Low-Fat Meats: Design Strategies and Human Implications

Kingdom

Preface

H.D. Hafs; R.G. Zimbelman

This book deals broadly with how to reduce fat in animals and in meat from animals by experimental technologies as well as by methods that have been used for many years. It also focuses on the potential human health, economic, ethical, and social issues associated with implementing such changes, as well as on the concerns of animal well-being. This book will be useful for faculty and students in animal science, food/meat science, economics, marketing, and sociology; for organizations of meat producers, packers, purveyors, and retailers; for animal science and meat/food science researchers in university, government, and industry laboratories; and for people charged with making, changing, and implementing public policy regarding our meat supply.

People generally understand that red meat is a readily available source of some critical nutrients. In the United States, beef, pork, veal, and lamb together supply about 27.6% of dietary protein, but only 15.5% of the calories, and they offer concentrated sources of B vitamins, iron, and zinc. In the past, most consumer concerns have been related to the cost, wholesomeness, and sensory qualities of meat. Increasingly, however, consumers worry that excessive dietary fat can lead to cardiovascular disease and possibly to colon cancer. Nevertheless, the meat marketing structure has rewarded producers for producing fat animals.

Thus far, producers have been able to alter the carcass composition of meat animals only through nutrition, breeding, and adjusting the age at slaughter. Unfortunately, until recently, producers have not been given incentives to use these tools to adjust for changing consumer demands. On the other hand, meat processors have been increasingly sensitive to consumer demands, creating an impressive array of low-fat meats now routinely available from most retailers, especially for pork. Also highly sensitive to consumer demands, meat purveyors and retailers have trimmed excessive fat from meat before cuts are presented in retail stores or restaurants. Unfortunately, this wastes resources because fat is energetically much more costly than muscle to deposit in an animal. Ironically, at this time, some of the trimmed fat from beef is sold at up to $0.50/lb to be mixed with very lean beef for fast food hamburgers, thereby increasing the price paid for fat cattle. However, a significant portion of the trimmed fat is disposed of, used to make lard, or mixed with grains to be fed to animals.

Mounting consumer concerns about dietary fat and the inefficiencies of the present system have led to new ways of measuring fat in animals and meat, to serious efforts to change how farmers are paid for meat animals, and to imaginative new meat products processed to modify their composition. Some of these new technologies will require significant investments, and not everyone agrees with the proposed changes because they may lead to major changes in the entire meat marketing system.

While these meat marketing changes have been developing, two metabolic control systems by which animals normally partition nutrients between fat and muscle have been discovered. Somatotropin and β-adrenergic agonists are the key compounds in these systems. When given to animals, these partitioning agents dramatically reduce carcass fat, increase muscle mass, and reduce the amount of feed required to produce the carcass. Commercial versions of these partitioning agents could be available in the United States as early as 1995, allowing producers to respond quickly to value-based procurement of animals by packers.

Since partitioning agents reduce fat, increase lean meat, reduce the cost of producing meat, and produce meat that is more healthful, one might predict their rapid commercial adoption. However, new worries have arisen. The treated animals likely will require modified management for optimal well-being. For example, the dietary requirements of animals given partitioning agents are altered. And there are other critical issues. Have the sensory characteristics of the meat changed, and what is its nutrient value? How will the carcass composition changes affect human health, and what will be the socio-economic impact on the rural communities where meat animals are produced? Will meat packers reward producers for animals with less fat and more muscle, and are processors prepared for the dramatic changes in carcass composition? These issues and the public policy implications they raise are a major focus of this book.

Notwithstanding their promise to improve meat quality, partitioning agents should be viewed only as an interim solution. Partitioning agents have not yet been widely adopted, and alternatives are already on the horizon. For example, animals may be immunized to block some metabolic steps that normally lead to excessive fatness, and genetic engineering may speed permanent genetic changes to provide low-fat meat products. Although commercial applications of these two biotechnologies are more distant than the use of partitioning agents, they likely will lead to some of the same questions and public policy issues and are therefore discussed herein.

In recent years, consumer demand for low-fat meat has solidified. The question now is not whether the industry should provide low-fat meat but rather how to accomplish that goal and how quickly the industry will respond. The hope is that by facilitating public discussions, this book will lead to early identification of all of the issues and will speed consensus.

