fatty acids)

Musclea

C14:0

LL TB Both Both Both LL TB LL TB LL TB LL TB LL TB LL TB Both Both Both Both

C14:1 C15:0 C16:0 (palmitic) C16:1 C17:0 C18:0 (stearic) C18:1 t8 C18:1 t9 (elaidic) C18:1 t10 C18:1 t11 (TVA) C18:1 c/td C18:1 c9 (oleic) C18:1 c11

Musclea

Interaction eVectb

R2 (%), RSDc

¤¤¤

¤¤

89, 0.24

ns ¤ ¤¤¤ ¤¤¤

ns ns + ¤

87, 0.09 92, 0.04 92, 0.83 87, 0.31

¤¤¤

¤¤¤

¤¤¤

96, 0.08

¤¤¤

¤¤¤

¤¤¤

95, 0.69

¤¤¤

¤¤

¤¤¤

94, 0.05

¤¤¤

¤¤¤

¤¤¤

94, 0.05

¤¤¤

¤¤¤

¤¤¤

95, 0.25

¤¤ ns ¤¤¤ ¤¤¤

ns ns ns ns

91, 0.18 84, 0.07 98, 0.98 92, 0.18

Group US Cattle

NZAge

NZWt

EVectb

2.81c 2.09b 0.60b 0.43b 23.58 3.12c 3.79c 1.32c 1.13b 14.11a 11.05a 0.40c 0.26b 0.53c 0.40b 1.74c 1.15b 1.02a 0.37a 38.43a 2.38b

2.18b 1.95b 0.55ab 0.29a 23.07 2.96a 3.37b 0.75a 0.72a 15.02c 13.85b 0.12a 0.11a 0.23a 0.24a 0.01a 0.00a 1.55b 0.20b 39.99b 1.98a

2.17b 1.71a 0.44a 0.31a 23.43 2.94a 3.04a 0.77a 0.77a 15.92d 15.06c 0.11a 0.13a 0.25a 0.26a 0.00a 0.00a 1.55b 0.24b 38.90ab 2.09a

¤¤

LL

+ ¤¤¤ ns +

0.52 0.35 24.91

1.44 0.27 38.55 1.86

¤¤¤ ¤¤¤ + ¤¤¤

TB

0.55 0.33 21.81

1.30 0.27 39.67 2.44

EVectb

a LL, longissimus lumborum muscle; TB, triceps brachii muscle. Results are shown in separate rows for the two muscles only when the interaction was signiWcant (P < 0.05). b ¤¤¤, P < 0.001; ¤¤, P < 0.01; ¤, P < 0.05; +, P < 0.10; ns, P > 0.10. c Measures of goodness of Wt of the model are given by coeYcients of determination [R2 (%)] and residual standard deviations (RSD). Means for diVerent groups for a particular fatty acid do not diVer signiWcantly (P > 0.05) if they have a common letter or if they have no letters. d C18:1 c/t is an unresolved mixture of cis and trans C18:1 fatty acids.

Table 2 Concentrations of polyunsaturated fatty-acids in intramuscular fat from two muscles of feedlot-Wnished Angus-cross heifers from the USA (US cattle) and New Zealand pasture-Wnished Angus-cross heifers either of a similar age (NZAge) or a similar weight (NZWt) to the US cattle Fatty acid (% total fatty acidsd)

Musclea

C18:2 n ¡ 6 (linoleic)

LL TB LL TB Both Both LL TB Both Both Both

C18:3 n ¡ 3 (linolenic) C18:2 c9,t11 (CLA) C20:3 n ¡ 6 C20:4 n ¡ 6 (AA) C20:5 n ¡ 3 (EPA) C22:5 n ¡ 3 (DPA) C22:6 n ¡ 3 (DHA)

