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Colorado State University4, Fort Collins 80523 and U.S. Department of Agriculture, ARS, Clay Center, NE 68933
| Abstract |
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Key Words: Beef Cattle Breeds Carcass Composition Meat Quality Muscle Fibers
| Introduction |
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There is considerable variation in expression of double muscling within the Piedmontese breed; cattle generally are divided into three classes based on phenotype: normal, intermediate and hypertrophied. Masoero (1982)
reported that the registered herd makeup was 1% normal, 42% intermediate and 57% hypertrophied for males and 12% normal, 68% intermediate and 20% hypertrophied for females. Bulls classified as hypertrophied are preferred for use as herd sires, whereas cows classified as intermediate are considered to be most productive and, therefore, are preferred by breeders (Sartore and Chiappone, 1982
).
Piedmontese cattle recently were introduced into the U.S. Because of their superior muscularity and leanness, Piedmontese sires may be particularly well suited for use in crossbreeding systems designed to produce high-cutability market cattle. However, the performance of Piedmontese in U.S. production systems has not been documented. This study was conducted to compare carcass characteristics of F1 Piedmontese steers with those of contemporary steers sired by Gelbvieh and Red Angus bulls.
| Materials and Methods |
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Forty-five male calves were produced by matings of four Piedmontese (P) classified as hypertrophied, five Gelbvieh (G) and four Red Angus (RA) sires to British and Continental crossbred dams. Cows classified as British (n = 26) were mostly Hereford x Black or Red Angus, although four straightbred Hereford cows also were included in this group. Continental crossbred cows (n = 19) were 25 to 50% Simmental (S) or Charolais (C) and 25 to 75% Hereford (H) or Red Angus. Sixteen of the Continental crossbred cows were three-way crosses (S x H x RA or C x H x RA). British dams produced 9 RA, 6 P and 11 G steers; continental dams produced 6 RA, 9 P and 4 G steers. The choice of sire breeds permitted comparison of P (a breed selected for double muscling) with G (a European dual-purpose breed) and RA (a British breed selected for conventional beef characteristics).
The calves were born February through May of 1984 at the Eastern Colorado Research Center near Akron, Colorado. At birth, the calves were individually identified and birth dates were recorded. At branding (May 15, 1984), all calves were vaccinated, branded with a hot iron, castrated and dehorned (if necessary). All calves nursed their dams under similar pasture conditions until weaning (October 1, 1984). After weaning, the steers were backgrounded on corn silage. On January 9, 1985, the steers, none of which would be classified as double-muscled, were started on a high-concentrate finishing diet and fed for 124, 166 or 208 d prior to slaughter. During the finishing period, the steers had ad libitum access to cracked corn and were fed 4.5 kg com silage (wet weight) and .45 kg protein supplement per head daily.
Carcass Data Collection and Sample Preparation.
On each of the three slaughter dates, five steers from each sire-breed group were weighed individually (unshrunk) and transported 145 km to a commercial packing facility. Immediately following slaughter, hot carcass weight was recorded and the carcasses were chilled at approximately 2°C. Following a 24-h chill, each carcass was ribbed between the 12th and 13th ribs and factors used to determine USDA quality grade (carcass maturity and marbling) and yield grade (subcutaneous fat thickness, ribeye area and estimated kidney, pelvic and heart fat percentage) were obtained by a panel of three experienced carcass evaluators (USDA, 1989
). Additionally the panel assigned scores to each carcass for color, texture and firmness of lean based on visual and physical characteristics of the surface of the M. longissimus at the 12th-13th rib interface. The following day, one side of each carcass was transported to the Colorado State University meat laboratory.