Chapter 1

Improving Carcass Composition through Selective Breeding

William R. Lamberson

I Principles of Genetic Improvement

A Genetic Structure of the Livestock Industry

The livestock and poultry industry is organized as a multitiered genetic pyramid. The narrow peak of the pyramid represents independent seedstock producers and livestock and poultry breeding stock companies. It is at the peak of the pyramid that genetic improvement practices of performance testing and selective mating are applied, potentially resulting in continually improving breeding stock. The seedstock level of the pyramid contains only a small fraction of the animals and birds in the livestock and poultry industry, but it is in ultimate control of any genetic change that occurs. The broad base of the pyramid is composed of commercial livestock and poultry production units. Commercial producers purchase breeding stock, and with it changes in genetic merit, from the seedstock level. Breeding stock may move directly from seedstock to commercial enterprise or they may be separated by one or more intermediate multiplication layers. Time required for dissemination of genetic improvement from the seedstock to commercial levels may be as little as 1 to 2 years in poultry and swine, but as many as 5 to 10 (or more) years in cattle. The commercial units ultimately produce millions of market animals and birds to be processed and sold as meat and meat products to consumers. Their survival is dependent upon production of a desirable product, one that is in demand by consumers, at an economical price.

Return to the producer is determined by unit cost of production and by price received for the product. Genetic merit of the stock and skill of the livestock producer in managing the stock to express their genetic potential are major factors affecting return to the producer. Genetic merit of the animal affects return to the producer in three ways. First, cost of production is affected directly by cost of the breeding stock. In general, price increases as genetic merit improves. Second, cost of production is affected by productivity of the animal. Reduced cost of production is associated with superior genetic merit for high reproductive rate, rapid growth rate, and feed efficiency. Third, value of product is affected by genetic merit of the animal for carcass composition. Lean, heavily muscled animals currently command premium prices in the marketplace.

Producers must weigh the cost of obtaining improved genetic merit against the expected return. Genetic pressure will be applied to change traits where return is greatest. Often, it is more cost effective to change traits by management methods (i.e., changes in age at slaughter, restriction of feed intake, vaccination, or alteration of diet) than it is to change traits genetically. Relative importance of traits shifts depending upon management and marketing situations. In swine, for example, among 11 traits studied, decreasing percentage fat at slaughter was predicted to have the greatest impact on improving return to swine producers when pigs were marketed as weight of carcass lean (Tess et al., 1983). However, when marketed on a live-weight basis, without regard to composition of the carcass, reducing percentage fat decreased return to the producer. Improvements in reproductive rate and growth rate under that marketing system became the most important traits in improving return to the producer.

Many tools are available to seedstock producers to make genetic change in their populations. Since they market genetic merit to their commercial customers, they must be concerned with improving the traits which will yield greatest return to commercial livestock and poultry producers. Given that economics drive the commercial producer to seek seedstock giving greatest return for dollar invested, how do seedstock producers develop animals with the optimum genetic merit?

B Selection among Populations

The first tool utilized to provide excellence in genetic merit is selection among parentstock populations. Choice of animals from the best-suited breeds and strains has long been the most obvious and commonly utilized genetic improvement tool. Original cattle populations in the Southwest were Longhorns introduced by the Spanish. These original cattle were improved and replaced by introduction of the British breeds. Later, Brahman and Brahman crosses were introduced to provide heat tolerance. More recently, continental European breeds have been introduced into breeding programs to provide genetic merit for growth and size previously unavailable.

Large numbers of breeds and lines exist in cattle, swine, and poultry. New breeds and strains are continually being developed or imported and evaluated for introduction into our production systems. McLaren (1990) includes information on 201 breeds of swine as a partial listing of the total available worldwide. Of that total, 8 are currently important in the U.S. Samples of highly prolific breeds from China and very lean breeds from Europe have recently been introduced into the U.S. for evaluation. Although unlikely to be important as pure strains, these breeds may contribute in crossbreeding systems or in the formation of new breeds or strains. Enormous variation exists in productivity of different breeds. Ongoing attempts to identify breeds with potential to positively impact the U.S. livestock industry are a needed research effort.

