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J. Anim. Sci. 2003. 81:2206-2212
© 2003 American Society of Animal Science

Relationships between longissimus glycolytic potential and swine growth performance, carcass traits, and pork quality

D. N. Hamilton*, K. D. Miller*, M. Ellis*,1, F. K. McKeith* and E. R. Wilson{dagger}

* Department of Animal Sciences, University of Illinois, Urbana 61801 and and {dagger} Pig Improvement Company, Franklin, KY 42134


    Abstract
 Top
 Abstract
 Introduction
 Materials and Methods
 Results and Discussion
 Implications
 Literature Cited
 
The relationships between glycolytic potential and growth performance, carcass traits, and pork quality were investigated in a group of 72 pigs from the same genetic line. Glycolytic potential (GP) was determined on live-animal biopsy samples and postmortem samples taken from the longissimus muscle, and free glucose concentration was measured on the exudate from the longissimus muscle taken postmortem. The mean live-animal and postmortem GP and free glucose values were 201.6 µmol/g (range = 113.8 to 301.1), 149.8 µmol/g (range = 91.0 to 270.5) and 110.1 mg/dL (range = 30.0 to 406.0), respectively. Correlations between live-animal and postmortem GP and free glucose ranged from 0.47 to 0.70; however, all three measures were weakly related to growth and carcass traits (r = 0.03 to -0.22; P > 0.05). Correlations of GP and free glucose values with fresh pork quality measurements were moderate (r = 0.23 [P < 0.05] to -0.63 [P < 0.001]). Regression analysis suggested that a one standard deviation increase in live-animal and postmortem GP and free glucose resulted in an increase in L* values (0.99, 1.32, and 2.05, respectively) and drip loss (0.85, 1.10, and 1.39 percentage units, respectively), as well as a decrease in ultimate pH (0.05, 0.11, and 0.16, respectively). Correlations between GP and cooking loss and tenderness and juiciness scores ranged between 0.16 (P > 0.05) to 0.34 (P < 0.01). Free glucose concentration showed no relationship (P > 0.05) with cooking loss, tenderness, and juiciness. Regression analysis suggested that a one standard deviation increase in live-animal and postmortem GP increased cooking loss (1.26% and 1.65%, respectively) and would improve taste panel tenderness (0.54 and 0.44, respectively) and juiciness (0.40 and 0.48, respectively) scores. Increasing GP and free glucose was also associated with decreased longissimus fat and protein, and increased moisture contents (r = 0.14 [P > 0.05] to -0.45 [P < 0.001]). Overall, relationships with fresh meat quality characteristics were stronger for free glucose values than either live-animal or postmortem GP. Results from this study indicate that decreasing longissimus GP and free glucose concentrations may improve pork color and water-holding capacity.

Key Words: Glucose • Glycogen • Meat Quality • Pork


    Introduction
 Top
 Abstract
 Introduction
 Materials and Methods
 Results and Discussion
 Implications
 Literature Cited
 
Glycolytic potential (GP) is a measure of all compounds present in the muscle that can be converted into lactic acid. It is an index of the muscle’s capacity for postmortem glycolysis, and, therefore, of the potential extent of muscle pH decline after slaughter (Monin and Sellier, 1985Go). Glycolytic potential can be measured on muscle samples taken from either the live animal or from the carcass. Also, Lundstrom and Enfalt (1997)Go showed a strong relationship between longissimus GP and glucose concentrations measured in the exudate from a longissimus sample, indicating that postmortem glucose concentration may be a reliable and rapid method to predict pork quality. A major factor associated with variation in muscle GP and pork quality is the Rendement Napole gene, which has been widely investigated (Enfalt et al., 1997Go; Hamilton et al., 2000Go; Miller et al., 2000bGo). However, relationships between muscle GP and pork quality attributes across a wide range of GP values have not been examined. Such relationships are of value to understand the extent to which changes in GP will impact animal performance, carcass characteristics, and pork quality. Thus, the objective of this study was to characterize relationships of live-animal and postmortem longissimus GP and free glucose concentrations measured on muscle exudate with growth performance, and carcass and meat quality characteristics within a population of pigs that had a wide range of GP values.