Within 96 h postmortem, each carcass side was fabricated into primal and subprimal cuts and each cut was dissected into muscle, fat and bone (bone plus cartilage and major tendons) using procedures described by Abraham et al. (1980)
. Individual weights were recorded for each cut and its dissected components. Component weights corresponding to a particular tissue type (muscle, bone or fat) were aggregated to provide the weights of muscle, bone and fat in the entire side. Additionally, weights for individual fat components from each side were aggregated to provide the weights of the following fat depots: 1) subcutaneous fat -external fat covering the peripheral musculature, including scrotal fat and fat beneath the M cutaneous truncii and M. cutaneous omobrachialis; 2) intermuscular fat - fat between the muscles and between the musculature and bones or tendons; and 3) internal fat – fat within the body cavity, including the kidney, pelvic and heart depots together with fat removed from the internal surfaces of the ribs. Carcass side weight was expressed as the aggregate of all component weights.
Immediately following carcass dissection, a portion of the M. longissimus was removed from each short loin immediately posterior to the 12th rib. These samples were trimmed of all subcutaneous fat, vacuum-packaged individually, placed in an insulated shipping container and transported to the U.S. Meat Animal Research Center at Clay Center, Nebraska for proximate analysis, muscle fiber typing and intramuscular collagen assays. In addition, the 6th to 8th rib section of the M. longissimus from each carcass was vacuum-packaged, aged at 5°C until the 11th d postmortem and stored at –20°C to be used for taste panel testing.
Proximate Analysis and Collagen Assays.
Triplicate tissue samples (2 g) of the 12th rib section of the longissimus muscle were weighed, dried at 100°C for 12 h and reweighed to determine moisture content. The dried samples then were extracted for 48 h using diethyl either in a soxhlet extraction apparatus. The extracted samples were redried and weighed to determine lipid content (AOAC, 1980
).
Freeze-powdered longissimus samples (4 g) were heated for 70 min at 77°C in .25-strength Ringers solution (Hill, 1966
). Supernatant fluid and residue fractions were hydrolyzed in 6 N HCl for 20 h at 115°C. Following neutralization, hydroxyproline content of each hydrozylate was determined using spectrophotometric methods described by Bergman and Loxley (1963)
. Collagen content was calculated by multiplying the hydroxyproline content of the residue by 7.25 and that of the supernatant fluid by 7.52 (Cross et al., 1973
). Percentage of soluble (heat labile) collagen was calculated by dividing the collagen content of the supernatant fluid by the collagen content of the entire sample.
Palatability Determinations.
Sensory evaluation was performed by a 10-member descriptive panel. Panelists were selected and trained in accordance with the AMSA Guidelines for Cooking and Sensory Evaluation of Meat (AMSA, 1978
).
Three steaks (2.54 cm thick) were removed from the frozen 6th to 8th rib section of the longissimus from each carcass. The steaks were thawed at 2°C and broiled on Farberware Open-Hearth broilers to an internal temperature of 70°C (monitored using copper-constantan thermocouples and a recording potentiometer). The 8th rib steak was used for sensory evaluation; the 6th and 7th rib steaks were used for subsequent shear force measurements.
Upon reaching the desired internal temperature, steaks for sensory evaluation were removed from the broiler and portioned into sections of uniform dimensions (approximately 1.3 cm x 1.3 cm x 1.9 cm). The warm sections were selected randomly and served immediately to the panel. Panelists assigned scores to each sample for juiciness, myofibrillar tenderness, connective tissue amount and flavor intensity using 8-point, structured rating scales.
Steaks for shear force determinations were cooled to 20°C and a minimum of six 1.27-cm cores were removed parallel to the longitudinal orientation of the muscle fibers for Warner-Bratzler shear force measurements.
Muscle Fiber Type Determinations.
Histological samples were removed from the medial, central and lateral sections of the 13th rib section of the longissimus muscle. The samples were frozen in liquid nitrogen, wrapped in aluminum foil and stored at –63°C. Transverse sections (10 µm thick) of each muscle sample were cut using a cryostat and stained for alkali-stable ATPase using procedures described by Guth and Samaha (1970)
. Serial sections were stained for succinate dehydrogenase activity according to procedures outlined by Troyer (1980)
. Photomicrographs were obtained for each sample and individual muscle fibers were counted and classified as red, intermediate or white based on staining intensity. The mean area of 10 fibers of each type was determined using a Bioquant particle size analyzer. Data from the three muscle locations were averaged prior to statistical analysis.
