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Improving the Sensory and Nutritional Quality of Fresh Meat

Improving the Sensory and Nutritional Quality of Fresh Meat

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Improving the Sensory and Nutritional Quality of Fresh Meat

1,487 pages
17 heures
Jan 22, 2009


Understanding of the scientific basis of quality attributes in meat is becoming more advanced, providing more effective approaches to the control of meat eating and technological quality. This important collection reviews essential knowledge of the mechanisms underlying quality characteristics and methods to improve meat sensory and nutritional quality.

Part one analyses the scientific basis of meat quality attributes, such as texture and tenderness, colour, water-holding capacity and flavour development. Chapters on the nutritional quality of meat and meat sensory evaluation complete the section. Part two discusses significant insights into the biology of meat quality obtained from genomic and proteomic perspectives, with chapters focussing on different types of meat. Parts three and four then review production and processing strategies to optimise meat quality, considering aspects such as production practices and meat nutritional quality, dietary antioxidants and antimicrobials, carcass interventions, chilling and freezing and packaging. Methods of meat grading and quality analysis are also included.

With its distinguished editors and international team of contributors, Improving the sensory and nutritional quality of fresh meat is a standard reference for those industrialists and academics interested in optimising meat quality.
  • Reviews methods to improve meat sensory and nutritional quality considering the effects of different production practices such as chilling, freezing and packaging
  • Analyses the scientific basis of meat quality attributes covering texture, tenderness, colour and water-holding capacity
  • Examines production and processing strategies to optimise meat quality, including the current state of development and future potential
Jan 22, 2009

Lié à Improving the Sensory and Nutritional Quality of Fresh Meat

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Improving the Sensory and Nutritional Quality of Fresh Meat - Elsevier Science


Part I

Understanding meat quality


Chapter 1: Trends in meat consumption and the need for fresh meat and meat products of improved quality

Chapter 2: Biology and regulation of carcass composition

Chapter 3: Fresh meat texture and tenderness

Chapter 4: Meat color

Chapter 5: Flavour development in meat

Chapter 6: Fresh meat water-holding capacity

Chapter 7: The nutritional quality of meat

Chapter 8: Sensory evaluation of fresh meat


Trends in meat consumption and the need for fresh meat and meat products of improved quality

M.D. Aaslyng,     Danish Meat Research Institute, Denmark


Meat is one of our main food items and most recipes are named after the meat ingredients. During the last fifty years, meat consumption has increased by aproximately 50% in both USA and Europe. Meat is eaten due to its culinary status (we like meat) but also due to its nutritional value. This chapter focuses on the meat consumption pattern and on the factors important for both the gastronomic and nutritional value of meat.

Key words

meat consumption

eating quality

nutritional quality





1.1 Introduction

Meat is one of our main food items. Looking in a cook book, most recipes are named after the meat ingredients and meat is in general seen as a central part of the meal (Horowitz, 2006). Already by the Middle Ages, eating meat was seen as a sign of prosperity and wealth (Anon, 2007), and following the increased wealth in the community, the amount of meat eaten increased (Horowitz, 2006). From the 1950s to the 1990s meat consumption per inhabitant increased by approximately 50% in both the USA and Europe (Fagt et al., 1992; Horowitz, 2006), and also in the last ten years increasing meat consumption has been seen (Breadsworth et al., 2004). In the southern part of Europe, meat has now overtaken fish as the main protein source and actually more meat is eaten today in countries around the Mediterranean compared with the northern part of Europe, which by tradition used to be heavily meat eating (Marques-Vidal et al., 2006; Naska et al., 2006). Even though some people regard meat negatively (Holm et al. 2000) 98–99% of the population eat meat regularly (Garemo et al., 2007; Guenther et al., 2005), even though newer surveys show a gradual decrease in meat consumption in the last years, at least in Denmark (Fagt et al., 2002).

The main meat types are beef, poultry (chicken and turkey), and pork. In an investigation in the USA of what kind of meat was eaten in a two-day period in 1994–1996, beef was the most frequently eaten meat, followed by chicken, with pork the least popular. However, increase in meat consumption in the USA is most noticeably an increase in consumption of chicken (Sloan, 2003), and today this type of meat might be the most popular. Also, in Norway the consumers prefer white to red meat (Kubberød et al., 2002a).

However, behind these facts about meat consumption, variations are present. It is reported in many countries that boys and men eat more meat compared with girls and women (Cosgrove et al., 2005; Guether et al., 2005; Lyhne, 2005). Also the type of meat that the two genders prefer, differ. Men prefer red meat while women eat more white meat and even take a dislike to red meat (Kubberød et al., 2002a, 2006). Also the attitude towards meat is more positive for men compared with women (Kubberød et al., 2002b).

An age difference in meat consumption pattern also exists. Young consumers eat less meat and prefer, to a large degree, white meat to red meat compared with more adult consumers (Aaslyng et al., 2007b; Koizumi et al., 2001; Kubberød et al., 2002a); and families with children eat more meat than families without children (Groth et al., 2003). It also seems that more educated consumers eat less meat (Groth et al., 2003). In general, young well-educated women are among the least meat-eating consumers while adult, less educated men are among the most meat-eating.