Musclea

Group b

LL

TB

EVect

b

Interaction eVectb

R2 (%), RSDc

US Cattle

NZAge

NZWt

EVect

2.43b 4.64d 0.29a 0.48b 0.37a 0.21 0.53a 1.25c 0.24a 0.50a 0.03a

1.70a 2.33b 1.26c 1.56d 0.50b 0.21 0.84b 1.05bc 0.80b 0.96b 0.14c

1.58a 2.88c 1.23c 1.81e 0.40ab 0.24 0.66a 1.16bc 0.72b 0.87b 0.09b

¤¤¤

¤¤¤

¤¤¤

92, 0.53

¤¤¤

¤¤¤

¤¤¤

97, 0.14

+ ns ns

0.39 0.14

0.46 0.30

¤¤ ¤¤¤ ¤¤¤

ns ns ¤¤

77, 0.10 75, 0.11 82, 0.24

¤¤¤ ¤¤¤ ¤¤¤

0.47 0.61 0.07

0.70 0.94 0.11

¤¤¤ ¤¤¤ ¤¤¤

+ ns ns

91, 0.14 87, 0.18 84, 0.04

a LL, longissimus lumborum muscle; TB, triceps brachii muscle. Results are shown in separate rows for the two muscles only when the interaction was signiWcant (P < 0.05). b ¤¤¤, P < 0.001; ¤¤, P < 0.01; ¤, P < 0.05; +, P < 0.10; ns, P > 0.10. c Measures of goodness of Wt of the model are given by coeYcients of determination [R2 (%)] and residual standard deviations (RSD). Means for diVerent groups for a particular fatty acid do not diVer signiWcantly (P > 0.05) if they have a common letter or if they have no letters. d AA, arachidonic acid; EPA, eicosapentaenoic acid; DPA, docosapentaenoic acid; DHA, docosahexaenoic acid.

Levels of the trans C18:1 showed strong interactions between groups and muscles with lower levels in the TB muscle being more apparent for the US cattle. Group diVerences for these acids diVered markedly, with samples from US cattle being higher for t8, t9 (elaidic acid) and t10, but lower for t11 (trans-vaccenic acid, TVA). This group of

acids is of interest because on the one hand, some members (particularly elaidic acid) are considered undesirable because of associations with high levels of plasma LDL cholesterol, while TVA as a precursor to CLA is considered beneWcial (Lock & Bauman, 2003). In this respect, the pattern of trans isomers favours the pasture-Wnished cattle.

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Table 3 Concentrations of groups of fatty-acids and ratios between groups of fatty acids in intramuscular fat from two muscles of feedlot-Wnished Anguscross heifers from the USA (US cattle) and New Zealand pasture-Wnished Angus-cross heifers either of a similar age (NZAge) or a similar weight (NZWt) to the US cattle Group or ratio

Musclea

Musclea

Group US Cattle

NZAge

NZWt

EVect

Groups of fatty acids SFA LL TB MUFA Both PUFA LL TB LC n ¡ 3 Both tMUFA-TVA LL TB

43.97d 36.75a 48.86b 4.33a 8.24d 0.78a 2.67c 1.81b

42.63c 38.72b 47.87ab 6.21c 8.02d 1.91b 0.36a 0.35a

44.51d 39.50b 46.79a 5.55b 8.71d 1.67b 0.36a 0.39a

¤

Ratios of fatty-acid groups PUFA:SFA LL TB MUFA:SFA LL TB n ¡ 6/n ¡ 3 Both C18:2:C18-3 Both

