Genetics of Pork Quality Allan P. Schinckel Department of Animal Sciences, Purdue University There has been a shift in consumer attitudes towards their purchases of meat products. Greater emphasis is being placed on palatability characteristics such as tenderness, juiciness, and flavor versus issues such as cost, food safety or leanness. The food industry is becoming more consumer driven which is driving segmentation of the market to specific products. For example, upper scale restaurants and hotels demand consistent pork products with above average quality in terms of tenderness and flavor. The export market has above average requirements in terms of color and marbling. To be able to produce pork products to meet specific market demands, we must identify and select specific genetic populations. It is also important that the quality of our U.S. fresh pork be competitive with alternative meat products. Otherwise, the pork consumption share of the domestic market will decrease. Pork Quality Pork quality is a general term including a combinations of traits that provide for a wholesome product, attractive in appearance and is appetizing, nutritious, and palatable to eat (Kauffman et al., 1990). To be consistently evaluated and interpreted, the best measures of pork quality are quantitative and repeatable (day to day, lab to lab, etc.). The seven sets of pork quality measurements are: (1) color; (2) firmness; (3) water holding capacity (drip loss); (4) ultimate pH; (5) marbling - intramuscular fat; (6) tenderness - shear force; and (7) palatability traits such as taste panel flavor or juiciness. 1 Genetic Variation and Relationships The heritability of many of these pork quality measurements have been estimated. Intramuscular fat, like measures of carcass composition are highly heritable (.40 to .50, Table 1). Ultimate pH, Hunter L scores and tenderness are moderately heritable. In each case, subjective visual scores (color-marbling) have a slightly lower heritability than the objective quantitative measurement (Hunter L, Japanese color score). The measures of pork quality are genetically correlated (Table 2). Ultimate (45h) pH is highly correlated with drip loss, color score and L*. Thus, lower pHu is associated with light colored loin muscle. Also, pHu is negatively correlated with percent lean (-.37) and positively correlated with backfat depth (.21). It is important to realize that the proportion of the total genetic variation in one trait can be explained by the variation in another as the R2 coefficient of variation. Based on this relationship, 40-60% of the genetic variation in drip loss and color can be accounted for by 24h pH. The relationships between percent lean and pork quality are only weak to moderate in magnitude. Thus, selection for improved pork quality and percent lean is possible. Impact of Lean European Lines In the spring and fall of 1995, two large swine lean growth trials were conducted at Purdue University. Participants had the choice of providing pigs in sets of either 24 or 32 per genotype and sex. In the spring of 1995, the terminal cross pigs of four new genetic sources were evaluated with four U.S. terminal crosses (2-Hampshire-Duroc by Yorkshire-Landrace, 1-Duroc by Yorkshire-Landrace and 1-Hampshire by Yorkshire-Landrace). The European pigs had been selected for lean efficiency and contained a higher percent Large-White. The European terminal cross pigs had lower feed intake (13.7%) and had lower carcass fat gain (28.7%; Table 3). The 2 lean European lines required 21.2% less feed per unit fat-free lean gain from 60-250 lbs live weight. Assuming grow-finish feed costs represent 48% of total costs and other costs were the same, the lean European lines had about 10% lower cost per lb than that of the 1995 U.S. terminal cross. However, the lean European lines had lower color scores, firmness scores and