M. Enser

Fatty acid content and composition of UK beef and lamb muscle in relation to production system and implications for human nutrition.

Although ruminant meats normally have a low ratio of polyunsaturated fatty acids (PUFA) to saturated fatty acids (P:S ratio), the muscle contains a range of C(20) and C(22) PUFA of both the n-6 and n-3 series of potential significance in human nutrition. However, information on the amounts of these fatty acids in muscle and how they are modified by production system is limited In this study, the content and composition of fatty acids was determined in several muscles from beef steers fed grass (grazed) and bulls fed cereal concentrates. These are the two main types of beef production in the UK and Europe. Muscle fatty acids were also determined in lambs fed grass (grazed on pasture). The total fatty acid content of all muscles studied was less than 35 g kg(-1). The percentages in total fatty acids of all n-3 PUFA were higher in muscles from steers fed grass than from bulls fed concentrates whereas all n-6 PUFA were higher in the latter. The gluteobiceps muscle contained the largest amounts of fatty acids including PUFA and the m. longissimus dorsi the least amounts of PUFA in beef and lamb, and m. longissimus contained the lowest percentages of PUFA. Arachidonic acid was the major fatty acid in the C(20) + C(22) PUFA in beef from both production systems with twice as much in muscles from bulls fed concentrates. The P:S ratios were higher in the latter animals, range 0.21-0.34 compared with 0.08-0.13 in the steers fed grass. However, the n6:n-3 ratio was much less desirable in the bulls, 15.6-20.1 compared with 2.0-2.3 in the steers fed grass. These effects of production system in ruminants are larger than previously reported. Lamb muscle P:S ratios resembled those in grass-fed beef but the n-6:n-3 ratios were lower. The percentage of trans unsaturated 18:1 fatty acids was similar in both cattle production systems but lamb muscles contained twice as much as beef. Although the concentrations of the C(20) and C(22) PUFA are much lower than in fish, maintaining high n-3 levels in ruminant meats through grass feeding may be advantageous in human nutrition since meat is more widely consumed.

Factors influencing fatty acids in meat and the role of antioxidants in improving meat quality

Meat has been identified, often wrongly, as a food having a high fat content and an undesirable balance of fatty acids. In fact lean meat is very low in fat (20-50 g/kg), pork and poultry have a favourable balance between polyunsaturated and saturated fatty acids (P:S) and grazing ruminants produce muscle with a desirable n-6:n-3 polyunsaturated fatty acid ratio. In all species, meat fatty acid composition can be changed via the diet, more easily in single-stomached pigs and poultry where the linoleic, alpha-linolenic and long-chain polyunsaturated fatty acid content responds quickly to raised dietary concentrations. Recent work in pigs has attempted to manipulate the n-6:n-3 ratio by feeding higher levels of alpha-linolenic acid (e.g. in rapeseed) or its products eicosapentaenoic acid (20:5) and docosahexaenoic acid (22:6) present in fish oils. In ruminants the challenge is to increase the P:S ratio whilst retaining values for n-6:n-3 found in cattle and sheep fed on forage diets. The saturating effect of the rumen can be overcome by feeding polyunsaturated fatty acids which are protected either chemically, by processing, or naturally e.g. within the seed coat. Some protection occurs when grain-based or grass-based diets are fed normally, leading to relatively more n-6 or n-3 fatty acids respectively. These produce different flavours in cooked meat due to the different oxidative changes occurring during storage and cooking. In pigs and poultry, high n-3 fatty acid concentrations in meat are associated with fishy flavours whose development can be prevented with high dietary (supranutritional) levels of the antioxidant vitamin E. In ruminants, supranutritional vitamin E delays the oxidative change of oxymyoglobin to brown metmyoglobin and may also influence the characteristic flavours of beef and lamb.

Fatty acid content and composition of english beef, lamb and pork at retail.

We have determined the fatty acid content and composition of retail samples of meat and assessed them with respect to UK dietary recommendations. Fifty beef sirloin steaks, pork chops and lamb chops were purchased from four supermarkets on separate occasions. The percentage of muscle (boneless basis) in the samples was 84.4 ± 4.3, 69.8 ± 7.7 and 78.9 ± 7.1 for beef, lamb and pork, respectively, with fatty acid contents of 3.84 ± 1.3, 4.73 ± 1.66 and 2.26 ± 0.7 g per 100 g muscle, respectively. Adipose tissue fatty acid contents were 70.0 ± 8.2, 70.6 ± 8.6 and 65.3 ± 9.4 g per 100 g tissue. A range of C20 and C22 polyunsaturated fatty acids (PUFA) was present in the muscle of all three species and pork adipose tissue but their concentrations in lamb and beef adipose tissue were too low to measure. The mean P:S ratios for beef, lamb and pork muscle were (adipose tissue values in parentheses): 0.11 (0.05); 0.15 (0.09) and 0.58 (0.61), and the n-6:n-3 ratios were 2.1 (2.3), 1.3 (1.4) and 7.2 (7.6). We conclude that the muscles of red meat species are a valuable source of PUFA, particularly the C20 and C22 n-3 fatty acids, in the human diet and that, considered as part of a varied diet, the low P:S ratio of the ruminant muscle, the high n-6:n-3 ratio of pork and the total fatty acid contents do not detract significantly from the nutritional value of lean meat.