0.10a 0.23c 1.09a 1.36d 4.58b 9.46b

0.15b 0.21c 1.10a 1.26c 1.14a 1.41a

b

LL

TB

Interaction eVectb

R2 (%), RSDc

¤¤¤

¤¤¤

95, 1.19

EVect

b

d

0.12ab 0.22c 1.04a 1.21b 1.21a 1.42a

¤¤ +

46.93

48.75

¤¤¤ ¤¤¤

ns ¤

87, 1.10 86, 1.16

¤¤¤ ¤¤¤

1.15

1.76

¤¤¤ ¤¤¤

ns ¤¤¤

89, 0.35 96, 0.30

ns

¤¤¤

¤

86, 0.04

¤¤

¤¤¤

¤¤¤

95, 0.05

ns ¤¤

ns ns

99, 0.18 98, 0.80

¤¤¤ ¤¤¤

2.32 3.83

2.31 4.36

a LL, longissimus lumborum muscle; TB, triceps brachii muscle. Results are shown in separate rows for the two muscles only when the interaction was signiWcant (P < 0.05). b ¤¤¤, P < 0.001; ¤¤, P < 0.01; ¤, P < 0.05; +, P < 0.10; ns, P > 0.10. c Measures of goodness of Wt of the model are given by coeYcients of determination [R2 (%)] and residual standard deviations (RSD). Means for diVerent groups for a particular fatty acid do not diVer signiWcantly (P > 0.05) if they have a common letter or if they have no letters. d SFA, saturated fatty acids; MUFA, monounsaturated fatty acids; PUFA, polyunsaturated fatty acids; LC n ¡ 3, long-chain n ¡ 3 acids (EPA, DPA, and DHA); tMUFA-TVA, trans monounsaturated fatty acids except TVA.

The most abundant C18:1 fatty acids present were the cis isomers, particularly c9, which was slightly lower for the US cattle, and c11, which was higher for the US cattle. For both these acids the levels were higher in the TB muscle. 3.1.2. Polyunsaturated fatty acids All the polyunsaturated fatty acids in Table 2 were present at higher levels in the TB muscle. The strong group diVerences shown were generally consistent with those shown in other studies where beef from pastureWnished cattle has been compared with that from cattle Wnished on high-grain diets, with the latter group having higher levels of C18:2 n ¡ 6, and lower levels of both C18:3 n ¡ 3, and the longer-chain n ¡ 3 fatty acids (EPA, DHA, DPA) (Elmore et al., 2004; Engle & Spears, 2004; Enser et al., 1998; French et al., 2000; Itoh, Johnson, Cosgrove, Muir, & Purchas, 1999; Marmer, Maxwell, & Williams, 1984; Realini, Duckett, Brito, Dalla Rizza, & De Mattos, 2004a; Realini, Duckett, & Windham, 2004b; Yang, Lanari, Brewster, & Tume, 2002). The eVects on levels of C18:2 n ¡ 6 have varied considerably in size, signiWcance and direction between studies, but the eVect on the n ¡ 3 fatty acids has been a large and consistent eVect that has been attributed to the high levels of C18:3 n ¡ 3 in grasses (Body & Hansen, 1978; Engle & Spears, 2004; French et al., 2000; Shorland, 1961). Smaller non-signiWcant eVects reported for some studies can probably be attributed to lower levels of con-

centrates in the non-pasture diet or to shorter trial periods (Varela et al., 2004). The only CLA detected consistently in the current study was the c9 t11 isomer (Table 2), which is the one that has been shown to possess signiWcant anti-carcinogenic properties (Enser, 2001; Lock & Bauman, 2003). Levels were higher in the TB muscle and also by 21% in beef from pasture-Wnished cattle, although this diVerence was marginally signiWcant. TVA as a precursor to CLA, was, however, about 50% higher in the beef of the pasture-Wnished groups (P < 0.001) (Table 1). Several other reports have shown that CLA c9 t11 is present at signiWcantly higher levels in beef from cattle Wnished on pasture relative to those Wnished on grain-based diets (Engle & Spears, 2004; French et al., 2000; Realini et al., 2004a, 2004b; Yang et al., 2002). Elmore et al. (2004) found no signiWcant diVerences for beef from cattle Wnished on grass silage relative to those on a 79% concentrate diet, but dairy cows fed fresh pasture had higher levels of CLA in their milk (Lock & Garnsworthy, 2003). Diets containing grain tend to lead to a lower rumen pH which results in less conversion of TVA to stearic acid (Bauman, PerWeld, de Veth, & Lock, 2003). 3.1.3. Groups and ratios Results for the levels of groups of fatty acids and for ratios between the groups in Table 3 essentially provide a summary of the results from Tables 1 and 2. SFA were higher for the LL muscle, but showed only small

R.W. Purchas et al. / Meat Science 70 (2005) 597–603

601

Table 4 Relationships between the concentrations of fatty acids in intramuscular fat of two muscles and the level of intramuscular fat (%), and the eVect of adjusting for intramuscular fat levels on the signiWcance of group and muscle diVerences Changea with fat % adjustment in