    Materials and Methods
 Top
 Abstract
 Introduction
 Materials and Methods
 Results and Discussion
 Implications
 Literature Cited
 
Animals
This study was conducted at the Swine Research Center at the University of Illinois and used progeny of line 355 sires and Camborough 22 dams (PIC USA, Franklin, KY). The sires were of Hampshire ancestry, and previous research indicated that they were likely to be heterozygous for the Rendement Napole gene (Miller et al., 2000aGo). The protocol used was approved by the Institutional Animal Care and Use Committee.

Biopsy Sampling
Glycolytic potential values were determined prior to the start of the performance test to ensure that the population of pigs used in this study had the desired range of GP values. At approximately 70 kg of live weight, a total of 153 animals from this population had a longissimus muscle biopsy sample removed using spring-loaded biopsy equipment (Biotech PPB-U, Nitra, Slovakia). The depth of the cannula was set to penetrate 5.8 cm, and samples were taken from the longissimus muscle at the level of the last rib, 5 cm from the midline, from the right side of the animal. Muscle samples were trimmed of fat and skin, weighed, and immediately frozen in liquid nitrogen and subsequently freeze-dried.

Glycolytic Potential Determination
Full details of the assay procedures used to estimate GP have been described by Miller et al. (2000c)Go. In summary, GP was calculated using the formula from Monin and Sellier (1985)Go as follows:


The combination procedure used for the determination of glycogen, glucose-6-phosphate, and glucose was previously described by Keppler and Decker (1974)Go and Dalrymple and Hamm (1973)Go. Lactate analysis was carried out using Sigma Diagnostics kits for lactate (Kit 826-A, Sigma Diagnostics, St. Louis, MO) (Bergmeyer, 1974Go).

Trial Design and Performance Test
A total of 72 pigs (equal numbers of barrows and gilts) were performance tested beginning at approximately 80 kg of BW and ending at approximately 125 kg of BW. Pigs were allotted on the basis of weight to like-sex groups of four animals. During the study, pigs were housed in a controlled-environment finishing house that had partially slatted floors and provided a space allowance of 1.2 m2 per pig. Pigs had ad libitum access to water and a corn-soybean meal-based finisher diet (15.8% CP; 0.67% lysine 3,329 kcal/kg of ME) from a two-hole feeder. Pigs and feeders were weighed weekly and feed additions were recorded. Pens of pigs were taken off test and sent to harvest when their average weight was within 3 kg of the target weight of 125 kg. On the afternoon prior to harvest, the animals were weighed before loading and transported to a packing plant located approximately 280 km from the farm. At the plant, the pigs were held in lairage overnight (approximately 16 h before slaughter) without food but with access to water. Pigs were mixed in one group during transport and in the lairage. Pigs were harvested using standard commercial procedures. Pigs were electrically stunned, exsanguinated, and dehaired. Following evisceration, the head, kidneys and leaf fat were removed. Carcasses were split down the midline and placed in a chiller (4°C) approximately 60 min postmortem and were held there overnight.

Carcass and Meat Quality Measurements
At 24 h postmortem, left sides of carcasses were ribbed between the 10th and 11th ribs and fat depth (3/4 measurement) and longissimus area were recorded. The longissimus muscle from the right side of each carcass was subsequently removed, vacuum packaged, and transported on dry ice to the University of Illinois Meat Science Laboratory for further analysis.

Ultimate pH was determined on a longissimus sample (approximately 3 g) taken at the level of the l0th rib with an 8-mm cork bore and homogenized in 10 mL of distilled water. The pH of the homogenate was measured using an Orion model 720A pH meter fitted with a Ross Sure Flow 81-72 electrode (Orion Research, Boston, MA). Objective color was measured on the cut surface of the longissimus section using a Hunter LabScan Spectrocolormeter (model XE, Hunter Associates Laboratory, Inc., Reston, VA) set at the D65 and 10° angle of reflection and CIE L*, a*, and b* were then calculated (CIE, 1978Go). Three 2.5-cm-thick chops were cut from the longissimus muscle immediately posterior to the 10th rib. One chop was weighed, placed in a Whirl-pak bag (NASCO, Modesto, CA), suspended in a 4°C cooler for 48 h, and then reweighed, and drip loss percentage calculated. A second chop was trimmed of connective tissue and external fat, homogenized, placed in a Whirl-pak bag, and frozen (-20°C) for subsequent chemical analysis. The remaining chop was used for GP determination using the procedures previously described for the biopsy sample. A 10-cm section of the longissimus muscle was taken and frozen (-20°C) for a minimum of 7 d prior to shear force and taste panel evaluations.