marbling scores (Table 4). During the mid 1990's U.S. purebred producers did three things to genetically reduce backfat thickness and increase carcass lean percentage. First, real-time ultrasound use increased which was more accurate than previous A-mode ultrasound. Second, selection pressure was placed on pigs that was reflected by substantial STAGESī genetic trends for reduced backfat thickness and increased pounds of lean. Third, lean sires from Europe, Sweden, Britain, and Denmark were incorporated in the U.S. purebred populations. In 1998, a genotype by environment interaction trial was completed. The European terminal cross was the top European Hampshire Duroc line mated to one of the leanest European maternal lines based on previous Purdue and NPPC trials. The U. S. Duroc × YorkshireLandrace pigs were produced by sampling numerous Duroc sires from four sources mated to average U.S. purebred derived Yorkshire-Landrace females. The trial included 288 pigs reared in two health environments. The differences between the European and U.S. pigs have been substantially reduced (Tables 5, 6, and 7) for all performance, carcass measurements, and measures of pork quality. Selection for leanness and incorporation of lean European lines into the U.S. purebred populations has produced pigs more similar to the European terminal crosses. Genetic by environmental interactions were significant for measures of growth rate, feed intake tenderness and juiciness. 3 Breed Differences Research has found substantial genetic variation amongst different genetic populations for pork quality. Berkshires, Chester Whites and Durocs have better pork quality than Hampshires, Landrace and Yorkshires (Table 8). Increasing the percentage of the three higher pork quality breeds into terminal crosses will improve pork quality. Major Genes Two major genes have been identified and characterized that impact pork quality. The effects of the porcine stress gene (PSS) on carcass composition and quality are well documented (Tables 9 and 10). More than a decade ago, PSS was described as acting as a complete recessive possibly because the results of halophane tests (positive stiff as a board or no response) and measures of muscle damage (CPK levels). However, data in the 1990's demonstrated that carrier pigs had lower pork quality and a substantially higher probability of PSE pork. Based on these results the vast majority of seedstock producers have estimated the stress gene through DNA testing. The other major gene is called the Napole gene. The Napole gene is currently considered to act as a completely dominate gene. In the past, pigs carrying one or two copies of the Napole gene were identified by having substantially highly glycolytic potential (GP) where (GP) = 2 (glycogen + glucose + glucose-6-phosphate + lactate). In the fall of 2000, the DNA test for the Napole gene was made commercially available. The DNA test will allow better research into the effects of the Napole gene. The Napole gene in Hampshires results in lower ultimate pH, increased drip loss and cooking loss (Tables 11 and 12). However, the shear force was lower for the Napole pigs. It is interesting to note that the Napole gene changes in 24h Ph vary from trial to trial. 4 Issues to be Dealt With The field of genetics makes a number of underlying assumptions. Sometimes these assumptions are not totally correct. Within line selection for pork quality assumes that the improvement realized in the purebreds will be realized in the crossbreds. Also, it assumes that the genetic improvement realized under one set of conditions (processing methods, nutrition, post-slaughter handling, rate of chill) will be realized under different conditions. One trial examining these concepts was recently completed at the University of Tennesses (Van Laack et al., 2001). In their study, Berkshire, Duroc, and Hampshire