Statistical Methods.
The data were analyzed using a least squares model that included the fixed effects of sire breed and dam breed-type. Additionally, linear regressions of traits on initial age (age at the onset of the finishing period) and days on feed were included in the model and variation due to the interaction between the days on feed regression and sire breed was partitioned. Subclass days on feed regressions were used to adjust the means for an additional comparison of the breed groups at a common degree of marbling using procedures outlined by Koch et al. (1979)
. When F-tests for sire breed were significant, subclass means were compared using Tukeys w procedure (Steel and Torrie; 1960
).
| Results |
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Mean squares for slaughter weight and estimated degree of maturity at slaughter are presented in Table 1
. Age-constant least squares means for these traits, corresponding to the effects of sire breed, are provided in Table 2
.
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Breed type of dam was a significant (P < .05) source of variation in slaughter weight, but it did not affect (P > .05) estimated degree of maturity at slaughter (Table 1
). Steers produced by Continental crossbred dams were heavier than those produced by British dams (528.5 kg vs 502.2 kg). Slaughter weight and estimated degree of maturity increased (P > .05) with increased time on feed (Table 1
).
Dressing Percentage.
Least squares means for dressing percentage (Table 2
) showed that P-sired steers had 1.2 and 1.6% higher dressed yields than G- and RA-sired steers, respectively. However, due to the limited number of animals used in this study, these differences in dressing percentage were not of sufficient magnitude for statistical significance (P = .15). Dressing percentage was not significantly affected by dam breed type or days on feed (Table 1
).
In a review of double muscling and its effects on meat production traits, Boccard (1981)
concluded that dressing percentages for homozygous double-muscled animals normally are about 5% higher than those for normal animals of the same breed, sex and weight. Boccard (1981)
attributed this advantage in dressing percentage to hypotrophy of several major internal organs, including the spleen, liver and digestive tract. The weights of these organs have been shown to be from 14 to 37% lower for double-muscled than for normal animals (Boccard, 1981
). Thiessen and Rollins (1982)
compared British crossbred calves that were heterozygous for double muscling with normal calves from similar matings. In their study, heterozygous bulls and heifers had 1.7% higher dressing percentages than did normal bulls and heifers.
Information concerning dressed yields of Piedmontese F1 cattle is limited. Siccardi (1973)
reported results from three small crossbreeding studies in which Piedmontese sires were mated to Dutch Friesian, Brown Swiss and Valdostana P.R. dams. Progeny (male and female) from these matings were compared to purebred progeny (male and female) of the three dam breeds. The comparisons of F1 Piedmontese with Dutch Friesian (DF) and Valdostana P.R. (V) involved calves slaughtered at very light weights (F1 = 174.3 kg vs DF = 163.7 kg and F1 = 206.7 kg vs V = 184.3 kg). Compared to DF and V calves, the F1 calves had 4.3 and 4.2 higher dressing percentages, respectively. In the third comparison, F1 Piedmontese (slaughtered at 492 kg) had 9.9% higher dressed yields than did purebred Brown Swiss (slaughtered at 434 kg).
Bonsembiante et al. (1975)
compared dressed yields of crossbred bulls (500 kg) produced by mating Red Pied, Piedmontese, Romagnola, Chianina and Marchigiana bulls to Brown Alpine and Friesian cows. In their study, F1 Piedmontese bulls had from 1.4 to 2.1% higher (P < .01) dressing percentages than did F1 bulls from the other four sire breeds. These differences in dressed yield are similar in magnitude to the mean differences observed between steers sired by P bulls and steers sired by G or RA bulls in our study.
Carcass Grade Traits.
