Nigel D. Scollan

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.

Recent developments in altering the fatty acid composition of ruminant-derived foods

There is increasing evidence to indicate that nutrition is an important factor involved in the onset and development of several chronic human diseases including cancer, cardiovascular disease (CVD), type II diabetes and obesity. Clinical studies implicate excessive consumption of medium-chain saturated fatty acids (SFA) and trans-fatty acids (TFA) as risk factors for CVD, and in the aetiology of other chronic conditions. Ruminant-derived foods are significant sources of medium-chain SFA and TFA in the human diet, but also provide high-quality protein, essential micronutrients and several bioactive lipids. Altering the fatty acid composition of ruminant-derived foods offers the opportunity to align the consumption of fatty acids in human populations with public health policies without the need for substantial changes in eating habits. Replacing conserved forages with fresh grass or dietary plant oil and oilseed supplements can be used to lower medium-chain and total SFA content and increase cis-9 18:1, total conjugated linoleic acid (CLA), n-3 and n-6 polyunsaturated fatty acids (PUFA) to a variable extent in ruminant milk. However, inclusion of fish oil or marine algae in the ruminant diet results in marginal enrichment of 20- or 22-carbon PUFA in milk. Studies in growing ruminants have confirmed that the same nutritional strategies improve the balance of n-6/n-3 PUFA, and increase CLA and long-chain n-3 PUFA in ruminant meat, but the potential to lower medium-chain and total SFA is limited. Attempts to alter meat and milk fatty acid composition through changes in the diet fed to ruminants are often accompanied by several-fold increases in TFA concentrations. In extreme cases, the distribution of trans 18:1 and 18:2 isomers in ruminant foods may resemble that of partially hydrogenated plant oils. Changes in milk fat or muscle lipid composition in response to diet are now known to be accompanied by tissue-specific alterations in the expression of one or more lipogenic genes. Breed influences both milk and muscle fat content, although recent studies have confirmed the occurrence of genetic variability in transcript abundance and activity of enzymes involved in lipid synthesis and identified polymorphisms for several key lipogenic genes in lactating and growing cattle. Although nutrition is the major factor influencing the fatty acid composition of ruminant-derived foods, further progress can be expected through the use of genomic or marker-assisted selection to increase the frequency of favourable genotypes and the formulation of diets to exploit this genetic potential.

Effects of breed and a concentrate or grass silage diet on beef quality in cattle of 3 ages. I: Animal performance, carcass quality and muscle fatty acid composition.

An increase in the intake of the n-3 series polyunsaturated fatty acids (PUFA) is recommended by nutritionists for the human diet and beef is a significant source of these fatty acids. Enhancing the n-3 PUFA content of beef is important in view of the generally saturated nature of fatty acids in ruminant meats and the potentially negative effect this can have on human health. This study examined the effects of breed and diet on the fatty acid composition of beef M. longissimus. Ninety-six steers were used, 48 Aberdeen Angus cross (AA) and 48 Holstein-Friesian (HF). At 6months of age, 3 groups were identified, to be slaughtered at 14, 19 and 24months, respectively. Each group consisted of eight steers of each breed fed on a concentrate or a grass silage diet, rich in n-6 and n-3 PUFA, respectively. The intake of the concentrate diet was restricted so that steers of each breed grew at a similar rate on each diet. The early maturing AA produced heavier, fatter carcasses with better conformation. Animals fed grass silage had higher carcass fatness and conformation scores and higher levels of neutral lipid and total lipid in muscle than those fed concentrate. When all animals were pooled, a decline in PUFA% as total muscle lipid increased was evident. Feeding a grass silage diet rich in α-linolenic acid (18:3n-3) increased levels of this fatty acid in muscle neutral lipid by a factor of about 3.0 compared with the concentrate diet, as well as enhancing the synthesis of the n-3 series long-chain C20-22 PUFA in the phospholipid fraction, including docosahexaenoic acid (DHA, 22:6n-3). In contrast, both levels and proportions of linoleic acid (18:2n-6) and the n-6 series C20-22 PUFA were higher in animals fed the concentrate diet. The proportions of 18:1trans and conjugated linoleic acid (CLA) in muscle neutral lipid were higher in animals fed concentrate compared with silage in all 3 groups. This was partly due to increased consumption of 18:2n-6. The ratio of PUFA to saturated fatty acids (P:S) in muscle was reduced by feeding grass silage, partly as the result of increased fat deposition. However, the increase in levels of n-3 series fatty acids with silage-feeding resulted in beneficially low n-6:n-3 ratios in muscle in all age groups (approximately 1.2 compared with 12.0 in the concentrate diet). Subtle breed differences in PUFA amounts and proportions were noted. Holstein-Friesians had higher proportions of PUFA and higher P:S ratios compared with AA, partly due to a higher proportion of phospholipid in total lipid. In phospholipid itself, HF in the 19 and 24months groups had higher proportions of most n-3 PUFA. In all age groups the ratio of DHA to its precursor, 18:3n-3 was higher in HF.

