Influence of dietary protein and recombinant porcine somatotropin administration in young pigs: III.

Muscle fiber morphology and shear force M. B. Solomon, T. J. Caperna, R. J. Mroz and N. C. Steele J Anim Sci 1994. 72:615-621.

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Influence of Dietary Protein and Recombinant Porcine Somatotropin Administration in Young Pigs: 111. Muscle Fiber Morphology and Shear Force112
M. B. Solomon*, T. J. Capernat, R. J. Mroz*, and N. C. Steelet
*Meat Science Research Laboratory and +Nonruminant Animal Nutrition Laboratory, Beltsville Agricultural Research Center, ARS, USDA, Beltsville, MD 20705-2350

ABSTRACT:

Sixty crossbred barrows (average 30 kg) were used in a 5 x 2 factorial treatment array to examine interactions between dietary protein concentration (11, 15, 19, 23, or 27% CP) and recombinant porcine somatotropin (rpST: 0, excipient buffer vs 100 pgkgl.d-l) for 42 d on muscle fiber morphology and meat tenderness. Diets were isocaloric (3.8 Mcal of DE/kg) and of equal lysine (4.9 g/Mcal of DE) achieved by diluting soybean meal with cornstarch and by addition of crystalline lysine. Dosage of rpST and feed intake (80% of predicted ad libitum) were adjusted weekly. Four muscles (longissimus = LM; semimembranosus = SM; semitendinosus = ST; triceps brachii = TB) were evaluated. Percentages of muscle for fiber types ( OR, aR, CYW) the LM, SM, and TB were not influenced by rpST treatment. More aR and

fewer CYW fibers were found in the ST muscle of rpSTtreated pigs. No interactions were observed between rpST treatment and dietary protein for muscle fiber type distribution. Dietary protein had no consistent influence on the distribution of muscle fiber types in all four muscles. Area of fibers generally increased in rpST-treated pigs compared with controls when diets contained 19% or more CP. The LM shear force was increased ( 13%) by rpST treatment for chops frozen after 5 d of storage in the cooler, but not in those chops frozen within 1.5 h postmortem. Dietary protein had a variable influence on tenderness. These data indicate that muscle fiber growth (hypertrophy) in pigs is positively influenced by rpST treatment. Marginal dietary protein intake reduces muscle fiber growth responses to rpST.

Key Words: Pigs, Somatotropin, Dietary Protein, Muscle Fibers, Tenderness

J. Anim. Sci. 1994. 72:615-621

Introduction
Administration of exogenous porcine somatotropin ( pST,regardless of whether it is pituitary or recombinantly derived) resulted in altered nutrient partitioning that stimulated protein accretion and reduced fat deposition in pigs (Etherton et al., 1987; Campbell et al., 1988; Evock et al., 1988; McNamara et al., 1991). Campbell et al. (1988) found that carcass composition and the rate of tissue accretion were dependent on dietary energy intake in pST-treated pigs.

'Special appreciation is extended to D. Meisinger for his assistance in obtaining recombinant porcine somatotropin from Pittman-Moore, Inc. 'Mention of specific equipment and trade name does not imply endorsement by USDA. Received May 27, 1993. Accepted November 3, 1993.

Caperna et al. (1990) examined interactions between dietary protein intake and the anabolic effects of rpST administration. Based on growth performance and body composition analyses, all pigs responded similarly to pST regardless of dietary protein concentration with no interactions. However, the magnitude of the response to rpST treatment was lowest among pigs fed 11%protein. Easter (1987) suggested that higher levels of dietary protein should be required t o meet the demand for enhanced lean tissue growth in finishing pigs treated with pST. Campbell et al. (1990) found dietary protein content to have a marked effect on the magnitude of the changes in growth performance and carcass composition elicited by exogenous pST administration to boars. The stimulatory effect of pST was probably present, but the constraints imposed by inadequate amino acid availability prevented the stimulatory response from being manifested. On the contrary, Boyd et al. (1991) showed that young barrows and gilts do not require additional protein intake to accommodate maximum response t o pST if nutrient requirements of the untreated pigs are met by the diet provided.
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Little is known about the influence of dietary protein intake in combination with pST administration on muscle fiber morphology. Therefore, the present experiment was conducted to investigate the interrelationships between exogenous pST administration and dietary protein content on porcine muscle fiber type percentage, cross-sectiodarea and shearforce values.

h, at which time standard carcass measurements, which included loin eye area (10th rib location), were recorded on the right side as described by Caperna et al. (1990).

