D.W. Pethick

Intramuscular fat content in meat-producing animals: development, genetic and nutritional control, and identification of putative markers.

Intramuscular fat (IMF) content plays a key role in various quality traits of meat. IMF content varies between species, between breeds and between muscle types in the same breed. Other factors are involved in the variation of IMF content in animals, including gender, age and feeding. Variability in IMF content is mainly linked to the number and size of intramuscular adipocytes. The accretion rate of IMF depends on the muscle growth rate. For instance, animals having a high muscularity with a high glycolytic activity display a reduced development of IMF. This suggests that muscle cells and adipocytes interplay during growth. In addition, early events that influence adipogenesis inside the muscle (i.e proliferation and differentiation of adipose cells, the connective structure embedding adipocytes) might be involved in interindividual differences in IMF content. Increasing muscularity will also dilute the final fat content of muscle. At the metabolic level, IMF content results from the balance between uptake, synthesis and degradation of triacylglycerols, which involve many metabolic pathways in both adipocytes and myofibres. Various experiments revealed an association between IMF level and the muscle content in adipocyte-type fatty acid-binding protein, the activities of oxidative enzymes, or the delta-6-desaturase level; however, other studies failed to confirm such relationships. This might be due to the importance of fatty acid fluxes that is likely to be responsible for variability in IMF content during the postnatal period rather than the control of one single pathway. This is evident in the muscle of most fish species in which triacylglycerol synthesis is almost zero. Genetic approaches for increasing IMF have been focused on live animal ultrasound to derive estimated breeding values. More recently, efforts have concentrated on discovering DNA markers that change the distribution of fat in the body (i.e. towards IMF at the expense of the carcass fatness). Thanks to the exhaustive nature of genomics (transcriptomics and proteomics), our knowledge on fat accumulation in muscles is now being underpinned. Metabolic specificities of intramuscular adipocytes have also been demonstrated, as compared to other depots. Nutritional manipulation of IMF independently from body fat depots has proved to be more difficult to achieve than genetic strategies to have lipid deposition dependent of adipose tissue location. In addition, the biological mechanisms that explain the variability of IMF content differ between genetic and nutritional factors. The nutritional regulation of IMF also differs between ruminants, monogastrics and fish due to their digestive and nutritional particularities.

Nutritional and hormonal regulation of energy metabolism in skeletal muscles of meat-producing animals

In meat-producing animals, skeletal muscle is a tissue of major economic importance. Energy metabolism of skeletal muscle should be assessed in relation to its main physiological functions which are growth as well as posture, physical activity and thermogenesis. These functions have specific energy requirements which may entail competition in the utilization of energy-yielding nutrients, and modify the efficiency of ATP production and use. In this review, the energy requirements of muscles will be presented as well as the main metabolic pathways involved in the utilization of carbohydrate and lipids for energy supply and deposition. Regulation of metabolism will also be tackled in relation to the major physiological functions of skeletal muscle in various domestic species. Special focus will be given to the energy metabolism of growing muscle and its interactions with meat quality. The question will be asked whether the optimization of the efficiency of ATP production and utilization in muscle for nitrogen retention is compatible with an improvement of meat quality (as partly determined by glycogen and lipid deposition)? For this purpose, the interactions between muscle carbohydrate, lipid and protein metabolism will be reviewed.

Effects of dietary factors and other metabolic modifiers on quality and nutritional value of meat

A number of technologies that increase feed efficiency and lean tissue deposition while decreasing fat deposition have been developed in an effort to improve profitability of animal production. In general, the mode of action of these metabolic modifiers is to increase muscle deposition while often simultaneously reducing fat deposition. However, there have been some concerns that the focus on increasing production efficiency and lean meat yield has been to the detriment of meat quality. The aim of this review is to collate data on the effects of these metabolic modifiers on meat quality, and then discuss these overall effects. When data from the literature are collated and subject to meta-analyses it appears that conservative use of each of these technologies will result in a 5-10% (0.3-0.5kg) increase in shear force with a similar reduction in perception of tenderness. However, it should be borne in mind that the magnitude of these increases are similar to those observed with similar increases in carcass leanness obtained through other means (e.g. nutritional, genetic selection) and may be an inherent consequence of the production of leaner meat. To counter this, there are some other metabolic factors and dietary additives that offer some potential to improve meat quality (for example immuncastration) and it is possible that these can be used on their own or in conjunction with somatotropin, approved β-agonists, anabolic implants and CLA to maintain or improve meat quality.