Manipulating the fatty acid composition of muscle and adipose tissue in beef cattle

Enhancing the n-3 polyunsaturated fatty acid (PUFA) content of beef is important in view of the generally saturated nature of fatty acids in ruminant meats and the negative effect this can have on human health. This study examined the effects of different sources of dietary n-3 PUFA on the performance of steers and the fatty acid composition of m. longissimus thoracis muscle and associated subcutaneous adipose tissue. Animals were fed ad libitum on grass silage plus one of four concentrates (60:40 forage:concentrate on a DM basis) containing differing sources of lipid: Megalac (16:0), lightly bruised whole linseed (18:3n-3), fish oil (20:5n-3 and 22:6n-3) and a mixture of linseed and fish oil (1:1, on an oil basis). Diets were formulated so that total dietary oil intake was 6 %, approximately half of which was from the experimental test oil. Linseed feeding not only increased the levels of 18:3n-3 in muscle phospholipid from 9.5 to 19 mg/100 g muscle but also enhanced the synthesis of 20:5n-3, the level of which increased from 10 to 15 mg/100 g muscle. Linseed also increased the proportion of 18:3n-3 in muscle neutral lipid and in adipose tissue lipids by a factor of 1.64 and 1.75 respectively. Fish oil feeding doubled the proportion of 20:5n-3 and 22:6n-3 in muscle phospholipids. The proportion of 18:1 trans in muscle neutral lipid was higher on the n-3 PUFA diets than the control diet, 0.04 and 0.02 respectively. Despite the implied modification to rumen metabolism, lipid source did not affect feed intake, growth rate, cold carcass weight or carcass fatness, but carcass conformation score was higher on fish oil treatments (P < 0.05). However, total muscle fatty acid content was not different between treatments and ranged from 3.5-4.3 % of tissue weight. The increase in n-3 PUFA in the meat produced by feeding linseed or fish oil lowered the n-6:n-3 ratio but had little effect on the P:S ratio.

Manipulating meat quality and composition.

Meat quality describes the attractiveness of meat to consumers. The present paper focuses on two major aspects of meat quality, tenderness and flavour. Both aspects of quality can be influenced by nutrition, principally through its effects on the amount and type of fat in meat. In several countries, high levels of intramuscular fat (marbling fat), i.e. above 30 g/kg muscle weight in longissimus, are deemed necessary for optimum tenderness, although poor relationships between fat content and tenderness have generally been found in European studies, where fat levels are often very low, e.g. below 10 g/kg in UK pigs. Muscle lipid may be a marker for red oxidative (type 1) muscle fibres which are found at higher concentrations in tender muscles and carcasses. Nutritional treatment can be used to manipulate the fatty acid content of muscle to improve nutritional balance, i.e. increase the polyunsaturated (PUFA): saturated fatty acid value and reduce the n-6:n-3 PUFA value. Increasing PUFA levels may also change flavour because of their greater susceptibility to oxidative breakdown and the generation of abnormal volatile compounds during cooking. This situation particularly applies to the n-3 PUFA which are the most unsaturated meat lipids. In pigs, a concentration of 3 mg alpha-linolenic acid (18:3)/100 mg in muscle and fat tissue fatty acids can easily be achieved by including whole linseed in the diet. This level has led to abnormal odours and flavours in some studies, but not in others. In cattle and sheep, feeding whole linseed raised 18:3 concentrations in muscle fatty acids from about 0.7 mg/100 mg to > 1 mg/100 mg. As with pigs, this diet also increased levels of long-chain n-3 PUFA formed from 18:3, including eicosapentaenoic acid (20:5). Although this increase led to greater oxidative breakdown of lipids during storage and the generation of large quantities of lipid-derived volatile compounds during cooking, there were no deleterious effects on odour or flavour. When 18:3 levels are raised in lamb and beef because of grass feeding, the intensity of the flavours increases in comparison with grain-fed animals which consume and deposit relatively more linoleic acid (18:2). In ruminants, very high levels of 18:2 produced by feeding protected oil supplements cause the cooked beef to be described as oily, bland or pork-like.

Flavour perception of oxidation in beef.