Regression

Group eVect

Muscle eVect

R2 (%)b

Individual saturated and monounsaturated fatty acids (% of total fatty acids) C14:0 +0.151 ¤¤¤ C14:1 +0.071 ¤¤¤ C15:0 +0.023 ¤¤¤ C16:0 +0.122 ¤¤¤ C16:1 +0.188 ns C17:0 +0.044 ¤¤¤ C18:0 +0.063 ns C18:1 t8 +0.029 ¤¤¤ C18:1 t9 +0.005 ¤¤¤ C18:1 t10 +0.123 ¤¤¤ C18:1 t11 (TVA) +0.055 ns C18:1ct ¡0.016 ¤¤ C18:1 c9 +0.278 ¤¤¤

¤¤ ! ¤ + ! ns ¤¤¤ ! ¤¤¤ ns ! ns + ! ns ¤¤¤ ! ¤¤¤ ¤¤¤ ! ¤¤¤ ¤¤¤ ! ¤¤¤ ¤¤¤ ! ¤¤¤ ¤¤¤ ! ¤¤¤ ¤¤¤ ! ¤¤¤ ¤¤¤ ! ¤¤¤ +!+

¤¤¤ ! ¤¤¤ ns ! ¤¤ ¤ ! ns ¤¤¤ ! ¤¤¤ ¤¤¤ ! ¤¤¤ ¤¤¤ ! ns ¤¤¤ ! ¤¤¤ ¤¤ ! ns ¤¤¤ ! ns ¤¤¤ ! ns ¤¤ ! ns ns ! ns ¤¤¤ ! ¤¤¤

89 ! 92 87 ! 92 92 ! 94 92 ! 92 87 ! 90 96 ! 97 95 ! 95 94 ! 95 94 ! 94 95 ! 96 91 ! 92 85 ! 85 86 ! 87

Individual polyunsaturated fatty acids (% of total fatty acids) C18:2 ¡0.443 C18:3 ¡0.105 C18:2 c9 t11 (CLA) +0.034 C20:3 n ¡ 6 ¡0.043 C20:4 n ¡ 6 (AA) ¡0.194 C20:5 n ¡ 3 (EPA) ¡0.120 C22:5 n ¡ 3(DPA) ¡0.157 C22:6 n ¡ 3 (DHA) ¡0.021

¤¤¤ ¤¤¤ ns ¤¤¤ ¤¤¤ ¤¤¤ ¤¤¤ ¤¤¤

¤¤¤ ! ¤¤¤ ¤¤¤ ! ¤¤¤ +!¤ ns ! ns ns ! ns ¤¤¤ ! ¤¤¤ ¤¤¤ ! ¤¤¤ ¤¤¤ ! ¤¤¤

¤¤¤ ! ¤¤¤ ¤¤¤ ! ¤¤¤ ¤¤ ! ¤ ¤¤¤ ! ¤¤ ¤¤¤ ! ¤¤¤ ¤¤¤ ! ¤¤ ¤¤¤ ! ¤¤¤ ¤¤¤ ! ¤

92 ! 96 97 ! 98 77 ! 79 75 ! 77 82 ! 89 91 ! 95 87 ! 93 84 ! 87

Groups of fatty acidsc (% of total) and ratios PUFA ¡0.969 SFA +0.422 PUFA:SFA ¡0.028 MUFA:SFA ¡0.004 n ¡ 6/n ¡ 3 +0.139

¤¤¤ ¤¤¤ ¤¤¤ ¤¤¤ ¤¤¤

+ ! ns ¤!¤ ns ! ns ¤¤ ! ¤ ¤¤¤ ! ¤¤¤

¤¤¤ ! ¤¤¤ ¤¤¤ ! ¤¤¤ ¤¤¤ ! ¤¤¤ ¤¤¤ ! ¤¤¤ ns ! +

86 ! 92 95 ! 95 86 ! 92 95 ! 95 99 ! 99

CoeYcient

a b c

EVecta

¤¤¤, P < 0.001; ¤¤, P < 0.01; ¤, P < 0.05; +, P < 0.10; ns, P > 0.10. CoeYcient of determination. SFA, saturated fatty acids; PUFA, polyunsaturated fatty acids; MUFA, monounsaturated fatty acids.