sires were mated to Yorkshire-Landrace F1 females. Half of the pigs were within 2h of transport and the other half withheld from feed 14h prior to transport. The means for the sire line and feed treatments are shown in Table 13. Within the Duroc breed, a significant negative correlation (.42) existed between intramuscular fat percentage and shear force. No significant relationship existed between shear force and intramuscular fat in the Berkshire or Hampshire sired pigs. In Hampshires there was a positive correlation between ultimate pH and shear force and negative relationships in the Duroc and Berkshire sired pigs. The 14h fast increased the ultimate pH of the Duroc and Berkshire sired pigs; but, had no impact on the Hampshire sired Napole carrier pigs. A significant sire line by storage time (2, 7 or 14d) interaction was found for shear force (Table 14). This data shows that "general" relationships amongst the pork quality measurement may vary amongst different genetic populations (pure lines or crosses). The differences in quality between genetic populations may vary somewhat depending on the handling, storage and processing of the pork products. Further research needs to be conducted to evaluate the consistency of the genetic and phenotypic correlations when estimated within line and in crosses. For example, if purebred 5 Durocs are a breed likely to be selected as a high quality terminal sire, then a trial evaluating the genetic and phenotypic parameters of the pork quality and carcass composition variables should be completed within the purebred Durocs and when the sires are used to produce crossbreds. Then, the Duroc breeder must ask how do the Duroc sired pigs need to be improved to better meet the specifications of the identified value added market. The traits measured and selected for must move the genetics of the Durocs such that the crossbreds achieve the specifications set by the purchaser of their commercial customers' pigs. Summary As pork quality specifications are established, commercial producers will have to decide which market will they are producing pigs for. If it is for a high quality value, added market, the pigs will have to contain a higher percentage of lines superior in pork quality. Seedstock producers will have to determine which market or markets their breeding stock is going to fit. The selection must be geared towards their purelines producing terminal cross pigs that meet established pork quality specifications. Literature Cited Kauffman, R. G., W. Sybesma and G. Eikelenboom. 1990. In Search of Quality. J. Can. Inst. Food Sci. Technol. Vol. 23, No. 4/5. van Laack, R. L. J. M., S. G. Stevens, and K. J. Stalder. 2001. The influence of ultimate pH and intramuscular fat content on pork tenderness and tenderization. J. Anim. Sci. 79:392-397. 6 Table 1. Heritability of Pork Quality Traits and Performance. Pork Quality h2 pH - 1 hr .19 pH - 24 hr .19 Hunter L* .31 a* .47 b* .42 Color score-Japan .22 Drip loss .10 Intramuscular fat .44 Marbling score .24 Eating Quality Tenderness, kg .26 Tenderness/sensory .29 Flavor .09 Juiciness .08 Overall acceptability .25 Eating Quality Tenderness, kg .26 Tenderness/sensory .29 Flavor .09 Juiciness .08 Overall acceptability .25 Copenhafer Literature Review, 2001. Table 2. Genetic Correlations pH 24 Drip Color score-Japan IMF Marb Score % Lean Drip -.71 Color score Japan -.62 -.41 IMF -.20 -.21 n/a Marb Score .26 n/a .25 n/a % Lean -.37 n/a -.15 -.12 .02 BF .21 n/a -.02 n/a .31 -.77 L* -.80 n/a -.99 .02 -.23 -.18 Copenhafer Literature Review, 2001. 7 Table 3. 1995 Spring Lean Growth Trail Performance Trait European US % Difference ADG, lbs/d 2.02 2.11 -4.3 ADFI, lbs 4.41 5.34 -13.7 Live Wt. F/G 2.28 2.53 -9.9 Fat Depth, in2 .71 1.18 -39.1 Fat-Free lean gain, lbs/d .74 .67 +10.5 Carcass Fat gain g/d .45 .63 -28.7 6.23 7.91 -21.2 Lean Feed Conversion (lbs Feed/lbs F.F. Lean) Test Period 60-250 lbs Mean of three European terminal crosses and four U.S. terminal crosses with either 24 or 32 pigs per genetic cross and sex. Table 4. 