Lipid oxidation is a major factor in meat quality. In order to relate human perceptions of lipid oxidation, as determined by a trained taste panel, to a chemical measurement of oxidation, we studied meat from animals with a wide range of potential oxidation through differences in their PUFA composition and by displaying the meat in high oxygen modified atmosphere packs for varying lengths of time. Meat was obtained from 73 Angus- and Charolais-cross steers from different trials that had been raised on 10 different diets: grass silage (high in C18:3, n-3), cereal concentrate (high in C18:2, n-6), three diets with 3% added fat consisting of three levels of protected lipid supplement (high in C18:2, n-6 and C18:3, n-3, ratio 1:1), a control with Megalac(®) (relatively saturated), three diets with three levels of inclusion of protected fish oil (high in C20:5 n-3 and C22:6 n-3) plus a constant amount of unprotected fish oil and a final diet with an unprotected fish oil control. The longissimus dorsi muscle was excised from the left carcass side, aged vacuum packaged for 10-13 days depending on the projects and frozen for less than eight months. TBARS and sensory analyses were performed on steaks displayed for 0, 4 or 9 days under simulated retail conditions, exposed to light in modified atmosphere packaging (CO(2):O(2); 25:75). Meat oxidation increased throughout display for each of the diets, as shown by a rise in TBARS values. This increase was not linear, differences between 0 and 4 days of display were smaller than between 4 and 9 days of display. The lowest TBARS and lowest increment occurred in the two control diets and the grass-fed animals, probably due to the more saturated fat of meat from animals fed the control diets and the higher content of vitamin E. Sensory attributes were also influenced by time of display. Positive attributes, such as beef flavour or overall liking, decreased throughout display, whereas negative attributes, such as abnormal and rancid flavours, increased. The correlations between sensory and analytical attributes were high. TBARS were a good predictor of the perception of rancidity (Spearmans rho=0.84). Panellist preferences were related to the presence of beef flavour (rho=0.93) and to the absence of abnormal (rho=-0.88) and rancid flavours (rho=-0.83). Under the experimental conditions used, a TBARS value of around 2 could be considered the limiting threshold for the acceptability of oxidised beef.

Feeding linseed to increase the n-3 PUFA of pork: fatty acid composition of muscle, adipose tissue, liver and sausages

Eighty pigs, male and female littermate pairs, were fed a control or a test diet from 25 to 95 kg live weight. The diets, as fed, contained 15.5 g/kg linoleic acid (18:2) and 1.9 g/kg α-linolenic acid (18:3) (control) or 10 g/kg linoleic acid and 4 g/kg α-linolenic acid (test). The test diet, with added linseed, was, therefore, high in the main n-3 polyunsaturated fatty acid (PUFA) 18:3 and low in the main n-6 PUFA 18:2. Making this relatively small change led to a 56% increase in the content of 18:3 in muscle and major increases in the contents of the beneficial longer chain PUFAs EPA (20:5n-3) (100% increase) and DHA (22:6n-3) (35% increase) which are synthesised from 18:3n-3. Levels of EPA and DHA in pigmeat adipose tissue were also increased by the test diet. In liver, the test diet resulted in an 18:3 level 4× higher than in muscle, with 10× more EPA and 20× more DHA. Sausages, analysed after 6 months frozen storage also had high n-3 PUFA levels, due to the contribution of these fatty acids from both muscle and adipose tissue. From a health perspective these results confirm the potential of pigmeat to supply valuable n-3 PUFA to the human diet. The test diet produced a PUFA:saturated FA ratio in muscle of 0.4, close to the minimum recommended value for the diet as a whole and an n-6:n-3 ratio of 5, a significant improvement on the current average for pigmeat (7). It is estimated that the test diet would provide 12 g of long chain n-3 PUFA to the human diet per annum at current pigmeat consumption levels in the UK, about a third of that from oily fish.

Fatty acid composition and eating quality of lamb types derived from four diverse breed production systems

Carcass composition, muscle fatty acids and eating quality of loin chops were examined in ram lambs from four diverse breed × production system groups: pure Welsh Mountain off upland flora, pure Soays off lowland grass, Suffolk crosses off lowland grass and Suffolk crosses off concentrates. The two Suffolk groups had heavier and better muscled carcasses than the others and Soays were particularly lean. Fatty acid composition was different between the groups. The forage-fed lambs all had high concentrations of n-3 polyunsaturated fatty acids (PUFA) including 18:3 (α-linolenic acid) and 20:5 (eicosapentaenoic acid) compared with Suffolks-concentrates which had high concentrations of the n-6 PUFA 18:2 (linoleic acid) and 20:4 (arachidonic acid). Soays were high in both n-3 and n-6 PUFA. Flavour characteristics in grilled chops were similar in Welsh Mountain and Suffolks-grass which differed from Soays and Suffolks-concentrates. The latter two groups had low scores for lamb flavour and overall liking and high scores for abnormal lamb flavour, metallic, bitter, stale, and rancid. Soays had the highest score for livery. These results extend previous findings of the association between feed, PUFA composition and lamb flavour profile and confirm that forage-fed lamb is preferred by UK taste panellists. They also identify a specific breed effect on the quality of meat from lambs raised on forage. These findings suggest that possibilities exist for the production of meat with specific quality characteristics.