Mean squares for carcass yield grade traits are presented in Table 1
. Breed of sire was a significant source of variation in adjusted fat thickness, longissimus area and yield grade but had no effect on hot carcass weight or estimated percentage of kidney, pelvic and heart fat. Fat thickness was highest (P < .05) for RA-, intermediate for G-and lowest (P < .05) for P-sired steers (Table 2
). Sire breed groups ranked in the reverse order (P > G > RA) for longissimus area. Differences in yield grade (RA > G > P) reflected the effects of sire breed on fat thickness and longissimus area (e.g., the P-sired steers were leanest and had the largest longissimus areas and therefore had the lowest numerical yield grades). Effects of dam breed on carcass yield grade traits were not significant (Table 1
). The significant effect of days on feed for fat thickness, longissimus area, hot carcass weight and kidney, pelvic and heart fat percentage (Table 1
) reflected an increase (P < .05) in each trait with increased time on feed.
Results for carcass quality grade traits are presented in Tables 1
and 2
. Significant effects of sire breed, dam breed type, days on feed and the interaction between sire breed and days on feed were observed for marbling score (Table 1
). On an age-constant basis, steers sired by RA bulls had higher (P < .05) marbling scores than G- and P-sired steers (Table 2
); marbling scores for the latter two groups (G and P) were not different (P > .05). Steers out of B dams had higher (P < .05) marbling scores than those produced by Cx dams (British = 432.5, Cx = 398.4).
Marbling score increased as time on feed increased. Moreover, tests of significance for differences among subclass regression coefficients revealed that RA-sired steers responded to increased feeding time (days) with greater (P < .05) increases in marbling score (numerical value) than either of the other two sire-breed groups (bRA = 1.32, bP = .80, bG = .71). Because of this difference in rate of marbling deposition, RA-sired steers required fewer days on feed to deposit a "small" degree of marbling (the minimum amount of marbling required for A maturity carcasses to grade Choice). In fact, RA-sired steers already had attained a "small" degree of marbling by the first slaughter date (124 d on feed). Steers sired by P and G bulls required approximately 170 and 190 d on feed, respectively, to attain a "small" degree of marbling. Carcass maturity increased (P < .01) as days on feed increased but was unaffected by either sire breed or dam breed type (Table 1
).
West et al. (1973)
compared carcass grade traits of heterozygous double-muscled steers with those of normal steers. In their study, the two genotypes did not differ in marbling score or USDA quality grade; however, heterozygous double-muscled steers had significantly less fat thickness, larger longissimus areas and higher estimated cutability (lower yield grades) than normal steers. Thiessen and Rollins (1982)
reported similar differences in yield grade traits among carcasses produced by heterozygous double-muscled and normal bulls and heifers, except that in their study differences in fat thickness were not significant. Carroll et al. (1978)
compared carcass quality grade traits of heterozygous double-muscled heifers and bulls with those of normal heifers and bulls. Heifer carcasses of the normal genotype had more marbling and higher quality grades than carcasses produced by heterozygous heifers, whereas carcasses from heterozygous and normal bulls did not differ in marbling score or quality grade.
Liboriussen (1982)
compared carcass traits of young crossbred bulls produced by double-muscled (Piedmontese and Belgian Blue), "muscular" Charolais and normal (Angus, Gelbvieh, South Devon, Braunvieh and West Flemish Red) sires. Bulls produced by double-muscled sires had larger longissimus areas and less fat thickness than did bulls produced by normal sires. Bulls produced by Piedmontese and Belgian Blue sires had similar values for longissimus area and fat thickness. Charolais crosses had intermediate values for longissimus area but were similar to double-muscled crosses with respect to fat thickness. Bonsembiante et al. (1975)
found that F1 Piedmontese bulls had larger longissimus areas than did F1 bulls produced by Red Pied, Romagnola, Chianina and Marchigiana sires. No information concerning carcass quality grade traits of Piedmontese cattle could be found.
Carcass Composition and Partitioning of Carcass Fat.
Results from analysis of variance for proportions of separable carcass components are presented in Table 3
. Percentages of separable muscle, bone and fat and carcass muscle-to-bone ratio were influenced by sire breed, but not by dam breed type.