Histochemistry
A l-cm3 fragment of tissue removed from each frozen sample was mounted on a cryostat chuck with a few drops of water so that muscle fiber orientation was perpendicular t o the cutting blade of the microtome. Mounted samples were allowed to equilibrate to -20C. Sections (12 pm thick) were cut with a microtome (Damon Minotome@ Microtome Cryostat, Needham Heights, MA). Sections were treated with the combination myofibrillar (acid) ATPase and succinic dehydrogenase staining procedure described by Solomon and Dunn (1988). The stained slides were observed with a Zeiss Standard #16 photomicro-scope (Carl Zeiss, New York). Fibers were classified according to Ashmore and Doerr (1971) on the basis of stain reaction (PR, crR, aW). All fibers inside a template (4.7 cm2 in area) were counted (approximately 75 to 100 fibers) and measured for cross-sectional area using a Zeiss Interactive Digital Analysis System (Carl Zeiss) as described by Solomon and Montgomery (1988).

Materials and Methods

A nimalslDie tslTrea tme n ts
Sixty crossbred (Yorkshire x Duroc x Landrace) barrows were used in a 5 x 2 factorial treatment array. Five levels of dietary protein (11, 15, 19, 23, and 27%) and two doses of recombinant pST ( rpST, 0 and 100 pgkg-l.d-l) were used (Pittman-Moore, Mundelein, IL). Pigs were housed in individual pens maintained at 22C and allotted randomly to 1 of 10 groups (six pigs per group) at approximately 30 kg live weight. Pigs were hand-fed once daily approximately 80% of ad libitum (based on predicted intake values for control pigs from previous work by Campbell et al. [19881) using weekly live weight measurements (1.28 kg/d at 30 kg BW and 2.11 kg/d at 55 kg BW) as described by Caperna et al. (1990). Because protein and lipid deposition are linearly related to energy intake in control and pST-treated pigs (Campbell et al., 19881, it was our intention to feed these pigs on a restriction scale that would provide equivalent daily energy intake for all pigs at similar metabolic body weights. Therefore, direct comparison of control and rpST-treated pigs could be achieved without the possible confounding effects that may arise when pigs are fed different daily levels of dietary energy. Diets were isocaloric (3.8 Mcal of DE/kg) and of equal lysine content (4.9 g/Mcal of DE). Details on the diets, performance, body composition, and hormonal status for the pigs were described by Caperna et al. (1990). All injections containing rpST or diluent (controls) were performed between 0800 and 1000, and pigs were fed after injections. Pigs were treated for 42 d. The experiment was terminated over a 10-d period following the 42 d and only one pig from each group was killed on any given day. Pigs were electrically stunned, exsanguinated, dehaired, and eviscerated. Samples (approximately 1 cm x 1 cm x 3 cm) of the medial portion of the longissimus (LM) at the anterior end of the 13th rib location and the medial location of the semimembranosus ( SM) , semitendinosus (ST) dark portion, and triceps brachii (TB) muscles were excised within 1.5 h postmortem and immediately restrained on flat sticks. Lengths of muscles were measured before excision to maintain proper fiber length when restrained. Muscle samples subsequently were frozen in liquid N2. Frozen samples were stored at -70C until histochemical analyses were performed. Carcasses were chilled (2C ) for 24