The cutting of the meat has an influence on its popularity. Minced meat is often preferred, as the chance of success in cooking is high (Holm et al., 2000). A Swedish survey showed that 98% of children eat minced meat at least once a week (Garemo et al., 2007). In Denmark, a geographic difference is seen regarding which form of pork is mostly eaten. In a city close to the capital, whole meat pieces, such as chops or roasts, are often bought, while far from the capital, minced meat is more often chosen (unpublished data).

Which meat-based meal is the most popular differs during a lifetime. In Sweden, 83% of children had sausages at least once a week (Garemo et al., 2007), and also in Denmark sausages are mainly eaten by children. Figure 1.1 shows the popularity of five meals, depending on the consumer’s age. The results are from a survey in which consumers indicated which meals they had eaten in the past week. Please note that this procedure (having more pork meals than chicken meals to choose between) means that chicken seems most popular, even though pork is eaten more often.

Fig. 1.1 The popularity of five selected meals in Denmark (National Danish Dietary Survey, 2000–2002; Fagt, Pers. Comm., 2007).

Beef bolognaise and other stews, and chicken, are the two most popular meals, independent of age. Sausages are very popular up to 14 years of age, at which point their popularity drops dramatically. However, from 25–34 years they regain their popularity and are the fourth most popular meal. This corresponds to the period in which many people have small children – who eat sausages! Whole beef roasts and steaks are most popular with consumers from 35–65 years old, after which they drop again. This last decrease can be due either to a reduced chewing capacity or to a reduction in income and thereby in the amount of disposable money for buying food.

The way in which meals with meat are composed can be either very traditional or more varied. When pork is served as a whole roast in Denmark, the accompaniments are often very traditional, including potatoes, gravy, pickled red cabbage and occasionally salad. In contrast, pork chops are served with a much larger variety of accompaniments. The basic accompaniment is mainly potatoes and sometimes rice, while pasta is seldom served. In addition to this, salad, bread and a large variety of vegetables both hot and cold are served with the chops (Aaslyng et al., 2007a). The same pattern seen for chops has been seen for beef steaks. They are most often served with potatoes, combined with a large variety of vegetables and salad (unpublished data).

In this way, some cuts seem much more traditional in the way they are served compared with others, in which the serving can be more varied. It is not established how these traditions are kept or broken. Who decides what is trendy? Chefs experiment with meals – not only the cooking of the meat, in which trends using low cooking temperatures for long times, often combined with a low internal temperature, have arrived in the last years – but also in the way the whole meal is composed. However, these trends are seldom converted directly to ordinary consumers. Meals prepared in the home are more focused on taste, convenience, health and nutrition. The meals must be easy to shop for and easy to prepare after a long day at work. Also, a high success rate in cooking is important as we all prefer food of good taste. To achieve this, well-known recipes are often chosen and inspirations for new meals are sought from cook books and food magazines. These meals are often made for special occasions and, if they are a success, transferred fully or partly to everyday meals. (Andersen, Pers. Comm., 2007).

But why do we eat meat and what are the trends in meat-eating habits? These questions are important in order to understand the consumption of this food group. We can eat food for several reasons. Pleasure (we actually like meat) and nutrition (meat is a good protein source with several nutritionally beneficial factors) are two of the main reasons for including meat in meals.

1.2 Eating meat for pleasure

Eating meat for pleasure means we like meat. For steaks, chops or roasts, three sensory attributes are of major importance for the hedonic value of the meat: Tenderness, juiciness, and flavour (both the presence of fried flavour and absence of off-flavours) (Aaslyng et al., 2007b; Bryhni et al., 2003; Miller et al., 2001). Which attribute is considered the most important depends on factors such as geographic location (Aaslyng et al., 2007b) and on the level of the other attributes (Huffman et al., 1996; Killinger et al., 2004). With tender meat, variations in juiciness or flavour have a larger impact on preference compared with less tender meat, in which increased tenderness is of more importance (Miller et al., 2001). However, these three parameters can only be assessed on cooked meat. When buying meat, we have to guess the eating quality from the appearance. The appearance of the raw meat is therefore very important for the consumer.