Effect of dietary lipid on the content of conjugated linoleic acid (CLA) in beef muscle

We have examined the effect of dietary fats containing n-3 polyunsaturated fatty acids on the conjugated linoleic acid (CLA) content of beef m. longissimus lumborum. Four groups of eight Charolais steers were given, for 120 days, grass silage plus a barley/sugar-beet feed concentrate containing one of four fat supplements: Megalac (saturated), linseed (high 18: 3),fish oil (high 20: 5 n-3, eicosapentenoic acid and 22: 6 n-3, docosahexaenoic acid) or linseed plus fish oil. The concentrates supplied 400 g/kg dry-matter (DM) intake and were designed to supply 45 g/kg of the total dietary fat calculated to be 60 g/kg of DM and to contain similar amounts of linoleic acid. Muscle from steers given the Megalac supplement contained 11·3 mg CLA per 100 g muscle and this was increased two- to three-fold in animals given the more unsaturated fat supplements. The increased deposition of CLA was similar for both linseed and fish oil supplements although the concentrations of total n-3 polyunsaturated fatty acids in the fish oil diet were much less than in the linseed diet. This suggests potent inhibition of conversion of CLA to trans vaccenic acid by fish oil fatty acids or their rumen metabolites.

Biohydrogenation and digestion of long chain fatty acids in steers fed on different sources of lipid

Scollan, N. D., Dhanoa, M. S., Choi, N.-J., Maeng, W. J., Enser, M., Wood, J. D. (2001). Biohydrogenation and digestion of long chain fatty acids in steers fed on different sources of lipid. Journal of Agricultural Science, 136, (3), 345-355

Effect of breed on the deposition in beef muscle and adipose tissue of dietary n-3 polyunsaturated fatty acids.

This study investigated the effects of dietary n-3 polyunsaturated fatty acids (PUFA) on muscle fatty acid composition and content in two cattle breeds; the Holstein-Friesian, a dairy breed, and the Welsh Black, a traditional beef breed. Animals were kept on ad libitum grass silage and one of three concentrates varying in the source of fat; Megalac (high in C16 : 0), formaldehyde treated whole linseed (high in C18 : 3 n-3) and a combination of formaldehyde treated whole linseed and fish oil (high in C20 : 5 n-3 and C22 : 6 n-3). The concentrates were offered at approximately 0·4 of total dry matter intake. Samples of m. longissimus thoracis from Holstein-Friesians contained higher total amounts of fatty acids ( P P P P P P

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

This paper compares the volatile compound and fatty acid compositions of grilled beef from Aberdeen Angus and Holstein-Friesian steers slaughtered at 14 months, each breed fed from 6 months on either cereal-based concentrates or grass silage. Linoleic acid levels were higher in the muscle of concentrates-fed animals, which in the cooked meat resulted in increased levels of several compounds formed from linoleic acid decomposition. Levels of α-linolenic acid, and hence some volatile compounds derived from this fatty acid, were higher in the meat from the silage-fed steers. 1-Octen-3-ol, hexanal, 2-pentylfuran, trimethylamine, cis- and trans-2-octene and 4,5-dimethyl-2-pentyl-3-oxazoline were over 3 times higher in the steaks from the concentrates-fed steers, while grass-derived 1-phytene was present at much higher levels in the beef from the silage-fed steers. Only slight effects of breed were observed.

As yet uncultured bacteria phylogenetically classified as Prevotella, Lachnospiraceae incertae sedis and unclassified Bacteroidales, Clostridiales and Ruminococcaceae may play a predominant role in ruminal biohydrogenation