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Differences among sire groups in percentages of separable bone (G > P) and muscle (P > G > RA) reflected the variation in fatness and muscularity discussed above. Bone percentage normally is inversely related to fat percentage and muscle-to-bone ratio, whereas muscle percentage is negatively correlated with fat percentage and positively correlated with muscle-to-bone ratio (Berg and Butterfield, 1966
). Due to a superior combination of leanness and muscularity, P-sired steers had 2.4% and 6.5% higher yields of separable muscle than did G- and RA-sired steers, respectively.
Significant effects of days on feed observed for percentages of separable carcass components (Table 3
) reflected changes in carcass composition associated with variation in length of the concentrate feeding period. Increased time on feed was associated with an increase in separable fat percentage and decreases in separable muscle and bone percentages. The fact that muscle-to-bone ratio was not affected (P > .05) by length of the feeding period (Table 2
) indicated that the primary influence of days on feed was exerted on the fat component of the carcass. Moreover, the lack of significance for the interaction between sire breed and days on feed indicated that compositional changes during finishing occurred at similar rates for P-, and G- and RA-sired steers.
Siccardi (1973)
reported a limited amount of information from three different crossbreeding studies concerning carcass composition of F1 Piedmontese cattle. These studies, which were described earlier, involved comparisons of F1 Piedmontese with purebred Dutch Friesian, Valdostana and Brown Swiss calves. Carcasses from the F1 Piedmontese consistently yielded higher percentages of muscle (from 4.6 to 5.7% higher) and lower percentages of fat (from .7 to 1.5% lower) and bone (from 2.8 to 5.3% lower) than did their purebred contemporaries. Muscle-to-bone ratios were from 1.1 to 2.3 units higher for the F1 calves.
Bonsembiante et al. (1975)
compared carcass composition of F1 bulls produced by Piedmontese (P), Red Pied (RP), Chianina (C), Marchigiana (M) and Romagnola (R) sires at a constant slaughter weight of 500 kg. Carcasses produced by F1 Piedmontese yielded more (P < .01) lean (P = 71.97%, RP = 63.15%, C = 62.91%, M = 64.63%, R = 64.12%) and less (P < .05) fat (P = 12.47%, RP = 21.67%, C = 20.08%, M = 19.67%, R = 19.35%) than carcasses produced by the four other breed groups. In a similar study, Borghese et al. (1978)
compared carcass composition yields of F1 bulls produced by mating Piedmontese (P), Chianina (C), Marchigiana (M), Limousin (L) and Charolais (Ch) bulls to Friesian cows. Piedmontese-sired bulls had the highest muscle percentage (P = 71.35%, C = 66.58%, M = 68.18%, L = 67.55%, Ch = 66.56%) and the highest muscle-to-bone ratio (P = 3.94, C = 3.07, M = 3.32, L = 3.52, Ch = 3.32).
Variance components and least squares means for relative proportions of separable carcass fat partitioned into the subcutaneous, intermuscular and internal depots are presented in Tables 2
and 3
. Sire breed had no effect (P > .05) on partitioning of separable carcass fat, indicating that the differences observed for fat thickness and percentage of total separable fat were associated with differences in overall degree of carcass fatness and not with differences in partitioning of fat into the different carcass depots. Other reports of fat partitioning data for F1 Piedmontese cattle could not be found.
Palatability Attributes, Collagen Content and Proximate Composition of the Longissimus.
Sire breed was a significant source of variation in shear force and sensory panel ratings for myofibrillar tenderness, flavor intensity and connective tissue amount (Table 4
). Longissimus steaks from carcasses produced by P- and RA-sired steers received higher (P < .05) ratings for myofibrillar tenderness, flavor intensity and connective tissue amount than did steaks produced by G-sired steers (Table 5
). Additionally, steaks from RA-sired steers had lower shear force values than did steaks from steers by G sires. Longissimus samples from steers in the three sire breed groups did not differ (P > .05) in intramuscular collagen content or moisture percentage. However, longissimus samples from RA-sired steers had the highest percentage of intramuscular lipid (Table 5
). Sensory panel ratings for myofibrillar tenderness and connective tissue amount increased and shear force values decreased as time on feed increased (Table 4
). Collagen content of the longissimus was not affected (P > .05) by days on feed; however, percentage of moisture decreased (P < .01) and percentage of intramuscular lipid tended to increase (P = .10) with increased time on feed (Table 4
).