1.2.1 Appearance

The appearance of meat at the point of buying is important for choice. For pork, colour and subcutanous fat (fat layer) are the two main drivers of acceptability. The consumer associates colour with freshness while a thin fat layer is associated with health. Overall, both dark and light meat colour can be preferred, depending on the country and consumer segment within the country (Ngapo et al., 2007). As an example, most consumers in Australia and Poland prefer light coloured pork chops while the opposite is true for Germany and Taiwan. For all countries, a low fat cover is preferred by most consumers. High marbling of pork chops was only important for most consumers in Australia and Ireland. For the rest of the 21 investigated countries, marbling was not important for choice of chop (Ngapo et al., 2007). In beef, it has been seen that, at the moment of purchase, consumers prefer steaks with low marbling, but when actually tasting the meat, they prefer steaks with a higher degree of marbling (Jeremiah et al., 1992; Madsen, Pers. Comm., 2007)

1.2.2 Tenderness

Tenderness has often been described as the most important factor for high eating quality, especially in beef. Consumers can distinguish between tender and tough beef steaks, whereas variations in tenderness in the middle groups are more difficult to separate (Sivertsen et al., 2002). It has also been shown that a certain level of tenderness is crucial to be acceptable (Huffman et al., 1996) and that tenderness of beef is so important for consumers, they are actually willing to pay more for tenderness (Boleman et al., 1997). An increased preference with increasing tenderness has also been reported in pork (Aaslyng et al., 2007b).

Tenderness of meat can be due to many different factors and there are many ways to achieve tenderness (Aaslyng et al., 2004a). Ageing is a well known way of increasing tenderness of meat. During ageing, proteolytic enzymes degrade the proteins and loosen up the structure of the meat. This can be seen by an increased Myofibrillar Fragmenting Index or increased degradation of specific proteins such as the cytoskeletal protein desmin (Kristensen et al., 2003, 2006; Therkildsen et al., 2002a). The effect of ageing depends on the time and temperature during ageing, and also on the growth rate of the animal up to slaughter, as a fast growthrate increases the activity of proteolytic enzymes in the meat (Kristensen et al., 2006; Therkildsen, 1999; Therkildsen et al., 2002a,b).

Also intramuscular fat (IMF) is an important factor in tenderness (Aaslyng et al., 2004b; Brewer et al., 2001; Fortin et al., 2005). The increase in tenderness with increasing IMF in pork is not linear but can more be seen as a limit at between 1.5–3% (depending on the country), below which the tenderness is regarded as unacceptable (Fortin et al., 2005). In beef, a ‘window of acceptability’ has been defined as at least 3% being necessary to gain acceptable palatability, at least 5% to get a medium palatability, while an IMF content of at least 7% is necessary for a high palatability (Savell et al., 1988).

The sarcomere length can vary depending on the chilling regime and on the suspension of the hot carcass. A short sarcomere length induces tough meat. In pork, a linear relationship between sarcomere length and tenderness exists for sarcomeres below 2 μm, independent of muscle. Above 2 μm sarcomere length, the meat is always regarded as tender (Wheeler et al., 2000).

A popular way to increase tenderness is to marinate the meat, using a solution with NaCl or CaCl2, perhaps including spices, sugars or other ingredients. Injecting this into the meat increases tenderness, juiciness and flavour. This is especially popular in USA, but has also become more widespread in Europe. In beef, a linear relationship has been demonstrated, with decreasing shear force (corresponding to increasing tenderness) as salt concentration is increased (Baublits et al., 2006); and marinating of a potentially non-tender muscle such as knuckle (M. rectur femoris) can increase the tenderness up to the level of a non-marinated loin (Rosenvold et al., 2006). For a meat producer, it is important to ensure high tenderness to obtain meat of high acceptability. Which is the best way to gain tenderness (IMF, ageing, chilling) must be chosen from a knowledge of the individual production systems.

1.2.3 Flavour

Flavour is a very important eating quality attribute in all meats. Not only are the preferred flavours important – it is just as, or even more, important to avoid off-flavours.

Fried flavour is generated during heat treatment at high temperatures. The aroma compounds are formed by several pathways, e.g. lipid oxidation and Maillard reaction between reducing sugars and amino acids; and the reactions between compounds from these two systems results in a large number of volatile compounds (Mottram, 1994). Thermal degradation of other compounds, such as thiamin, also contributes to the flavour of meat (Tai et al., 1999). Non-volatile compounds, such as inosine monophosphate (IMP) and its degradation products, free amino acids, smaller peptides and lactic acid and ions, also have an impact on flavour (Fuke et al., 1991; Maga, 1994; Nishimura et al., 1988; Tikk et al., 2006). Even though flavour is important for preference (Aaslyng et al., 2007b), the consumers willingness to pay for improved flavour has not been investigated as thoroughly as for the tenderness.

Increasing IMF, to a degree, increases the fried flavour of both pork and beef (Brewer et al., 2001; Killinger et al., 2004). The main contributors to meaty volatiles are the phospholipids (Mottram et al., 1983) and the content of this fraction is constant, as they are the structural lipids present in the membrane of muscle cells. The effect of IMF on flavour could therefore be due not only to increased flavour development but also to improved flavour release. This agrees with findings showing only minor effects of fatty acid composition on the flavour of pork and beef (Elmore et al., 1999; Tikk et al., 2007; Wood et al., 2003) even though some fatty acids, especially omega-3-fatty acids, can induce off-flavours if present at too high concentrations (Aaslyng et al., 2007c; Elmore et al., 1999).