Shear Force
Two rib chops (2.5 cm in thickness) from the posterior end of the 13th rib location were removed within 1.5 h postmortem (early) and an additional two rib chops (2.5 cm thick) from the 10th rib location were removed 24 h postmortem (delayed) and vacuum-packaged. The early chops (boneless with subcutaneous fat removed) were placed in a -20C freezer immediately after vacuum packing and stored for subsequent shear-force determinations. The delayed chops (boneless) were refrigerated ( 2 "C) through 5 d postmortem and then frozen and stored at -20C for subsequent shear-force determinations. Chops were thawed and cooked (broiled) to an internal temperature of 75"C, turning once at 40C, using Farberware Open-Hearth broilers (Model 350A, Farberware Co., Bronx, NY ). Internal temperature was monitored using iron-constantan thermocouples attached to a recording potentiometer. Chops were allowed to cool to room temperature (25C) before coring. A minimum of four cores (1.27 cm in diameter) was removed from these chops, parallel to the muscle-fiber length orientation; these cores were used for shear-force determinations using a WarnerBratzler shear device mounted on a Universal Instron Testing Machine (Model 1122, Instron, Canton, MA).

Datu Analysis
Data were analyzed using GLM procedures (SAS, 1985). Final live weight was included as the covariate to determine the significance of variation among

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Table 1. Effects of dietary protein level and administration of recombinant porcine somatotropin (rpST) on the distribution of longissimus muscle fiber types
Dietary protein, 8
11 11

rpST, *gLkg-'.d-l

Final wt, kg 49.7 54.2 52.1 61.6

Fiber, %

OR
13.5 12.7 14.9 11.5 12.8 13.0 10.9 11.3 10.4 15.4
1.8

olR
33.1 32.9 26.8 26.0 28.0 28.7 29.5 38.1 29.2 30.3 2.6

OW r
53.5 54.3 58.2 62.4 59.1 58.2 59.6 50.5 60.3 54.2 3.3

0 100
0 100 0 100 0 100 0 100

15 15 19 19 23 23 27 27

SEMa

53.3 62.5 52.7 64.3 52.4 62.7 1.3

.01

Significance, P <: 0 rpST vs 100 rpST Dietary protein ( P 1 roST x P

.01 .OS

NSb NS NS

NS .05 NS

NS NS NS

aPooled standard error of the mean. ~ N S not statistically significant. =

treatments for a completely randomized 5 x 2 factorial arrangement. Least squares means were generated using SAS (1985).

Results and Discussion

Final weights for the pigs are presented in Table 1. Final weight was increased in rpST-treated pigs at all protein levels; however, the magnitude of this increase was greater with 15% or more CP in the diet ( P < .09). No significant interactions occurred between effects of rpST and dietary protein for the distribution of muscle fiber types for the LM, SM, ST, or TB (Tables 1,2, 3, and 4).In general, dietary protein had virtually no effect on the percentages of fiber types present in the four muscles evaluated. Daily administration of rpST had no influence on muscle fiber type distribution in the LM, SM, or TB muscles. We have previously reported that pST did not influence fiber type distribution in the LM of pigs (Solomon et al., 1988, 1990, 1991). However, administration of rpST resulted in significantly more aR fibers and fewer aW fibers in the ST muscle (Table 3 ) . Ashmore et al. ( 1972) concluded that aR fibers have the capacity to transform into aW fibers during postnatal growth. Transformation involves changes in energy-producing enzymes and is accompanied by a rapid increase in fiber size. Solomon (1989) suggested that the transformation of aR to aW fibers may be related to the physiological maturation rate of the muscle. Compositional differences between muscle fiber types exist between supportive and locomotive muscles (Beecher