Chemical Analysis
Fat and moisture contents were determined on the homogenized longissimus muscle sample using the procedures described by Novakofski et al. (1989)Go. Samples were oven dried (110°C for 48 h) and fat was extracted using a mixture of warm chloroform and methanol (4:1). Protein content was estimated using Kjeldahl procedures (AOAC, 1990Go).

Shear Force and Sensory Evaluation
Chops for Warner-Bratzler shear force were thawed for 24 h at 4°C and cooked on Farberware open-hearth grills (model 1 SSN, Walter Kidde, Bronx, NY) to an internal temperature of 70°C. Chops were turned mid-way through the cooking process when the internal temperature reached approximately 35°C. Temperatures were monitored using copper constantan thermocouples and a recording thermometer (Campbell Scientific, Logan, UT). Chops were weighed before and after cooking to calculate cooking loss percentage. Chops were cooled to 25°C and six 1.3-cm diameter cores were removed parallel to the muscle fibers and sheared once through the center with an Instron model 1122 Universal Testing Machine (Instron, Canton, MA) fitted with a Warner-Bratzler shear attachment. The Instron was equipped with a 10-kg load-cell and a crosshead speed of 200 mm/min. Chops for sensory evaluation were prepared and cooked using the same procedures as for shear force. Twelve panelists, consisting of faculty and graduate students at the Meat Science Laboratory, were trained according to the procedures for sensory evaluation (AMSA, 1978Go). Six of the twelve trained panelists were randomly selected for each taste panel session. Panelists evaluated juiciness, tenderness, and off-flavor intensity using a 15-cm structured line scale with anchors (0 cm = extremely dry, tough, and intense off-flavor to 15 cm = extremely moist, tender, and no off-flavor). Water was provided to panelists between samples to cleanse their palates.

Free Glucose Determination
A pipetter (Oxford Benchmate, Fisher Scientific, Pittsburgh, PA) was used to remove a 5-µL sample of the purge from the bag in which the longissimus sample used for GP determination had been stored. Samples were analyzed using a device developed to check blood glucose concentrations in diabetes patients (Accu-Chek Instant Blood Glucose Monitoring System, Boehringer Mannheim Corp., Indianapolis, IN). This process involved the addition of the 5-µL sample to a test strip and feeding the strip into the glucose analyzer.

Statistical Analysis
The PROC REG and PROC CORR procedures of SAS (SAS Inst., Inc., Cary, NC) were used to determine relationships of GP and free glucose concentrations with growth, carcass, and meat quality traits. Live-animal and postmortem GP and free glucose were used as the independent variables in the regression analysis. The initial model included linear, quadratic, and cubic terms. Subsequently, simpler models were derived from the initial model by dropping higher-order regression terms. The adequacy of simpler models was tested using a lack of fit test (Neter et al., 1985Go), and the simplest model without lack of fit, compared to the initial model, was derived for each variable.


    Results and Discussion
 Top
 Abstract
 Introduction
 Materials and Methods
 Results and Discussion
 Implications
 Literature Cited
 
Simple statistics and correlation coefficients for growth, carcass, and meat quality characteristics are presented in Tables 1Go and 2Go, respectively. The regression relationships between live-animal and postmortem GP and free-glucose are presented in Table 3Go. Regression relationships of GP and free glucose with growth, carcass, and fresh meat quality characteristics are presented in Table 4Go, and the regression relationships of GP and free glucose with palatability and chemical composition traits are presented in Table 5Go. For all variables, the linear model gave a similar fit (P > 0.05) to the data than the higher order regression models and only the linear regression relationships have been presented.