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West et al. (1973)
found that steaks from the longissimus and semimembranosus muscles of heterozygous double-muscled steers were more tender than corresponding steaks from normal steers. Carroll et al. (1978)
compared taste panel evaluations of tenderness for longissimus and semimembranosus samples from heterozygous double-muscled and normal heifers and bullocks. They found that, among bullocks, meat produced by heterozygous double-muscled animals was more tender whereas among heifers, meat from normal animals was more tender.
Borghese et al. (1978)
compared tenderness of longissimus samples from F1 bulls sired by Chianina, Piedmontese, Marchigiana, Limousin and Charolais bulls. Samples from F1 Piedmontese had the lowest Warner-Bratzler shear force values. Shear force values for the other breed groups were similar.
Muscle Fiber Types.
Sire breed was a significant source of variation in percentages of intermediate and white muscle fibers and in cross-sectional area of red muscle fibers in the longissimus muscle (Table 6
). Steers sired by P bulls had the highest (P < .05) percentage of white fibers, the lowest (P < .05) percentage of intermediate fibers and the smallest (P < .05) areas of red muscle fibers. Fiber profiles and sizes for G- and RA-sired steers were similar (Table 5
). Areas of red, intermediate and white muscle fibers increased (P < .05) but the percentage of intermediate fibers decreased as time on feed increased (Table 5
).
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West (1974)
reported data comparing fiber characteristics of homozygous double-muscled heterozygous double-muscled and normal females at 454 kg live weight. In the semimembranosus muscle, homozygous double-muscled heifers had a lower percentage of intermediate fibers, which was attributed to greater conversion of intermediate to white fibers.
Our findings generally are consistent with those of previous studies. In the present study F1 Piedmontese steers had a higher percentage of white fibers and a lower percentage of intermediate fibers compared with the other two sire breed groups. Moreover, the percentage of intermediate fibers decreased for all three breed groups as the steers became older. Greater conversion of intermediate to white fibers for P steers during the growth and finishing periods may be a plausible explanation for their different muscle fiber type profiles.
Comparisons at a Constant Marbling Score.
Least squares means for certain traits adjusted linearly to a constant degree of marbling (Small18) are provided in Table 7
. Because most cattle currently produced in the U.S. are slaughtered at a low Choice quality grade endpoint, these data have practical implications.
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The most exhaustive comparison of sire breeds used in U.S. crossbreeding systems is the Germ Plasm Evaluation Program (GPE) at the U.S. Meat Animal Research Center. Koch et al. (1976
, 1979)
compared F1 steers sired by bulls representing various British (Angus, Hereford, Red Poll, South Devon) and continental European (Charolais, Chianina, Gelbvieh, Limousin, Maine Anjou, Simmental) breeds at a constant marbling score (small). At the same degree of marbling, G and P steers in the present study had carcass weights similar to those of the continental-European crosses in the GPE studies. However, the G and P steers in our study were leaner and had larger ribeyes and lower numerical yield grades than the GPE steers. The RA steers in our study had carcass weights that were similar to the British crosses in the GPE studies, but again, our steers were much leaner and had larger ribeyes and lower yield grades. The GPE studies demonstrated very slight breed differences in muscle-to-bone ratio and no effect of breed on meat palatability, which is in contrast to our findings.
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| Footnotes |
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2 Present address: Heritage Lite Meat Corporation, 5147A 69th, Lubbock, TX 79424. ![]()
3 Present address: Bryan Foods, P.O. Box 1177, West Point, MS 39773. ![]()
Received for publication May 19, 1989. Accepted for publication August 21, 1989.
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