C Selection within Populations

Selection within populations is the primary method through which permanent genetic improvement is achieved. Selection is practiced within seedstock populations, usually composed of some pure genetic group. These seedstock populations make up a small fraction of the total number of animals in a livestock species. They are in the hands of individual producers or commercial breeding companies. Genetic improvement made in these seedstock populations passes through one or more levels of dissemination to commercial herds and flocks by sale of semen, males, and sometimes females. Seedstocks have traditionally been in the hands of many individual livestock producers, each of whom may have their own ideas regarding appropriate objectives for their herd. Variation in objectives reduces progress toward any one goal; however, it has the advantage of maintaining genetic diversity and allowing commercial producers to choose among populations and obtain seedstock particularly suited for their specific needs.

Progress from selection for any one trait decreases as the number of traits considered in the selection program increases. Most seedstock producers or breeding companies thus emphasize a small number of traits to enhance strengths or correct weaknesses of their particular breeding stock. Traits chosen for emphasis in genetic selection programs are a tradeoff between the value of expected change and the cost of selection. The potential to change a particular trait by selection is determined by the degree of genetic control of the trait or its heritability, the variability of the trait, usually measured by the standard deviation, and our ability to measure the trait or its accuracy. Another factor affecting response to selection is the proportion of the population that is retained (selected) to produce the next generation. The smaller the proportion selected, the greater their potential superiority. This factor is termed the intensity of selection.

The heritability is measured on a scale of zero to one with zero being no genetic influence and one representing complete genetic control. Alternatively, it can be defined as the proportion of the superiority of the selected parents, relative to that of the population as a whole, which is passed on to the next generation. Most traits associated with carcass composition have a relatively high degree of genetic control. The heritability of cutability traits usually falls in a range of 0.35 or higher. This compares to reproductive traits which characteristically have heritabilities in the range of 0.05 to 0.25 and growth traits which have heritabilities which range from 0.20 to 0.50 (Lasley, 1978).

When individuals are ranked on a specific criterion and the highest ranking individuals are selected, superiority of the selected individuals is proportional to the variability of criteria. Traits with high variability, even if the heritability is low, may still respond well to selection. Conversely, as variability declines, response to selection approaches zero. An example of a decline in response to selection associated with loss of variability would be selection for decreased fat thickness in swine. As the thickness of backfat declines, the variability also declines, the magnitude of superiority of the selected group is reduced, and the selection response approaches zero. A limit is reached which may be physical, e.g., it may be impossible to measure backfat in very lean animals, or the limit may be biological; i.e., the animal with less fat cannot survive or reproduce.

Selection within a population has been shown experimentally to be an effective means of improving carcass composition. Several studies in laboratory animals and swine have shown the effectiveness of selection in improving carcass composition. Backfat in pigs was reduced by 2.6% and 4.0% per generation in two studies in which backfat was the single selection criterion (Gray et al., 1968; Berruecos et al., 1970). Total fat in the carcass decreased with the reduction in backfat.

D Mating Systems

A mating system is the method of choosing which selected males are mated to each selected female. Two general variations in mating systems exist that can be used to advantage for genetic improvement. Inbreeding is the mating of individuals which are related. Inbreeding frequently reveals detrimental genes carried by seedstock lines which can then be eliminated from the population by selection. Mild inbreeding combined with selection has been heavily utilized in the development of breeds of livestock. More extreme inbreeding, combined with selection and crossing, has been used successfully by corn breeders to produce improved varieties of hybrid seed corn. Poultry breeders have also attempted to use this method of genetic improvement. Unfortunately, success similar to that of corn breeders has not occurred in livestock breeding. Selection is less intense in livestock species relative to plants because of their lower reproductive rate. Selection when combined with inbreeding has resulted in less genetic improvement than selection without inbreeding, thus inbreeding has lost favor with most livestock breeders.

Crossbreeding includes a collection of mating systems which involves mating males and females less related than the average relationship across all animals in a population. Crossing of existing lines or breeds has attained great favor with livestock producers. Virtually all commercial poultry result from line crosses. Approximately 95% of market hogs are crossbred. Smaller proportions, but still a majority, of market cattle are crossbred. Two advantages are associated with crossbreds. First is heterosis or hybrid vigor, a general improvement in fitness and performance associated with crossbreeding. This improvement in performance is primarily associated with reduced mortality, increased reproduction, and faster and more efficient growth. Unfortunately, heterosis is also associated with increased fatness. The second advantage of crossbreeding is utilization of breed complementation. That is the mating of sire and dam lines in such a way to take advantage of specific maternal and paternal strengths of the lines. It is in this way that crossbreeding can be an advantage in improving carcass composition. Males from lines with excellent carcass merit can be mated to females from lines with superior genetic merit for reproduction and maternal performance. This type of system can lead to performance for both characters better than that which can be obtained in any single pure line.