et al., 1965; Solomon and West, 1985). Thus, responses to rpST observed for the muscles in the present study may be a result of differences between the function of these muscles. Beermann et al. (1990) examined the relationship between exogenous pST administration and ST muscle growth (both deep medial and superficial portion); pST did not alter fiber type percentages from pigs slaughtered at 100 kg live weight. However, pigs in the present study treated with rpST were approximately 40 kg lighter (and less mature) at slaughter than the pigs used in the study by Beermann et al. (1990). The results from the present study are in general agreement with those of Lefaucheur et al. (19921, who reported that 100 pgkg of pST dose administered daily to growing pigs had no effect on LM fiber type. In contrast, Whipple et al. (1992) reported that the percentage of aR fibers tended to decrease and percentage of aW fibers increased in the LM with the use of pST. They found no effect as a result of pST administration on SM fiber type distribution. Miller et al. (1975) found that total fiber number in the LM accounted for more of muscle mass variation than fiber diameter, and that faster-growing pigs possessed more but smaller muscle fibers than slower-growing pigs. Krausgrill et al. (1990) observed a significant interaction for fiber type distribution between pST and genotype; faster-growing pigs responded more to pST than did slower-growing pigs. Pigs treated with and pST had higher percentages of CYW lower percentages of CYR than did controls. Significant interactions between the effects of rpST and dietary protein level were observed for the area of

Table 2. Effects of dietary protein level and administration of recombinant porcine somatotropin (rpST) on the distribution of semimembranosus muscle fiber types
Dietary protein, % rpST, p g k-. d-' g' Fiber, 8
PR

OR
33.4 29.2 29.4 30.3 26.9 30.2

OW r 52.9 58.6 59.0 57.8 54.1 47.9 55.9 50.3 55.9 52.8
3.9

11 11 15 15
19 19

0 100 0 100
0 100

13.7 12.1 11.5 11.9 18.9 21.9 12.9 12.3 11.8 15.5 3.6

23 23 27 27

SEMa

0 100 0 100 -

31.2 37.4 32.2 31.7 2.9

Significance, P <: 0 rpST vs 100 rpST Dietary protein (PI rpST x P

NSb .05 NS

NS NS NS

NS
NS

NS

aPooled standard error of the mean. ~ N S not statistically significant. =

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Table 3. Effects of dietary protein level and administration of recombinant porcine somatotropin (rpST) on the distribution of semitendinosus muscle fiber types
Dietary protein, %
1 1 1 1

Table 4. Effects of dietary protein level and administration of recombinant porcine somatotropin (rpST) on the distribution of triceps brachii muscle fiber types
rpST, &g.kg-l.d-l
0 100

rpST, pg.kg-'.d-'
0 100 0 100 0

Fiber, 9'0

OR
38.7 32.6 33.2 34.9 36.9 38.8 37.2 38.5 31.5 34.3 2.6

ffR
32.9 39.3 32.9 38.3 32.8 32.2 34.2 34.9 29.7 35.8 2.5 .01

CW Y
28.3 28.1 33.8 26.8 30.3 28.9 28.7 26.6 38.8 29.9 2.6 .01 .05

Dietary protein, %
1 1 1 1

Fiber, %

OR
28.2 26.6 26.1 30.5 19.0 21.6 21.9 20.8 25.1 20.7 4.3

CR Y
31.0 37.2 36.3 35.3 36.4 35.4 32.1 38.4 38.3 37.2 3.1

CW Y
40.8 36.1 37.6 34.2 44.5 43.0 45.9 40.7 36.5 42.1 3.9

15 15 19 19 23 23 27 27

15 15 19 19 23 23 27 27

0 100
0 100 0 100 0 100

100
0 100

0 100

SEMa Significance, P <: 0 rpST vs 100 rpST Dietary protein ( P ) rpST x P

SEMa

NSb

NS
NS

NS NS

NS

Significance, P <: 0 rpST vs 100 rpST Dietary protein ( P ) rpST x P

NSb NS NS

NS NS NS

NS NS NS

aPooled standard error of the mean. 'NS = not significantly significant.