As the Maillard reaction is essential for developing fried flavour during heat treatment, the content of carbohydrates and amino acids seems to be crucial. Meat has a high protein content and amino acids should be present in large amounts. Carbohydrates are therefore more likely to be the limiting factor. Investigations have therefore focused more and more on the content of carbohydrates in the meat (Aliani et al., 2005; Koutsidis et al., 2007; Meinert et al., 2006, 2007).

Several off-flavours have been reported. In pork, the boar taint is crucial as also is piggy flavour (Hansen et al., 2006; Meinert et al., 2006), sow flavour (Sindelar et al., 2003a,b) and sometimes simply off flavours/abnormal flavours (Wood et al., 1995, 2004). In beef and veal especially, livery flavours are described (Yancey et al., 2006), but also metallic, cowy, milky, grassy, painty and sour flavours are mentioned (Calkins et al., 2007).

1.2.4 Juiciness

Juiciness is said to be an important factor in the eating quality of meat. How important it is depends, to a large degree, on the meal composition, as a steak needs to be juicier than small slices in a stew. However, to get a tasty piece of meat requires some meat juice.

The main factor determining the juiciness of meat is the end-point temperature. The higher the end-point temperature, the higher is the cooking loss and the lower the juiciness (Aaslyng et al., 2003). The connection between cooking loss and juiciness is not simple. In pork, meat of low pH had a higher cooking loss than meat of normal pH but this difference was not reflected in the juiciness, as they were equally juicy (Aaslyng et al., 2003).

Increased amounts of intramuscular fat also increased juiciness, but only when the meat was fried to a high end-point temperature. At lower end-point temperatures, at which the meat was juicier, intramuscular fat had no effect on juiciness (Aaslyng et al., 2004b).

To increase juiciness of meat, the most important factor must be to educate consumers not to over-cook the meat. This can be done by advising consumers on cooking times (http://www.danishmeat.dk) or teaching them to use a frying thermometer (Edwards et al., 2005) to ensure a safe, well tasting meal.

1.3 Eating meat for nutrition

Meat is an important nutritional source in the diet of most people. It contains a high amount of protein, including essential amino acids, and other beneficial nutrients such as iron and vitamins. But it has some negative effects. Meat has a high content of saturated fatty acids, of which we need to reduce the intake to minimize the adverse effects of cardiovascular disease and the risk of cancer (NNF, 2004).

1.3.1 Protein

Eating meat means eating a huge amount of protein. Protein is an important nutrient, but in the western world lack of protein is not a problem. Investigations have shown that a diet high in protein can induce weight loss, perhaps because it is more satisfying compared with a carbohydrate-rich diet, so the intake of energy is less (Richelsen, 2006). In a world with increasing problems with obesity, this effect could be important. A EU-project – DIOGENES (Diet, Obesity and Genes) – is currently investigating these relationships (http://www.diogenes-eu.org/).

1.3.2 Fat content and fatty acid composition

We are advised to reduce the amount of fat we eat, especially saturated fat (NNF, 2004). Fat from animal sources (milk, meat, egg) is very saturated and the authorities in many countries therefore advise the population to reduce the intake of animal fat. This has been taken in by consumers, as most now prefer meat without visible fat.

Meat contains variable amounts of fat, depending on the source (chicken, beef, and pork) and the cutting (Danish Meat Association, 2007). A large amount of fat is visible, either between muscles (intermuscular fat) or as the fat layer (subcutaneous fat), and can be removed by the industry before sale or by the consumer before or after cooking. Most meat types are actually very lean and the amount of intramuscular fat should not be reduced too much as this will reduce the eating quality (Aaslyng et al., 2004b). Many consumers regard chicken as the most lean meat type (Andersen, Pers. Comm., 2007), but this is not strictly true. Pork and veal can be just as lean after removing the visible fat.

In minced meat, it is more difficult for the consumer to remove the fat, as it is mixed into the meat. The industry has therefore a responsibility to offer consumers lean minced meat as an option in the shops.

Not only the amount of fat, but also the fat quality is important with regard to health. The fatty acid composition of meat depends on the feed of the animal. Pork and chicken (monogastric animals) easily reflect the fatty acid composition of the feed in the fat (Affentranger et al., 1996; Gatlin et al., 2002; Nürnberg et al., 1999; Tikk et al., 2007), but also the fatty acid composition of beef can vary with varying feed (Elmore et al., 1999, 2004). Table 1.1 shows the general fatty acid composition of pork, beef and chicken.

Table 1.1

Fatty acid composition (%) in pork, beef and chicken. (Fødevaredatabanken version 6, Danmarks Fødevareforskning).

*This percentage is for both lean and fat meat cuts even though a lean piece of meat will contain a higher percent phospholipids and thereby less saturated fat compared with a fattier piece of meat.

The fatty acids of beef are more saturated than the fatty acids of pork and chicken. In comparison, the relationship between omega-3 and omega-6 fatty acids are better in pork and chicken as the proportions are below 4, as is often recommended (NNF, 2004). The proportion between omega-3 and omega-6 fatty acids in pork depends to a large extend on the feed, and ratios as low as 1.4 have been reported (Nuernberg et al., 2005).