II Improvement of Carcass Composition: Progress to Date

A Poultry

Recent surveys of commercial livestock suggest that improvements in carcass composition have been limited. The force that drives livestock producers in the commercial industry to seek genetic change is economic return. Until recently, insufficient economic incentive has been present to turn producers’ attention away from traits more closely associated with performance, such as growth, efficiency, and reproduction. Cycles of genetic change in carcass value have been in response to favorable price differentials for composition and quality.

The poultry industry has been very effective in utilizing within line selection to make improvement in traits of economic importance. The high reproductive rate of chickens allows high selection intensity yielding rapid selection response. The broiler industry has very successfully concentrated on improvement of growth rate and feed efficiency. The industry’s sharp focus on a small number of traits has also contributed to the development of improved lines. These selected lines are crossed to take advantage of heterosis in the production of commercial broilers.

Heavy selection emphasis was placed on a single trait: first, growth rate, and later, feed efficiency. Genetic response has been dramatic. Body weight at 8 weeks has nearly doubled and feed consumed to produce a pound of gain has been cut nearly in half since 1940 (Shoffner, 1989). Recently, concern regarding fat in broilers has gained attention in the poultry industry. It is anticipated that procedures for measurement of carcass value of the live bird will be implemented in selection programs. Selection should quickly correct problems with carcass quality or composition. In addition, vertical integration of the poultry industry combined with rapid generation turnover of poultry results in a rapid response to needs of the consumer.

B Swine

Only since the late 1980s have sufficient incentives been paid to demand that swine producers reduce fat on market hogs. Prior to that time, the trend in fatness of hogs was not consistently in a favorable direction. McLaren et al. (1990) examined carcasses from performance-tested pigs in Illinois from 1968 to 1989. Nearly 6,000 carcasses were examined. Backfat did not change significantly when measured on weight-constant carcasses between 1968 and 1983. From 1984 through 1987, a substantial unfavorable trend in fatness was observed. This unfavorable trend has since been reversed and fatness declined from 1987 through 1989.

These early trends were confirmed by David et al. (1985) in a study of selection procedures and results in seedstock swine herds. Data from over 100,000 swine in 18 herds from 1971 to 1979 were examined. A slightly favorable phenotypic trend in backfat was observed. Examination of selection procedures indicated that backfat received 19% of the total selection emphasis in males and 9% in females. Selection was practiced for reduction of backfat in nearly all herds but it received only about 16% of the attention which it potentially could have, had backfat been the only trait considered in making selection decisions.

Change in carcass composition of swine in Europe has been the subject of several recent reports. In general, changes in carcass traits have been more favorable in Europe than in the United States. Ratio of fat to eye muscle has improved by 5 to 10% of a phenotypic standard deviation per year from 1979 to 1987 in herdbook populations of four breeds of swine in Germany (Kovac and Groeneveld, 1990). Genetic and environmental trends both contributed to the improvement. Genetic trend in backfat thickness was significantly negative in French Large White but was negligible in French Landrace (Ollivier et al., 1991). Trends in Polish Large White nucleus herds were surveyed from 1979 to 1987 (Kaplon et al., 1991). Phenotypic trends were favorable with backfat decreasing at a rate of 0.065 mm per year. Genetic trend accounted for approximately one-seventh of the change. Genetic evaluation of on-farm tested Swedish Yorkshires and Landrace revealed improvements in the percentage of carcass in premium cuts, a trait which has the combined influence of muscling and fatness (Hofer et al., 1992). The genetic trend over 20 years was + 1.9% in Yorkshires and + 1.2% in Landrace.

Genetic trends in carcass composition in Europe were favorable, but only a part of favorable phenotypic trends. This suggests that Europeans have been successful in improving the genetic merit of their swine herds as well as adapting management strategies to increase leanness of pigs. Concentration of seedstock in fewer hands in Europe than in the United States may lead to more rapid implementation of new technology to evaluate genetic merit of pigs. Government subsidies may assist seedstock producers in achieving more rapid rates of genetic improvement.