Increasing viscosity of the intestinal contents alters small intestinal structure and intestinal growth, and stimulates proliferation of enterotoxigenic Escherichia coli in newly-weaned pigs

Sources of viscous soluble fibre, such as barley and oats, have often been included in the weaning diet of the pig to accelerate development of the large intestine. Inclusion of a non-fermentable, viscous compound, sodium carboxymethylcellulose (CMC), in a low-fibre weaning diet was tested to assess the influence of digesta viscosity on the gut in the absence of increased fermentation. Two CMC sources, of low and high viscosity, were added to cooked rice-based diet at 40 g/kg total diet. A third control rice diet did not contain any CMC. Diets were fed for 13 d following weaning at 3 weeks of age. Addition of CMC to the diet significantly increased the intestinal viscosity of digesta within the small (P<0.001) and large (P<0.05) intestine. No simple association was found between increases in intestinal viscosity and effects on intestinal morphology and whole-body growth. The average empty-body-weight gain and the small intestinal villus height increased with low-viscosity CMC, but decreased with the high-viscosity CMC group. The full large intestinal weight increased in all pigs fed CMC. Dietary CMC (both low- and high-viscosity) increased the percentage moisture of digesta and faeces, and was associated with increased faecal shedding of enterotoxigenic haemolytic Escherichia coli. Feed ingredients in weaning diets that excessively increase the viscosity of the intestinal digesta may be detrimental to pig health and production.

Growth, development and nutritional manipulation of marbling in cattle: a review

This review describes the pattern of intramuscular fat accretion in cattle and the potential for its manipulation during both the pasture (or backgrounding) and intensive grain-finishing phases of development. A growth curve for the development of marbling in British and Japanese Black type breeds is discussed with the conclusion that 3 phases of development exist: (i) a period of growth up to ∼200 kg hot carcass weight where intramuscular fat does not increase; (ii) a period of linear development as carcass weight increases from 200 to 450 kg; and (iii) the attainment of mature body size (∼500 kg carcass weight depending on genotype) at which intramuscular fat content appears to reachea maximum. Data are also presented to show that the intramuscular and other fat depots develop at similar rates indicating that intramuscular fat is not a late maturing depot. Pre-finishing growth checks reduce the initial intramuscular fat at the start of finishing and this is translated into lower levels at the end of finishing. It is argued that the greatest potential for the manipulation of intramuscular fat accretion during fattening is via an increase in the net energy of the ration. Increasing net energy can be achieved by increasing the cereal grain content of the diet (grain v. grass); by feeding processed cereal grain, which allows both maximal rumen fermentation and small intestinal digestion of starch, and by increasing the lipid content of the diet. In addition it is proposed that the substrate supply or hormonal milieu can also be optimised, along with the availability of net energy to maximise fat accretion. The role of lipolysis (fat turnover) as a regulator of fat accretion is also discussed.

Genetic and environmental effects on meat quality

In order for livestock industries to consistently produce high quality meat, there must be an understanding of the factors that cause quality to vary, as well as the contribution of genetics. A brief overview of meat tenderness is presented to understand how genotype and environment may interact to influence this trait. Essentially, meat tenderness is determined from the contribution of connective tissue, sarcomere length determined pre-rigor and rate of proteolysis during ageing, as well as contributions from intramuscular fat and post-mortem energy metabolism. The influence of mutations in myostatin, the callipyge gene, the Carwell or rib eye muscle gene as well as the calpain system on meat tenderness is presented. Specific examples of interactions between the production or processing environment and genetics are presented for both sheep and cattle. The day-to-day variation in tenderness is evident across experiments and this variation needs to be controlled in order to consistently produce tender meat.