Effects of dietary fat source and breed on the carcass composition, n -3 polyunsaturated fatty acid and conjugated linoleic acid content of sheep meat and adipose tissue

Seventy-two 8-week-old ram lambs from three breeds, Suffolk, Soay and Friesland, were offered one of four diets based on dried grass and formulated to have a similar fatty acid content (60 g/kg DM) and containing: Megalac (high in 16 : 0, control; Volac Ltd, Royston, Herts., UK), whole linseed (18 : 3n-3), fish oil (20 : 5n-3 and 22 : 6n-3) or whole linseed plus fish oil. The lambs were slaughtered at approximately half of their mature live weight (43, 21 and 43 kg for Suffolk, Soay and Friesland lambs, respectively). Fish oil reduced DM intake and lamb live-weight gain (P<0.001), while DM intake, live-weight gain and subcutaneous fat content were highest in Suffolk and lowest in Soay lambs. Linseed feeding doubled the proportion (x100) of 18 : 3n-3 in the longissimus dorsi from 1.4 to 3.1 and in the subcutaneous adipose tissue from 1.2 to 2.6 (P<0.001). Suffolk and particularly Soay lambs contained higher proportions of 18 : 3n-3 than Friesland lambs in the longissimus dorsi, while in the adipose tissue, Suffolk lambs had the highest level. Feeding fish oil increased the muscle proportion (x100) of 20 : 5n-3 from 0.7 to 2.3 and 22 : 6n-3 from 0.3 to 0.8 (P<0.001). By contrast, the proportions of the longer-chain n-3 polyunsaturated fatty acids were similar across all three breeds. All three lipid supplements containing n-3 polyunsaturated fatty acids increased the content of muscle trans-18 : 1 relative to the control values, but conjugated linoleic acid (cis-9,trans-11-18 : 2) only increased in the muscle of lambs fed linseed. Feeding linseed or fish oil lowered the n-6 : n-3 ratio in sheep meat, but neither diet nor breed had much effect on the polyunsaturated fatty acid: saturated fatty acid ratio.

Effects of dietary n-3 polyunsaturated fatty acids, breed and dietary vitamin E on the fatty acids of lamb muscle, liver and adipose tissue

The effect of feeding n-3 PUFA on the fatty acid composition of muscle, adipose tissue and liver of lambs was investigated. Groups of eight ram lambs per breed, SuffolkxLleyn (24 kg live weight) and Scottish Blackface (18 kg live weight), were each fed one of six diets containing one of three fat sources (50 g fatty acids/kg DM; Megalac((R)) (calcium soap of palm fatty acid distillate; Volac Ltd, Royston, Herts., UK) and formaldehyde-treated whole linseed (Trouw Nutrition UK, Northwich, Ches., UK) either alone or with fish oil (1:1, w/w) and either 100 or 500 mg alpha-tocopheryl acetate/kg DM. Feed was offered ad libitum until slaughter at approximately half breed mature live weight. The type of dietary fat had no effect on intake, growth rate or feed conversion ratio. The 3.0-fold higher concentration of 18 : 3n-3 in the linseed compared with the Megalac((R)) diet approximately doubled (P<0.001) the concentration in the neutral and polar lipid fractions of musculus semimembranosus and liver, and in adipose tissue it increased 2.5-fold. Feeding protected linseed also increased (P<0.001) concentrations of 20 : 5n-3 and 22 : 5n-3 in muscle polar lipids and both lipid fractions of liver. The linseed-fish oil raised the 20 : 5n-3 concentrations above those for the linseed diet and also increased 22 : 6n-3. Scottish Blackface lambs had lower concentrations of 18 : 3n-3 in all lipids compared with Suffolk x Lleyn lambs, but more 20 : 5n-3 in the polar lipids of muscle and liver. High levels of dietary vitamin E were associated with small decreases in the concentration of monounsaturated fatty acids and increases in PUFA. Linseed raised the PUFA : saturated fatty acid ratios in liver and adipose tissue but not in muscle, and improved the n-6 : n-3 fatty acid ratio, as did the linseed-fish oil. Different combinations of dietary fatty acids and better protection against rumen biohydrogenation are required to improve muscle PUFA : saturated fatty acids ratios.