diVerences between the groups, while MUFA and PUFA were higher in the TB muscle and also showed only small diVerences between the groups, although MUFA levels were higher for the US cattle than the NZWt group. The long-chain n ¡ 3 fatty acids were clearly higher in the pasture-Wnished groups, while the trans-MUFA fatty acids other than TVA were lower in these groups. Apart from the last Wnding, which does not appear to have been reported before, these results are generally consistent with those of other comparisons of intramuscular fat from cattle Wnished on pasture or concentrates, including those referenced in the previous section and the results of Steen, Lavery, Kilpatrick, and Porter (2003). The TB muscle contained higher levels of the LC n ¡ 3 acids, and lower levels of the trans-MUFA fatty acids other than TVA, although the latter eVect was only apparent for the US cattle (Table 3). 3.2. Fatty-acid levels adjusted for fatness In order to assess the extent to which some of the group and muscle eVects reported in Tables 1–3 were

due to diVerences in intramuscular fat levels (Fig. 1), intramuscular fat level as a percentage of muscle weight was Wtted Wrst as a covariate in the general-least-squares model (type I sums of squares). The results in terms of the regression coeYcient, the signiWcance of the relationship with intramuscular fat, and the change in the level of signiWcance of group and muscle eVects are given in Table 4. The change in the coeYcient of determination [R2 (%)] when the covariate was included is also shown. All the items shown in Table 4 were signiWcantly related to the level of intramuscular fat except CLA and its precursor TVA, and C16:1 and C18:0. The absence of a signiWcant relationship between intramuscular fat levels and the concentration of CLA has also been reported for beef by Knight et al. (2003), which suggests the CLA has similar concentrations in both neutral and polar lipids. For pigs, Demaree, Gilbert, Mersmann, and Smith (2002) reported that CLA was not preferentially incorporated into any subcellular fraction of muscle or adipose tissue. With the exception of C18:1 ct, all the saturated and monounsaturated fatty acids in Table 4 increased with

602

R.W. Purchas et al. / Meat Science 70 (2005) 597–603

Fig. 1. Means (§SE) for intramuscular fat in the longissimus lumborum (LL) muscle and the triceps brachii (TB) muscle of Angus-cross heifers that were Wnished on a high-grain feedlot ration in Washington State (US) or on pasture in New Zealand (NZAge and NZWt). Bars without a common letter above them diVer (P < 0.05).

increased levels of intramuscular fat, which suggests that they comprise a greater proportion in neutral depot lipids than polar structural lipids, as the proportion of polar lipids is known to decrease as intramuscular fat levels increase. Itoh et al. (1999), for example, reported a correlation of ¡0.90 between the proportion of polar lipids and the level of intramuscular fat in beef longissimus muscle. In contrast, all the PUFA in Table 4 except CLA showed a signiWcant negative relationship with intramuscular fat levels, indicating that they were primarily in the structural polar lipids. Even though there were signiWcant group diVerences in intramuscular fat levels, particularly for the LL muscle as shown in Fig. 1, the adjustment to a common fatness level brought about only small changes in the signiWcance of the group eVect (Table 4) and no changes in the ranking of the least-squares means that diVered signiWcantly. This suggests that diVerences between the production systems were not due to diVerences in intramuscular fat levels, but to other factors, with the diVerent diets probably being important. For diVerences between the two muscles, many of these were retained after adjustment to a common fat level, but there were also many where the level of statistical signiWcance was reduced, or where it became non-signiWcant. The group of trans C18:1 acids, for example, all became non-signiWcant with regard to muscle diVerences, as did C17:0 and C15:0. Adjustment to a common level of fatness did not

Fig. 2. Changes in the combined concentrations of three long-chain n ¡ 3 fatty acids (EPA, DPA, DHA) as a percentage of total fatty acids (TFA), with increasing levels of intramuscular fat. Results are shown for the LL muscle (circles) and the TB muscle (triangles) from the US cattle (Wlled symbols) and NZ cattle (open symbols).

change the ranking of the two muscles for those fatty acids where the diVerence was still signiWcant. An example of the nature of the relationship between intramuscular fat level and the concentration of fatty acids is provided for the sum of three long-chain n ¡ 3 fatty acids (Fig. 2), where higher levels of intramuscular fat were associated with lower concentrations of these muscles, presumably because they are mainly present in the polar structural lipids that will become a lower proportion of total fatty acids as intramuscular fat increases (Itoh et al., 1999). Extrapolating the present results to other situations involving comparisons between Wnishing diets based on either pasture or a grain-based high-concentrate feedlot diet needs to be done with caution, as these results apply strictly to only the speciWc diets involved and there were a number of confounding variables. It is reassuring, however, to note that many of the diVerences shown are consistent with other similar studies.