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Table 1. Simple statistics for growth, carcass, and longissimus muscle meat quality traits
 

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Table 2. Selected correlation coefficients between longissimus glycolytic potential (GP), exudate-free glucose, and growth, carcass, and longissimus meat quality traits
 

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Table 3. Regression equations between longissimus glycolytic potential (µmol/g) and exudate-free glucose (mg/dL)
 

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Table 4. Regression equations between independent (longissimus glycolytic potential [GP, µmol/g] and exudate-free glucose [mg/dL]) and dependent (growth performance, carcass traits, and longissimus muscle quality characteristics) variables
 

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Table 5. Regression equations between independent (longissimus muscle glycolytic potential [GP, µmol/g] and exudate-free glucose [mg/dL]) and dependent (longissimus meat quality characteristics) variables
 
Relationships of Glycolytic Potential with Free Glucose
Live-animal GP, postmortem GP, and free glucose values ranged from 113.8 to 301.1 µmol/g fresh tissue, from 91.0 to 270.5 µmol/g, and from 30.0 to 406.0 mg/dL, respectively (Table 1Go). The mean and range of live-animal and postmortem GP values reported in the current study were similar to values reported in other papers that used animals with a wide range of GP values (Hamilton et al., 2000Go, 2002Go; Miller et al., 2000aGo). In addition, the mean free glucose values reported in the current study were similar to values reported by Miller et al. (2000a)Go.

Live-animal GP values were correlated with postmortem GP and free glucose concentrations of the exudate (Table 2Go). However, the strongest correlation was between the postmortem measurement of GP and free glucose concentrations. This is not surprising given that postmortem GP and free glucose were measured on the same sample. Results from the regression analysis indicated that a one standard deviation increase in live-animal GP (44.88 µmol/g) resulted in a 26 µmol/g increase in postmortem GP and a 13 mg/dL increase in free glucose (Table 3Go). Additionally, a one standard deviation increase in postmortem GP (36.62 µmol/g) resulted in a 11 mg/dL increase in free glucose. Differences between live-animal and postmortem GP values are in agreement with other studies (Miller et al., 2000aGo,cGo) and are not unexpected given the many factors that can affect muscle metabolism and glycogen levels from the time the animal leaves the farm through to slaughter, including transport time and distance, time off feed, lairage time, and pig handling procedures.

Relationships of Glycolytic Potential and Free Glucose with Growth and Carcass Traits
Correlations between GP and free glucose and ADG and 10th rib backfat were weak (r <= -0.22, P > 0.05) (Table 2Go). Interestingly, longissimus muscle area was moderately related (P < 0.05) to postmortem GP but not (P > 0.05) to live-animal GP or free glucose. Regression analysis suggested that a one standard deviation increase in postmortem GP will result in a 1.76 cm2 decrease in longissimus muscle area (Table 4Go). Other studies have reported reduced backfat depths in high- vs. low-GP animals (LeRoy et al., 1996Go; Enfalt et al., 1997Go), but there are no reports of a relationship between muscle GP and longissimus muscle area.

Relationships of Glycolytic Potential and Free Glucose with Pork Quality
Relationships of live-animal GP, postmortem GP, and free glucose with longissimus objective color, drip loss, ultimate pH, and cooking loss were moderate to strong (range 0.23 to -0.63; P < 0.05; Table 2Go). Studies comparing populations with high and low GP values, resulting from the segregation of the RN gene, have demonstrated paler color and reduced water-holding capacity for the high-GP population (Hamilton et al., 2000Go; 2002Go; Miller et al., 2000aGo). Live-animal GP, postmortem GP, and free glucose were linearly related with L*, drip loss, cooking loss, and ultimate pH (Tables 4Go and 5Go). Thus, a one standard deviation increase in GP (live animal and postmortem) and free glucose was associated with an increase in longissimus L* values of 0.99, 1.32, and 2.05; an increase in drip loss of 0.85, 1.10, and 1.39 percentage units; an increase in cooking loss of 1.26, 1.65, and 1.88 percentage units; and a decrease in ultimate pH of 0.05, 0.11, and 0.16, respectively. This suggests that fresh meat quality traits generally deteriorated with increasing longissimus GP and/or free glucose concentration of the exudate. Somewhat surprisingly, free glucose content had the strongest relationships with meat quality parameters, whereas live-animal GP had the weakest relationships. It is reasonable to expect that both of the postmortem measurements would have stronger relationships to meat quality parameters than the measurement taken before harvest because they better reflect the metabolites present within the muscle at the completion of rigor mortis.