C Beef Cattle

Randomly sampled carcasses from 28 federally inspected beef packing plants were evaluated for USDA yield and quality grade factors from October to December 1991 (Lorenzen et al., 1993). A total of more than 7,000 carcasses were examined by experienced evaluators. Ribeye muscle area and fat thickness were measured and a marbling score was assigned. Data were compared to results of a similar study conducted in 1974. Carcasses were heavier (36.8 kg) and had larger ribeye muscle area (7.1 cm²) in 1991 than in 1974. Adjusted fat thickness on the carcasses had changed by less than 1 mm. Contemporaneous progeny resulting from semen from bulls from 1971 versus 1984 showed a slight improvement in quality grade but poorer yield grade indicating that fatness within breed had increased over that time period (Sanders et al.,1988). Thus, the limited changes in adjusted fat thickness observed by Lorenzen et al. (1993) while the carcass weight increased were likely due to an increased proportion of continental European breeds in the the cattle’s genetic makeup.

Further results presented by Lorenzen et al. (1993) point to the dilemma faced by beef producers. Carcass value is dependent on quality grade and yield grade as well as weight. One factor influencing quality grade is intramuscular fat (marbling). Increased marbling results in higher quality grade and higher price but is also associated with increased subcutaneous fat (Table I). Conversely, superiority in yield grade is commanded by leaner animals. Increased weight is also associated with increased fatness, thus feeding animals to heavier weights will generally result in improved quality grade but inferior yield grade. Strategies to optimize return from these complex relationships usually involve variation in length of time animals are fed, rather than changing the genetic makeup. Cost of feed also influences the relationship. Animals tend to have reduced efficiency of feed utilization for lean deposition as they reach heavier weights. When feed is inexpensive, it is more economical to feed animals to heavy weights than when feed is expensive.

Table I

Mean Fat Thickness by Quality Grade, Yield Grade, and Carcass Weight

From Lorenzen et al. (1993).

III Selection on Carcass Composition: Correlated Responses

One impediment to selection to improve carcass composition has been concerns regarding adverse responses in correlated traits. These concerns have most often been with regard to reproduction. Increased leanness in maternal livestock herds is thought by some stockmen to be associated with reduced fertility of the female. The reduction in fertility is primarily associated with delayed puberty and increased incidence of failure to rebreed. Reduced litter size may also be a concern. Reviews of the subject have not indicated a strong relationship between leanness and reproduction in swine in the absence of the porcine stress gene (Lamberson, 1990a). The stress gene is known to result in increased leanness and muscling and has also been associated with decreased litter size and poor maternal environment (Christian and Mabry, 1990). Selection for increased lean and decreased fat during the 1960s in the U.S. resulted in an increased frequency of the gene which caused declines in reproductive rate.

Since most commercial livestock are crossbred, specialized sire and dam lines can be utilized to reduce possible problems with reproduction in extremely lean genotypes. A system of using lean, muscular terminal sire lines mated to more moderate dam lines may prevent difficulties. In all livestock species, adequate nutritional management can be used to mitigate problems, as well.

IV Potential for Future Improvements in Carcass Composition

A Classical Breeding Methods

Long-term experiments designed to find limits to selection response have shown that genetic variation is virtually inexhaustible. There is genetic potential to achieve change well beyond that practical for conventional livestock production. Application of intense selection to reduce fat in livestock and poultry has potential to reach a biological limit at which the animal will no longer be able to reproduce or possibly survive under current production conditions. An example of a similar situation is selection for muscling in turkeys. Muscling has become sufficiently developed that reproduction by natural mating is nearly impossible. The industry has adopted artificial insemination as a normal production practice to accommodate the biological change in the bird. Likewise, new production practices might need to be adopted to accommodate biological needs of extremely lean genotypes of livestock.