It is possible to increase the content of omega-3 fatty acids by feeding pigs with, e.g. linseed oil (Nürnberg et al., 1999), and in France and Canada pork and pork products have been marketed with a health claim for a high content of omega-3 fatty acids. This is one way of meeting consumer demand for healthy meat products.

1.3.3 Minerals and vitamins

Iron is a main mineral in our body as it is essential in the transportation of oxygen around the body. Iron deficiency is well known, especially in young women (Samuelson et al., 2003). Meat is an excellent source of iron as it contains high amounts of the easily absorbable haem iron. At the same time a ‘meat factor’ is present as meat seems to enhance the absorption of non-haem iron (Bæch et al., 2003a,b). Even small amounts – ≥50 g a day – of meat enhance the iron uptake from food in general (Bæch et al., 2003a).

Selenium is a micronutrient necessary for some antioxidative enzymes. Pork has been shown to be an excellent source of selenium (Skibsted, 1997). The reported content of selenium in meat varies between countries, perhaps due to variation in the method of analysis (Strong, 2006).

The intake of D-vitamin is too low in many countries, and even though it can be synthesized by exposure to the sun, a higher intake through the diet is desirable. Meat contributes about 26% of the D-vitamin intake in a normal diet (Lyhne et al., 2005). D-vitamin is present in the fatty part of the meat and also in the rind of pork (Fødevaredatabasen, 2007). It is not known if exposure to sun of the live animal enhances the concentration of D-vitamin in the rind.

Sodium chloride is a salt often used in meat products for its taste and functionality, and for its effect of prolonging the shelf-life of a product. However, it is recommended that we reduce our intake of sodium-ion substantially (Loria et al., 2001; NNF, 2004). Meat products are responsible for a relatively high amount of our intake of sodium (NNF, 2004), and decreased use of sodium by the meat industry is therefore desirable. Countries such as Finland and UK have already worked for some time on mapping routes for reducing sodium chloride without reducing eating quality or shelf-life, and other parts of Europe and North America have now also started focusing on reducing sodium.

1.4 Variability in meat and meat products

Meat is a biological source, and natural biological variations between animals will always exist. How can a consumer be certain to get what he wants? Is there a need for improved quality?

A steak or a chop needs to be fried for a certain time depending on how thick it is. To increase the success rate for the consumer when frying the meat, the slices in a package needs to be similar in size. This sounds easy, but requires good workmanship from the industry.

Tenderness is, as stated in the introduction, an important factor for consumer satisfaction and possibilities abound to increase tenderness. Marbling is an important factor in both beef and pork, but again a large variation exists in the degree of marbling both between animals and within the same muscle (Aaslyng, 2002). Also, other factors can influence tenderness, such as the age of the animal (especially beef) and the chilling regime. The effect of these factors can not be seen on the raw meat and branding is therefore necessary to help the consumer to choose tender meat. The branding could be built upon a combination of sorting the carcasses to get marbled meat, of having a code of practice to ensure optimal chilling or ageing of the meat to increase tenderness. This aspect is discussed in more depth in Chapter 3 of this book.

To produce healthy products, the variation in the nutritional quality of the meat must not be too high. Feeding is known to alter the fatty acid content and distribution in the meat and restriction of the fat in the feed can therefore be relevant if the focus is on healthy products. This aspect is discussed in depth in Chapters 7 and 15 of this book.

Also, the technological quality has a natural variation. In pork, the pH variation in a survey of five Belgian slaughterhouses was between 5.56 ± 0.03 and 5.76 ± 0.04 (Lammens et al., 2007). This is reflected in a variation in drip loss and colour. Gentle handling of the pigs up to slaughter (Støier et al., 2001) combined with rapid chilling (Maribo et al., 1998), can improve the technological quality and reduce the natural variation. In beef, technological quality is reflected in the pH. The pH is measured the day after slaughter and is used as a quality attribute to screen out carcasses of low quality.

When producing meat products, other factors can be of relevance to obtain a high quality compared with those measured in fresh meat. To produce dry-cured ham products from pork, proteolytic enzyme activity is very important. This can, however, vary between animals depending on the growth rate up to slaughter (Kristensen et al., 2003; Therkildsen et al., 2002a,b). To produce bacon, the distribution between fat and meat influences the quality of the product, and variations are also seen here. To overcome this, sorting on fatness is a possibility. In the production of sausages, raw meat quality variation has an influence, especially the fatty acid composition. Too high a degree of unsaturation might be nutritionally beneficial, but the quality of the sausages can be reduced. The fatty acid composition depends on the feeding and also on the location of the fat in the carcass. The problem can be overcome either by restrictive feeding of the animals or by combining fat sources with different degrees of saturation.