1995 Spring Lean Growth Trial Pork Quality Traits Pork Quality Lean European U.S. Color 2.22 2.84 Firmness 1.71 2.90 Marbling 2.00 2.74 Table 5. Genetic by Environmental Interactions for Growth Rate and Feed Intake (GxE 3) Conv. Weaning-Cont Flow SEW Three Site EUR D-YL EUR D-YL Sign. 30-51 d ADG, lb/d .72 .90 1.09 1.03 G**E**, GxE** ADFI, lb/d .99 1.16 1.43 1.51 G**GxS* 51 d - 250lb ADG, lb/d 1.69 1.81 1.87 1.87 G**E**GxE*T**ExT* * ADFI, lb/d 4.60 5.04 5.04 5.22 G**E***GxE*T**ExT * Gain/Feed .367 .361 .373 .359 Days to 250lb 181.3 171.8 163.7 164.8 G*E**GxE**T**S** 8 Table 6. LSMEANS for the Carcass Traits (G x E 3) Conv. WeaningCont. Flow SEW Three Site EUR D-YL EUR D-YL FD 10 Rib in .74 .83 .80 .86 2 Loin Eye Area, in 7.30 7.15 7.48 7.38 % Lean 52.1 50.8 51.7 50.6 Color 2.13 2.00 2.26 2.16 Marbling 1.56 1.76 1.59 1.65 Firmness 2.15 2.29 2.20 2.22 Table 7. LS means for Pork Quality Traits (GxE 3) Conv. WeaningCont. Flow SEW Three Site EUR D-YL EUR D-YL Tenderness 6.79 6.81 7.00 5.65 Juiciness 6.42 6.22 6.94 5.33 Shear force 5.47 5.64 5.03 5.67 Cooking Loss, % 32.8 28.7 31.2 29.3 IMF, g 1.57 1.94 1.95 2.22 Ph24h Drop loss 24h Hunter L* 5.55 3.62 58.1 5.64 1.43 57.2 Table 8. Breed Differences - NBS® Growth ADG BF10 lb/day inches Berkshire 1.70 1.18 Chester 1.64 1.24 White Duroc 1.78 1.02 Hampshire 1.64 .91 Landrace 1.72 1.02 Yorkshire 1.69 1.00 5.43 3.74 57.9 5.53 2.39 57.2 Carcass LMA Minolta sq in reflect 5.39 22.9 5.59 23.3 5.76 6.49 5.97 5.89 24.6 22.5 27.6 24.7 Sign G**,GxT*,S*E* E,S***, GxExS** G**,S** E*, S** G*,S** S*,ExS* Sign G*,GxE**, GxT* G***, GxE**, GxT* G**, GxExT* G***, ExS* G**,E**,S** G**E** G*** GxS1 E, GxS* Meat Quality 24h pH IMF pH % 5.84 3.24 5.82 2.93 5.72 5.54 5.61 5.68 3.97 2.35 2.31 2.24 9 Table 9. Means for Stress Carriers versus Non-Carriers Nn NN Sign LS Means ADG 2.15 2.13 NS Gain:Feed .36 .33 .01 Dressing Percent 75.3 74.4 .001 10th Rib Fat Depth .94 1.02 NS 2 LEA, in 6.64 6.43 NS Ham Lean 15.21 14.55 .01 Pork Quality Means 45 min pH 6.4 6.6 .001 24h pH 5.6 5.7 .01 Color 2.2 2.7 .001 Firmness 2.2 2.9 .001 Minolta L* 45.7 42.0 .001 a* 9.5 9.2 NS b* 7.3 5.8 .001 Drip Loss 5.2 3.4 .001 Cooking Loss 27.1 25.6 NS Lipid Content 2.0 2.3 NS 144 stress carrier and normal 242, 275, and 309 lbs Leach et al., 2000. University of IL Table 10. NBS-Stress Gene Data Trait ADG LENGTH BF10 LMA MINOLTA Phu IMF Nn-NN -0.2 lb/day 0 inches -.05 inches .37 sq in 2.2 -.01 -.34% Quality effect of one n gene None None Better Better Poorer None Poorer Table 11. Effects of Napole in Hampshire Sired Pigs. Glycolytic Potential RN (198) rn rn (32) Biopsy 238.5a 145.7b a Post Mortem 224.7 165.5b Phu 5.42a 5.52b Drip Loss 5.87 4.32 Cooking Loss 25.5a 21.2b Shear Force, kg 2.85 2.99 (Miller et al., 1998. University of Illinois) 10 Table 12. U. S. Research on the Napole Gene Napole Napole Sign Carrier Free N 56 55 Dressing % 73.8 73.6 Backfat .93 .89 Ph 24th 5.26 5.50 .001 Color 1.90 2.24 .05 Firmness 2.26 2.00 Marbling 1.54 1.78 Minolta L* 50.3 54.5 .001 Minolta a* 8.71 8.75 Minolta b* 9.13 Drip Loss 7.02 WB-shear, kg 1.54 (Hamilton et al., 2000. Univ. of Illinois) 7.28 4.67 1.78 .001 .001 .05 Table 13. Effect of Sire Breed on Ultimate pH. Sire N Ultimate pH Berkshire 60 5.61 Duroc 54 5.63 Hampshire 82 5.51 RN__ 30 5.41 rn rn 32 5.60 Feed Unfed Fed 88 83 5.62 5.55 Van Laack, Stevens, Stalder, 2001 Table 14. Warner-Bratzler shear force (kg) of pork longissimus muscle as affected by sire genetic type and storage time (LS means) Storage time, d Sire genetic type 2 7 14 Berkshire 4.95av 4.28wx 3.57yz Duroc 5.18v 4.19wx 3.58yz w x Hampshire 4.53 4.01 3.39yz v wx Non-RN carrier 4.87 4.14 3.56yz RN carrier 4.24w 3.95xy 3.27z a Standard error is 0.2. Van Laack et al., 2001 11