Relationship between animal age, intramuscular fat, cooking loss, pH, shear force and eating quality of aged meat from sheep

The relationships between sensory traits (tenderness, juiciness, flavour and overall liking) and objective measures, such as shear force, intramuscular fat, cooking loss, pH and animal age, were derived for M. longissimus thoracis et lumborum (LL) from 471 lamb and sheep carcasses. Tenderness could be predicted with the most accuracy (R2 = 0.24) and flavour with the highest precision (r.s.d. = 7.5 units) when using the objective measures, which may be in part due to the small variation in the range of shear force values of the samples (all carcasses electrically stimulated and meat aged for 5 days) and the use of consumer panels for the assessment of sensory traits. The ultimate pH of the LL, the rate of decline in pH in the LL or the predicted temperature at pH 6.0 were not significant predictors of the sensory traits when tested on a subsample of the carcasses. The model coefficients indicated that all sensory traits (tenderness, flavour, juiciness and overall liking) declined as shear force and age increased, and as intramuscular fat percentage decreased. This translated into a decline of 16 points on average for tenderness and 13 points for overall liking when LL samples from 68.5-month-old sheep were compared with those from unweaned lambs, when adjusted to the same level of intramuscular fat and shear force. Predictions of the sensory traits at varying levels of shear force were made and show that at 49 Newtons (N), the overall liking score would be 51 and the tenderness score 48. Derived relationships between objective meat quality measures and sensory traits suggest that to achieve a failure rate of no more than 10% for loin meat when eaten, it must have a shear force of about 27 N or less.

Acetate supply and utilization by the tissues of sheep in vivo

1. The supply and utilisation of acetate has been estimated simultaneously in the whole animal and tissues of sheep using a combination is isotope-dilution and arteriovenous-difference techniques. Animals were made alloxani-diabetic and acetate metabolism was compared when stabilized to normal metabolite levels with insulin (ITA sheep) and when food and insulin had been withdrawn for 36 h (fasted, diabetic sheep). 2. Acetate was simultaneously produced and utilized by all tissues. The exogenous (or gut) supply of acetate was the most important determinant of circulating acetate level. Endogenous acetate was produced mainly in the liver; 77 and 94% in fasted, diabetic and ITA sheep respectively. The production of endogenous acetate remained fairly constant and was not related to ketogenesis, which supports the idea that circulating acetate is largely a produce of fermentation. The liver, gut and muscle utilized 17, 25 and 54% respectively (96% total) of the acetate entry rate in ITA sheep; a similar percentage utilization was found in fasted, diabetic sheep. 3. Acetate is largely oxidized to carbon dioxide in the gut and muscles of sheep and may account for 30-40% of their oxidative metabolism. This figure is similar to that for the whole animal. The total acetate taken up by the liver could account for 30% of the oxygen consumption; however, the liver may not directly oxidize all the utilized acetate. 4. The over-all conclusion from this study is that acetate is largely of dietary origin and the major factor determining its rate of utilization is the arterial concentration.

The impact of processing on sensory and objective measurements of sheep meat eating quality

The impact of processing factors on sheep meat eating quality was investigated in an experiment in which stimulation (stimulation and no stimulation), chilling rate (fast and slow chilling) and carcass suspension (tenderstretch and normally hung by Achilles tendon) treatments were overlaid on 80 lamb and 40 mutton carcasses processed over 2 days, at 2 different abattoirs. Within each carcass, 3 muscles (M. longissimus thoracics et lumborum, biceps femoris and serratus ventralis) were collected from both sides and aged for 2 of 3 ageing periods (2, 5 or 14 days), before sensory testing for tenderness, juiciness, like flavour and overall liking, using a consumer taste panel. Processing treatments of stimulation and chilling were variable in their effect on the rates of pH and temperature decline and temperature at pH 6 (temp@pH6), between the 4 slaughter groups. Therefore chilling rate and electrical stimulation were considered as tools by which temp@pH6 could be manipulated, rather than as treatment effects per se. Age category (lamb or mutton) had the largest impact on tenderness score, followed by muscle, ageing and carcass suspension. There were significant interactions for tenderness and overall liking scores between muscle × ageing, age category × muscle and carcass suspension × muscle (P 30°C) or low (<10°C) temp@pH6. There was a tendency for sensory scores (particularly for the like flavour score) to decline for the biceps femoris when it was stored for extended periods (14 days).

How might marbling begin

Marbling is an important meat quality trait, in that it contributes directly to the value of beef on international markets. The development of marbling is not well understood, though there have been some significant recent discoveries regarding adipogenesis in general. This article describes a working hypothesis around the early events of marbling. It attempts to rationalise findings from several mammalian experimental systems on hyperplastic growth of adipocyte precursor cells.