The natural variation between animals is thus a challenge for the industry, requiring a large degree of sorting or combination of raw materials to gain the optimum quality. On the other hand, it is also a possibility to take advantage of the variation to increase the quality by choosing the optimum for each product and to meet market demand.

1.5 Future trends

The meat industry has a challenge in the future to meet consumers’ demand for healthy products with high eating quality. How can the consumers and the industry meet? First of all the consumers demands must be known. Not just the demands of today, but also the expected demands in the future.

When buying meat, the consumer must either guess about the eating quality or trust the supplier. Quality branding for assured high eating quality is a way to make it easier for the consumer to choose between products when buying. Documentation of the history of the meat back to a farm or even to the specific animal is another way of convincing the consumer that the meat industry has control of the production and thereby the quality of the meat.

To meet the demand for healthy meats, development of low-fat products with optimal fatty acid composition is one of the future challenges for the industry. A further focus on developing meat products with low concentrations of sodium is also one of the major challenges for the industry.

Convenience is another future challenge for the meat industry, not only in producing ready meals or meal components, but also in ensuring a high degree of convenience of the fresh meat. It must be easy for the consumer to go home and prepare the meat.

Retailers demand a long shelf-life for the meat and meat products, even though the consumers perhaps prefer freshness. This is another challenge for the meat industry. How do we produce fresh meat with a superior quality and a long shelf-life? Case ready production can be a way to achieve this.

To meet the challenges of the future, co-operation between the industry and scientists within the fields of meat quality, sensory science and microbiology is therefore of importance to ensure that research and product development gain from each other.

1.6 Acknowledgement

I would like to thank Susanne Støier, Niels T. Madsen, Ina Clausen, Christian Vestergaard from Danish Meat Research Institute and Grethe Andersen from Danish Meat for valuable inspiration, discussions and help during the writing of this chapter. I would also like to thank Sisse Fagt from the Technical University of Denmark, the National Food Institute, for letting me use her results even before they are published.

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Biology and regulation of carcass composition

P.L. Greenwood,     NSW Department of Primary Industries, Australia

F.R. Dunshea,     University of Melbourne, Australia


This chapter provides an overview of the generally accepted understanding of the biology and regulation of carcass composition. More specifically, patterns of growth are briefly described and an overview of the biology of carcass tissue growth and development is provided. Regulation of compositional characteristics by nutrition, genotype and metabolic modifiers is also reviewed. Finally, future trends in research relating to the regulation of carcass composition and meat production are discussed.

Key words

carcass composition



metabolic modifiers

meat production trends

2.1 Introduction

The composition of carcasses and meat are important for consumer acceptance and health, and for meat processors due to the economics associated with meeting specific market requirements, maximising yield of saleable meat, and minimising wastage due to excessive fatness and bone content. Carcass and meat composition are important to producers of meat due to their importance for growth and the efficiency of utilisation of nutrients, and in meeting market specifications to maximise returns.

Defining carcass composition depends on the market segment for which the meat is destined. In a more traditional sense, carcass composition refers to the absolute or relative amounts of muscle, fat and bone, or of protein, lipid, ash and water. From a processor perspective, this definition can be further refined to encompass quantities of retail meat, fat trim and bone and, within the wholesale component of meat, into primal cuts that may be prepared with or without bone. However, the complexity of the modern marketplace has resulted in a redefining of carcass composition to include other factors such as those listed below and detailed elsewhere in this book.

Consumer requirements for meat vary widely depending upon socio-economic factors. In developing countries, there is an increasing demand for protein of which meat is an important and growing contributor. In developed countries, there is an increasingly segmented marketplace that demands meat products based on factors, detailed elsewhere within this book, that include for example: calorific, fat and protein content; specific dietary components such as omega-3 and other fatty acids, iron, zinc, antioxidants and other macro and micronutrients; intramuscular fat content or marbling; tender, juicy and flavourful meat; retail attractiveness including colour; residue free meat; organically produced meat; animal welfare considerations; environmental impact; and low cost meat.

For the meat producer, the primary objective is maximum efficiency of nutrient utilisation to achieve specific market requirements that maximise income. To meet this goal it is necessary to utilise a genotype capable of meeting market specifications, and to provide a suitable environment that includes enough nutrients of a quality appropriate for the various stages of growth and development of that genotype. The profitability of processing animals for meat depends upon a supply of product that is within specification, maximising the yield of saleable meat of an appropriate specification, and minimising the amount of fat that has to be trimmed from a carcass and the amount of bone relative to saleable meat.

In this chapter, we provide an overview of the generally accepted understanding of the biology and regulation of carcass composition. More specifically, we briefly describe patterns of growth, provide an overview of the biology of carcass tissue growth and development, and review regulation of compositional characteristics by nutrition, genetics and metabolic modifiers. Finally, we discuss future trends in research relating to carcass composition and its regulation.