4. Conclusions Intramuscular fat in beef from pasture-Wnished cattle diVers from that of cattle Wnished on a high-grain diet in several nutritionally beneWcial ways, including having a lower ratio of n ¡ 6/n ¡ 3 PUFAs, lower levels of trans C18:1 except for TVA, and higher levels of both an anticarcinogenic CLA as well as its precursor (TVA).

R.W. Purchas et al. / Meat Science 70 (2005) 597–603

The pattern of fatty acids diVers signiWcantly between the LL muscle and the TB muscle, with some of the diVerences being attributable to the higher intramuscular fat levels in the LL muscle. In this study, there were a number of signiWcant interactions between the experimental groups and the muscles, but these generally were not associated with diVerences in the ranking of either the groups or the muscles. Acknowledgements This research was carried out under contract to Meat Biologics Research Limited, New Zealand. The technical contributions by Mike Agnew and Andrea Death are gratefully acknowledged, as is the assistance of personnel at the meat plants of AFFCo Manawatu Ltd., and Washington Beef Inc. References Bauman, D. E., PerWeld, J. W., de Veth, M. J., & Lock, A. L. (2003). New perspectives on lipid digestion and metabolism in ruminants. In Proceedings of the 2003 Cornell nutrition conference for feed manufacturers (pp. 175–189). Ithaca, NY: Department of Animal Science, Cornell University. Body, D. R., & Hansen, R. P. (1978). The occurrence of C13 to C31 branched-chain fatty acids in the faeces of sheep fed rye grass, and of C12 to C34 normal acids in both the faeces and the rye grass. Journal of the Science of Food and Agriculture, 29, 107–114. Demaree, S. R., Gilbert, C. D., Mersmann, H. J., & Smith, S. B. (2002). Conjugated linoleic acid diVerentially modiWes fatty acid composition in subcellular fractions of muscle and adipose tissue but not adiposity of postweanling pigs. Journal of Nutrition, 132, 3272– 3279. Elmore, J. S., Warren, H. E., Mottram, D. S., Scollan, N. D., Enser, M., Richardson, R. I., & Wood, J. D. (2004). A comparison of the aroma volatiles and fatty acid compositions of grilled beef muscle from Aberdeen Angus and Holstein-Friesian steers fed diets based on silage or concentrates. Meat Science, 68, 27–33. Engle, T. E., & Spears, J. W. (2004). EVect of Wnishing system (feedlot or pasture), high-oil maize, and copper on conjugated linoleic acid and other fatty acids in muscle of Wnishing steers. Animal Science, 78, 261–269. Enser, M. (2001). The role of fats in human nutrition. In B. Rossell (Ed.), Animal carcass fats (Vol. 2). Oils and fats (pp. 77–123). Leatherhead, England: Leatherhead Publishing. Enser, M., Hallet, K. G., Hewett, B., Fursey, G. A. J., Wood, J. D., & Harrington, G. (1998). Fatty acid content and composition of UK beef and lamb muscle in relation to production system and implications for human nutrition. Meat Science, 49, 329–341. Folch, J., Lee, M., & Sloane-Stanley, G. H. A. (1957). A simple method for the isolation and puriWcation of total lipid from animal tissue. Journal of Biological Chemistry, 226, 497–509. French, P., Stanton, C., Lawless, F., O’Riordan, E. G., Monahan, F. G., CaVret, P. J., & Moloney, A. P. (2000). Fatty acid composition, including conjugated linoleic acid, of intramuscular fat from steers oVered grazed grass, grass silage, or concentrate-based diets. Journal of Animal Science, 78, 2849–2855.

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The effect of production system and age on ...

(P < 0.05). Aspects of the fatty-acid patterns that are of relevance to human nutrition tended to favour the .... Data analysis employed a block design within the.

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