Given that little reduction in fatness occurred over the past two decades, is there reason for optimism that classical breeding methods can be used to reduce fat in the future? Yes, because of two factors: incentive and means. Recent price incentives have increased demand for lean breeding stock, particularly in swine. Automated means of measuring fat content of carcasses are becoming incorporated in packing plants which allow packers to provide feedback to producers regarding quality of carcasses. The means to measure leanness in the live animal has been greatly improved by the introduction of real-time ultrasound. Accurate, direct measures of leanness and muscling of the live animal are now available. Purdue University and the University of Missouri have been leaders in using this technology to assist breeders in determining genetic merit of swine. Increasing vertical integration of the swine industry will also speed movement to leaner swine. Vertical integration improves the chain of communication from meat processors to commercial producer to seedstock breeder and thus coordination of genetic improvement programs.

The poultry industry is already vertically integrated. This industry will respond quickly to consumer or product fabricator demands for leaner birds. Development of technology to accurately measure fatness of the live bird would improve response to consumer demands for leaner birds.

Carcass traits have also drawn recent attention in the beef cattle industry. The American Angus Association and the American-International Charolais Association have recently introduced carcass information in their sire summaries. These estimates of genetic merit allow simultaneous, independent evaluation of factors affecting quality grade and yield grade. This information allows producers to select lean bulls which are superior for marbling or quality grade. Success in these two breeds will likely result in other breeds adopting similar evaluation procedures.

B Application of Molecular Genetics

Techniques in molecular genetics provide additional tools to enhance rates of genetic improvement. Mapping of the genomes of swine, cattle, and poultry is expected to isolate sites upon which to focus efforts. Preliminary efforts in marker-assisted selection have been practiced through use of markers associated with the porcine stress gene. Selection on markers associated with loci with measurable effects on traits of economic importance is currently utilized by some swine breeding companies. Commercial companies also offer services to screen for markers in cattle to aid in making selection decisions. Commercial applications in this area will, without doubt, increase in the future. These techniques are likely to be a supplement to, rather than a replacement for, classical genetic methods in livestock improvement programs.

Direct modification of the livestock genome through development of transgenic animals is a technique likely to be applied in animal improvement in the future. This technique shows potential for dramatically changing animal performance. Procedures for direct modification are reviewed by Rexroad and Pursel in Chapter 16. The interaction of direct modification of the livestock genome with classical breeding methods needs further investigation. It is likely that transgenic animals will be the basis for genetic lines that will be further improved by classical breeding methods and then used in cross-breeding.

C Potential for Reducing Variability

A ubiquitous concern of meat packers is variability of carcasses. Greater uniformity of carcasses would increase efficiency of procedures in packing plants. Hohenboken (1985) reviewed possible strategies for reducing genetic variability of metric traits. The most favorable procedure was to utilize first crosses of inbred or highly selected lines. Cloning individual genotypes was also suggested as a means of reducing phenotypic variation. Lamberson (1990b) reported that variation in backfat of littermate cloned swine would be reduced as much as 45% relative to full-sibs.

An effective and economical procedure for utilizing cloning technology would be crosses of male clones with genetic superiority as terminal sires with female clones with genetic superiority as maternal stock. This procedure would spread the cost involved in producing individual clones among their progeny. Mating of clones from terminal and maternal lines would be similar to repeatedly mating the same pair of animals. The relationship among offspring would be equivalent to that of full-sibs, but not all would be produced as littermates. This procedure would be expected to decrease the range backfat of pigs within herd by approximately 14%.

V Summary

Three primary methods are traditionally used to genetically improve livestock. (1) Choice of the best populations can involve fundamental decisions as simple as deciding among which lines to purchase from which breeding companies or as complex as deciding among breeds for import and testing to potentially be incorporated into stocks. (2) Selection within herds has been practiced for centuries to refine and improve breed characteristics. Experiments in livestock, poultry, and laboratory species have shown that selection is a powerful tool to cause genetic change. Selection is most powerful when applied to a limited number of traits which can be accurately measured. However, selection will not be applied to change a particular trait unless an economic incentive exists. Dramatic improvements in growth rate of broilers and milk production of dairy cattle are examples of effective use of selection. (3) Systems of mating such as crossbreeding can be used to improve performance of livestock and poultry. Nearly all of the poultry and more than 95% of swine marketed in the United States are crossbred. Crossbreeding results in substantial gains in performance for many traits, but unfortunately the improvement in performance due to heterosis is not accumulative; that is, it is not passed on to offspring. Thus selection within the best available stocks used to make up the crossbreds is necessary for continued genetic