2.2 Patterns of growth of carcass tissues

The carcass tissues follow well-defined patterns of growth, referred to as allometric or relative growth due to their differing rates of accretion during development, growth and maturation (Hammond, 1932; Palsson, 1955). Overall, relative growth of bone precedes that of muscle which precedes that of fat. Hence, at younger ages and lighter body weights bone is present at greater proportions than muscle and fat, respectively, than at older ages and heavier weights. Carcass lipid increases at an increasing rate compared to carcass protein and hence contributes to carcass energy content at an increasing rate during postnatal growth (Fig. 2.1). This relationship is more evident when assessed on a bodyweight- or carcass weight-specific basis (Fig. 2.2). The relative proportions of the carcass tissues will differ at a given age or weight due to genotype or gender, depending upon the maturity pattern and mature size, and on the supply of nutrients and growth rate. For example, later maturing, higher mature weight animals will have a greater proportion of lean tissues (muscle and bone) at the same age or weight than earlier maturing, lower mature weight animals, which will be fatter. Similarly, animals of the same genotype nourished to grow rapidly to a given age or weight will generally have a lower proportion of lean tissues and a higher proportion of fat than those nourished to grow more slowly (Fig. 2.3). This is due to the priority of use of available nutrients favouring bone over muscle over fat accretion when nutrients are limiting (Hammond, 1944; Palsson, 1955). Conversely, as the supply of nutrients increases, adipose tissue accretion increases by more, relative to muscle and bone.

Fig. 2.1 Pattern of growth of carcass components and the contribution of protein and fat to energy stored in the body of the pig. From Etherton and Walton (1986) in Mitchell et al. (2001). © J Anim Sci.

Fig. 2.2 The relationship between weight of protein and fat in lambs and (a) age and (b) fleece-free fasted body weight (FFFBW). Black (1983) in Oddy and Sainz (2002). Reprinted by permission of CSIRO Publishing and CABI, Wallingford, UK.

Fig. 2.3 Relationship between feed intake and growth of pigs from 20 to 45 kg and body composition at 45 kg. From Mitchell et al. (2001) using data from Campbell et al. (1983). Reprinted from Biology of the Domestic Pig, edited by Wilson G. Pond and Harry J. Mersmann. Copyright © 2001 by Cornell University. Used by permission of the publisher, Cornell University Press.

2.3 Biology of carcass tissue development and growth

2.3.1 Embryonic and fetal development


Most fetal growth occurs during the final third or so of pregnancy in cattle (Winters et al., 1942) and sheep (Wallace, 1948a,b; Joubert, 1956; Everitt, 1968), and from about day 70 onwards in pigs (Marrable and Ashdown, 1967), although genesis and growth of vital organs and body tissues commences during early gestation. Development of the vital organs generally precedes that of the carcass tissues (Palsson, 1955; Black, 1983). However, development of muscle is evident by about day 30 of gestation in sheep (Wilson et al., 1992), and of bone by about day 35 of gestation (Altschul, 1987) in sheep.


Growth of the skeleton, particularly of long bones, is the primary determinant of mature size, and is an important regulator of skeletal muscle growth due to stretch induced hypertrophy of muscle.

During early fetal life, bone commences developing on non-ossified, cartilaginous rods, plates and disks that act as templates for subsequent ossification and bone formation. In sheep, ossification commences from about day 35 for the jaw bones (Altschul, 1987), followed by commencement of mineralisation of long bones of the fore- and hind-limbs, in that order, and of other bones of the skull and thorax by day 41 of gestation (Wenham, 1981; Altschul, 1987). Ossification of the long bones commences in the first trimester in pigs (Hodges, 1953; Connolly et al., 2004) and cattle (Gjesdal, 1969; Sterba, 2004).

The three major cell types that contribute to formation of the skeleton are the chondrocytes, which are of mesenchymal origin and form cartilage, osteoblasts, which also originate from the mesenchyme and produce bone matrix, and the osteoclasts, which resorb bone and are of haematopoietic origin (Erlebacher et al., 1995).

Long bone development is characterised by endochondral ossification, with development of a solid, central primary core of ossification, the diaphysis, which develops towards the ends of the still growing cartilaginous model. Ossification also occurs within secondary sites, the epiphyses, which develop into a shell encompassing plates that become the epiphyseal growth plates between the diaphysis and the epiphyses at the ends of the long bone. This process is accompanied by continual remodeling as a result of resorption and apposition (anabolism) of the growing bone and development of a central core. This core contains blood vessels, reticuloendothelial cells, and connective tissue including adipocytes that form bone marrow.

Growth of bones occurs due to continued proliferation of cells known as chondrocytes that arise from progenitor cells within the growth plate, and which secrete the chondroid matrix. Existing cells become more distant from the proliferative zone of the growth plate as proliferation and bone growth continues, and subsequently hypertrophy and extrude calcium which mineralizes the chondroid or cartilage matrix. In turn, the mineralized chondroid matrix is invaded by blood vessels and osteoblasts which secrete osteoid and form bone. Concurrently, osteocytes remove residual chondroid matrix from the newly formed spongy bone, which is replaced by dense bone.