Microbial biohydrogenation of dietary poly-unsaturated fatty acids (PUFA) to saturated fatty acids (SFA) in the rumen results in the high ratio of SFA/PUFA in ruminant products, such as meat and milk. In vitro, Butyrivibrio proteoclasticus-related bacteria extensively biohydrogenate PUFA to SFA, yet their contribution in the rumen has not been confirmed. The aim of this study was to evaluate the role of Butyrivibrio proteoclasticus group bacteria in ruminal biohydrogenation and to assess the possible role of other bacteria. Fish oil at 0%, 1.5% and 3% dry matter intake was fed to eight Holstein × Friesian steers, in order to elicit changes in the extent of PUFA biohydrogenation. Fatty acid and B. proteoclasticus group 16S rRNA concentrations in rumen digesta were determined. Correlation between digesta 18:0 concentration and B. proteoclasticus group 16S rRNA concentration was low. Terminal restriction fragment length polymorphism and denaturing gradient gel electrophoresis (DGGE) coupled with multivariate statistics revealed that many terminal restriction fragments (T-RFs) and DGGE bands were linked to cis-9, trans-11 conjugated linoleic acid (CLA), 18:1 trans-11 and 18:0 ruminal concentrations. MiCA T-RF predictive identification software showed that these linked T-RFs were likely to originate from as yet uncultured bacteria classified as Prevotella, Lachnospiraceae incertae sedis, and unclassified Bacteroidales, Clostridiales and Ruminococcaceae. Sequencing of linked DGGE bands also revealed that as yet uncultured bacteria classified as Prevotella, Anaerovoax (member of the Lachnospiraceae incertae sedis family), and unclassified Clostridiales and Ruminococcaceae may play a role in biohydrogenation.

Shelf life and eating quality of beef from cattle of different breeds given diets differing in n-3 polyunsaturated fatty acid composition.

Two experiments were conducted with 32 Charolais cross steers (experiment 1) and 36 Holstein-Friesian and Welsh Black steers (experiment 2) to examine the effects of dietary n-3 polyunsaturated fatty acids (PUFA ) and breed on meat quality. Diets were comprised of grass silage and a concentrate (60 : 40 on a dry-matter basis). Lipid in the concentrate provided 30 g/kg diet dry matter derived from either linseed, fish oil or linseed/fish oil (experiment 1); and linseed or linseed/fish oil (experiment 2). These n-3 PUFA sources were compared with Megalac, a saturated fatty acid control, in both experiments. Diets were offered for 120 days (experiment 1) and 90 days (experiment 2). Measurements of meat quality included fatty acid composition, lipid oxidation and colour during retail display (i.e. shelf life parameters) and eating quality assessed by a trained taste panel. The samples examined were sirloin (longissimus) steaks and minced beef burgers produced from the forequarter muscles infraspinatus, supraspinatus and triceps brachii. The fatty acid results showed that linseed increased the proportion of 18 : 3 n-3 (a -linolenic acid) in neutral lipid of the burgers and approximately doubled it in the phospholipid. Increased synthesis of 20 : 4 n-3 and 20 : 5 n-3 also occurred, raising their levels in the phospholipid. Fish oil doubled the proportions of 20 : 5 n-3 and 22 : 6 n-3, with the linseed/fish oil results intermediate between linseed and fish oil. The Welsh Blacks in experiment 2 had higher proportions of 18 : 3 n-3 in neutral lipid and higher proportions of 18 : 3 n-3, 20 : 5 n-3 and 22 : 5 n-3 in phospholipid. This showed a clear genetic effect on fatty acid composition. The fish oil diet in experiment 1 greatly increased lipid oxidation during retail display in both overwrapped loin steaks and burgers packed in a modified atmosphere. Values for lipid oxidation in the burgers were much higher than in the steaks due to the intimate mixing of phospholipid fatty acids and prooxidants. Lipid oxidation was also greater in the linseed/fish oil samples than controls in both experiments but was similar in linseed and controls. Colour oxidation also proceeded more rapidly in the fish oil samples in experiment 1 and tended to be faster in the linseed/fish oil groups in both experiments. Eating quality was affected by diet in experiment 1 — the fish oil samples eliciting higher scores for ‘fishy’ and ‘rancid’ than controls and having lower ‘overall liking’ scores. Steaks from animals given the linseed diet (with high 18 : 3 n-3 proportions) had the highest ‘overall liking’ scores in experiment 1 but the cattle given linseed for the shorter time in experiment 2 were not different from controls. Despite their different PUFA profile in muscle, Welsh Blacks had similar eating quality scores to Holstein-Friesians which deposited more neutral lipid (marbling fat) in muscle.

The effect of clover silages on long chain fatty acid rumen transformations and digestion in beef steers

Lee, M. R. F., Harris, L. J., Dewhurst, R. J., Merry, R., Scollan, N. D. (2003). The effect of clover silages on long chain fatty acid rumen transformations and digestion in beef steers. Animal Science, 76, (3), 491-501.