Correlations with taste panel tenderness and juiciness scores were moderate for live-animal and postmortem GP values, but free glucose exhibited much weaker relationships with these traits (r <= 0.14; P > 0.05; Table 2Go). Taste panel juiciness and tenderness scores increased linearly (P < 0.05) as live-animal and postmortem GP increased (Table 5Go), indicating an improvement in palatability with increasing GP. Regression relationships suggest that a one standard deviation increase in live-animal and postmortem GP resulted in an improvement in taste panel tenderness (0.54 and 0.44, respectively) and juiciness (0.40 and 0.48, respectively) scores. Other authors have also shown an improvement in palatability traits with increasing muscle GP (Lundstrom et al., 1996Go; Miller et al., 2000aGo). Overall, the correlations between postmortem GP and the palatability traits were similar to those reported by Huff-Lonergan et al. (2002)Go but lower than those reported by van Laack and Kauffman (1999)Go.

Longissimus protein and moisture were moderately correlated to live-animal and postmortem GP, and free glucose; however, the correlations between live-animal GP, postmortem GP, free glucose, and longissimus muscle fat content were weaker (Table 2Go). Results from the regression analysis suggested that a one standard deviation increase in live-animal GP, postmortem GP or free glucose resulted in a 0.40, 0.37, and 0.33 percentage unit decrease in logissimus protein content, respectively, and a corresponding increase in longissimus moisture content of 0.27, 0.26, and 0.25 percentage units, respectively. Similar relationships have been documented by other researchers (Enfalt et al., 1997Go; Sutton, 1997Go; Miller et al., 2000bGo), and indicate that as glycogen reserves within the muscle increased, glycogen bound moisture also increased.

Relationships of Longissimus pH with Pork Quality
Ultimate pH was strongly related to longissimus muscle L* values and drip loss (Table 2Go). These correlations are similar to the findings of Bidner (1999)Go and van Laack and Kauffman (1999)Go, who both reported correlations greater than 0.70 between longissimus ultimate pH and L* values and drip loss. Longissimus ultimate pH was negatively correlated with cooking loss and shear force. Thus, lower ultimate pH values were associated with deterioration of fresh pork quality in terms of paler muscle with reduced water-holding capacity. These results are not surprising given that as muscle pH approaches the isoelectric point the water holding capacity of the muscle sharply decreases (Price and Schweigert, 1987Go).


    Implications
 Top
 Abstract
 Introduction
 Materials and Methods
 Results and Discussion
 Implications
 Literature Cited
 
Relationships evaluated in this trial suggest that decreasing glycolytic potential across the wide range studied would result in improvements in the color and water-holding capacity, but also in deteriorations in the juiciness and tenderness of fresh pork. Free glucose concentration in the exudate from longissimus muscle was more strongly related to fresh pork quality than glycolytic potential and is quicker and less expensive to measure.

1 Correspondence: 216 Animal Sciences Laboratory, 1207 W. Gregory Dr. (phone: 217-333-6455; fax: 217-333-7861; E-mail: mellis7{at}uiuc.edu).

Received for publication December 3, 2002. Accepted for publication May 14, 2003.


    Literature Cited
 Top
 Abstract
 Introduction
 Materials and Methods
 Results and Discussion
 Implications
 Literature Cited
 


AMSA. 1978. Guidelines for Cookery and Sensory Evaluation of Meat. Am. Meat Sci. Assoc.—Natl. Livest. and Meat Board, Chicago, IL.

AOAC. 1990. Official Methods of Analysis. 15th ed. Assoc. Offic. Anal. Chem., Arlington, VA.

Bergmeyer, H. U. 1974. Methods of Enzymatic Analysis. Academic Press, New York.

Bidner, B. S. 1999. The effects of RN genotype, feed withdrawal prior to slaughter, lysine-deficient diet, and sodium tripolyphosphate pumping on pork quality and sensory characteristics. M.S. Thesis, Univ. of Illinois, Urbana-Champaign.

CIE. 1978. Recommendations on Uniform Color Spaces—Color Differences Equations, Psychrometric Color Terms. Supplement No. 2, CIE Publication No. 15 (E1.3.1).

Dalrymple, R. H., and R. Hamm. 1973. A method for the extraction of glycogen and metabolites from a single muscle sample. J. Food Technol. 8:439–444.