Compared to soft tissues, the skeletal size of the fetus is relatively resistant to the growth-limiting effects of restricted nutrient supply due to maternal nutritional and/or placental limitations. Although some between-fetus variation in bone development and skeletal size can occur within the first third of pregancy (Hafez and Rajakoski, 1964), most of the increase in between-animal variation in size of the fetal skeleton occurs during the final third of gestation (McDonald et al., 1977; Greenwood et al., 2002b).

Bone represents a much higher proportion of body weight in newborn livestock than muscle, adipose, and other soft tissues. Bone growth continues during postnatal life until growth plate closure, which generally occurs shortly after puberty. At this time, progenitor cells cease proliferation and are consumed into mineralised tissue, and the growth plate is replaced by bone. Apart from gross differences in size and structure of bones between the livestock species, finer-scale structural differences also exist at birth and during postnatal growth (Mori et al., 2005).


Muscle fibres form as a result of the alignment and fusion of myoblasts, which arise from mesodermal somites, to form primary myotubes in the first instance (Hauschka, 1994). Subsequently, myoblasts align and fuse on the surface of established myotubes and/or myofibres to form secondary and/or tertiary myofibres. These latter phases account for most of the increase in the number of fibres within a muscle.

The timing of stages of myogenesis in livestock (Table 2.1) was previously reviewed by Picard et al. (2002) and Brameld et al. (2003). In sheep, primary myogenesis commences at about day 32 of gestation, secondary myogenesis by about day 38, and myogenesis is concluded by about day 110 (Wilson et al., 1992; Maeir et al., 1992; Greenwood et al., 1999), at which time the number of muscle fibres is established and hypertrophy of myofibres commences (Greenwood et al., 1999). In cattle, primary myotubes appear prior to day 47 of gestation (Russell and Oteruelo, 1981) and secondary myotubes are present from around day 90 (Russell and Oteruelo, 1981; Robelin et al., 1991), with myogenesis complete by about day 200 (Picard et al., 2002). In the pig, primary myogenesis is evident by day 35 of gestation, secondary myogenesis by day 55, and myogenesis is believed to be completed by day 90 (Wigmore and Stickland, 1983; reviewed by Novakofski and McCusker, 2001), although there is some evidence of a wave of tertiary myogenesis during early postnatal life in pigs (Lefaucheur et al., 1995).

Table 2.1

Timing (days of gestation) of muscle development in livestock (after Brameld et al., 2003)

*days postnatal

While it is believed that the number of primary myotubes influences the total number of myofibres formed within a muscle, the number of secondary and/or tertiary myofibres formed appears more susceptible to environmental influences. Evidence exists that the number of myofibres can be altered at least to some extent by maternal nutritional deprivation during early to mid pregnancy in sheep (Everitt, 1968; Zhu et al., 2004) and pigs (Wigmore and Stickland, 1983), but less so in cattle (Greenwood and Cafe, 2007). In pigs, improved nutrition (Dwyer et al., 1993; Gatford et al., 2003) during early to mid pregnancy may increase myofibre number, although this is not always the case (Nissen et al., 2003). Likewise, somatotropin treatment of sows during early to mid gestation increased fetal muscle fibre numbers in some studies (Rehfeldt et al., 1993, 2001) but not others (Gatford et al., 2003) (see reviews by Rehfeldt et al., 2004 and Rehfeldt and Kuhn, 2006). However, the extent to which these findings may also reflect differences in the growth of intrafascicularly terminating myofibres during early life is unclear (see Swatland and Cassens, 1973), and may account for differences in findings between apparently similar studies. Hence, an assessment of prenatal effects on myofibre number may be more reliable if undertaken later in life (for example, Greenwood and Cafe, 2007), when radial growth rather than elongation of myofibres is most responsible for myofibre hypertrophy (Swatland, 1994), provided that accurate measurement of muscle cross-sectional area can be made. By comparison, with studies on environmental or exogenous effects during early to mid pregnancy, in sheep severely growth-retarded during later fetal life due to a placental restriction on growth (Greenwood et al., 2000b), the number of myofibres formed was unaffected in various anatomical muscles relative to normally grown fetuses (Greenwood et al., 1999, 2000a; McCoard et al., 2000). However, runt pigs have fewer myofibres than their larger counterparts, demonstrating the severe influences on development that can occur during myogenesis (Hegarty and Allen, 1978; Rehfeldt and Kuhn, 2006)

Following completion of myogenesis, muscle satellite cell cycle activity is believed to be the sole source of new myonuclei in support of myofibre hypertrophy (MacConnachie et al., 1964; Shafiq et al., 1968) which continues, albeit at declining rates (Allbrook et al., 1971; Greenwood et al., 1999), until some time approaching mature muscle mass. By this time, the number of nuclei in muscle fibres, as indicated by the amount of DNA present, is maximal (Trenkel et al., 1978; Solomon et al., 1986; Di Marco et al., 1987; Novakofski and McCusker, 2001), while the ratio of protein to DNA which

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