Enfalt, A. C., K. Lundstrom, A. Karlsson, and I. Hansson. 1997. Estimated frequency of the RN- allele in Swedish Hampshire pigs and comparison of glycolytic potential, carcass composition, and technological meat quality among Swedish Hampshire, Landrace, and Yorkshire pigs. J. Anim. Sci. 75:2924–2935.[Abstract/Free Full Text]

Hamilton, D. N., M. Ellis, K. D. Miller, F. K. McKeith, and D. F. Parrett. 2000. The effect of the Halothane and Rendement Napole genes on carcass and meat quality characteristics of pigs. J. Anim. Sci. 78:2862–2867.[Abstract/Free Full Text]

Hamilton, D. N., M. Ellis, K. D. Miller, F. K. McKeith, A. D. Higgerson, and J. E. Beever. 2002. Comparison of the glycolotyic potential and DNA-based test for predicting Rendement Napole genotype. J. Anim. Sci. 80(Suppl. 2):45. (Abstr.)[Abstract/Free Full Text]

Huff-Lonergan, E., T. J. Baas, M. Malek, J. C. M. Dekkers, K. Prusa, and M. F. Rothschild. 2002. Correlations among selected pork quality traits. J. Anim. Sci. 80:617–627.[Abstract/Free Full Text]

Keppler, D., and K. Decker. 1974. Glycogen determination with amyloglucosidase. Pages 127–131 in Methods of Enzymatic Analysis. Vol. III. Verlag Chemie Weinheim, Academic Press Inc., New York.

LeRoy, P., G. Monin, J. M. Elsen, J. C. Cartiez, A. Talmant, B. Lebret, L. Lefaucheur, J. Mourot, H. Juin, and P. Sellier. 1996. Effect of the RN genotype on growth and carcass traits in pigs. Page 311 in Proc. 47th Mtg. Eur. Assoc. Anim. Prod. Vol. 2. Lillehammer, Norway.

Lundstrom, K., A. Anderson, and I. Hansson. 1996. Effect of the RN gene on technological and sensory meat quality in crossbred pigs with Hampshire as terminal sire. Meat Sci. 42:145–153.

Lundstrom, K., and A. C. Enfalt. 1997. Rapid prediction of RN phenotype in pigs by means of meat juice. Meat Sci. 45:127–131.

Miller, K. D., M. Ellis, B. S. Bidner, F. K. McKeith, and E. R. Wilson. 2000a. Porcine longissimus glycolytic potential level effects on growth performance, carcass, and meat quality characteristics. J. Muscle Foods 11:169–181.

Miller, K. D., M. Ellis, F. K. McKeith, B. S. Bidner, and D. J. Meisinger. 2000b. Frequency of the Rendement Napole RN- allele in a population of American Hampshire pigs. J. Anim. Sci. 78:1811–1815.[Abstract/Free Full Text]

Miller, K. D., M. Ellis, D. S. Sutton, F. K. McKeith, and E. R. Wilson. 2000c. Effects of live-animal sampling procedures and sample storage on the glycolytic potential of porcine longissimus muscle samples. J. Muscle Foods 11:61–67.

Monin, G., and P. Sellier. 1985. Pork of low technological quality with a normal rate of muscle pH fall in the immediate post-mortem period: The case of the Hampshire breed. Meat Sci. 13:49–63.

Neter, J., W. Wasserman, and M. H. Kutner. 1985. Applied Linear Statistical Models: Regression, Analysis of Variance, and Experimental Designs. 2nd ed. Irwin, Homeland, IL.

Novakofski, J. S., S. Park, P. J. Bechtel, and F. K. McKeith. 1989. Composition of cooked pork chops: effect of removing subcutaneous fat before cooking. J. Food Sci. 54:15–17.

NPPC. 1991. Procedures to Evaluate Market Hogs. 3rd ed. Natl. Pork Prod. Council, Des Moines, IA.

Price, J. F., and Schweigert, B. S. 1987. The Science of Meat and Meat Products. 3rd ed. Food and Nutrition Press, Westport, CT.

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van Laack, R. L., and R. G. Kauffman. 1999. Glycolytic potential of red, soft, exudative pork longissimus muscle. J. Anim. Sci. 77:2971–2973.[Abstract/Free Full Text]


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