aPooled standard error of the mean. ~ N S not statistically significant. =

muscle fiber types for all four muscles (Tables 5 , 6, 7, and 8). In the LM muscle of pigs fed the three diets highest in protein, exogenous rpST administration resulted in larger muscle fibers. The administration of rpST had no effect when dietary protein level was 11% CP. In the majority of pST studies using the LM muscle (Solomon et al., 1988, 1990, 1991; Lefaucheur et al., 1992), pST administration, regardless of any other treatment involved, resulted in an increase in muscle fiber area. Recent research by Whipple et al. (1992) reported the lack of hypertrophic effect on LM muscle fiber types. They measured and reported fiber size as the least fiber diameter rather than total area. Whether this method of fiber size determination is directly comparable to the method reported here remains to be determined. Loin eye area, an indicator of carcass muscling, increased in size as a result of rpST treatment. This was evident a t all of the dietary protein levels except the 11%level. Loin eye areas of rpST-treated pigs protein, whereas no difference were smallest with 11% in size was observed among the four higher protein levels. This is in agreement with the findings for LM fiber area. Caperna et al. (1990) reported that carcass protein increased linearly as dietary protein was increased in both control and rpST-treated pigs. In the study by Campbell et al. (19901, in which the interrelationship between exogenous pST administration and dietary protein level was studied, an increase in loin eye area (measured at last rib) was noted in boars fed 14.5, 20.7, and 23.8% protein and administered pST. No effect of pST administration was observed at 8.3, 11.4, and 17.6% protein in the diet.

In the SM muscle, pigs given the three diets higher in protein and also receiving exogenous rpST had larger aR and CYW fibers. Negligible effects of rpST on OR fiber area were observed at all five protein levels. No rpST effect was found at 11% protein for SM fiber area (all three fiber types). Whipple et al. (1992) reported that pST had no effect on SM fiber diameter

Table 5. Effects of dietary protein level and administration of recombinant porcine somatotropin (rpST) on longissimus muscle fiber area
Dietary protein, 8
1 1 1 1

rpST, pg.kg-l.d-l
0 100 0 100 0 100 0 100 0 100

Fiber area, pm2

PR
2,156 2,005 2,429 2,592 2,505 2,858 1,817 2,643 2,445 2,708 191

CR X
2,219 2,052 2,754 2,916 2,460 2,899 1,967 2,725 2,349 2,913 22 1 .01 .01 .05

ffw

LEA, cm2a
19.4 21.5 22.5 27.0 21.9 28.0 22.5 27.6 20.4 27.8 2.1

3,695 3,479 3,822 4,249 3,353 4,240 3,480 4,647 3,574 4,359 244 .01

15 15 19 19 23 23 27 27

SEM~

Significance, P <: 0 rpST vs 100 rpST Dietary protein ( P ) rpST x P

NSC
.05 .05

NS
.05

.01 .01

NS

aLoin eye area, 10th rib location. bPooled standard error of the mean. CNS = not statistically significant.

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Table 6. Effects of dietary protein level and of recombinant porcine somatotropin (rpST) administration on semimembranosus muscle fiber area
Dietary protein, %
1 1 1 1

Table 7. Effects of dietary protein level and of recombinant porcine somatotropin (rpST) administration on semitendinosus muscle fiber area
Dietary protein, %
1 1 1 1

rpST, pgkg-'.d-'
0 100
0 100 0 100
0 100

Fiber area, pm2

rPST,
p g kg-l. d-l

Fiber area, pm2

PR
2,790 3,029 2,373 3,032 2,442 3,256 2,715 2,715 2,716 3,208 262

BR
1,671 1,775 2,477 2,327 2,224 2,436 2,087 2,225 2,365 2,291 217

CR Y
1,811 1,990 2,572 2,865 2,171 3,121 2,376 2,746 2,385 3,065 243

CW Y
3,425 3,254 4,562 4,193 3,517 4,599 3,674 4,341 3,740 4,281 368 .01 .05 .05

CtR
2,452 2,907 2,723 2,623 2,636 3,253 2,615 2,825 2,675 2,685 305 .01

CW Y
3,097 3,246 2,726 2,887 2,948 3,572 2,596 2,942 2,657 3,275 410

0 100 0 100 0 100 0 100 0 100

15 15 19 19 23 23 27 27

15 15 19 19 23 23 27 27

0 100

SEMa
Significance, P <: 0 rpST vs 100 rpST Dietary protein ( P ) rpST x P

SEMa
Significance, P <: 0 rpST vs 100 rpST Dietary protein ( P )

NSb .01

NS

.01 .01 .05

.01

.01

NSb
.05
~~

NS
.05

NS
.05

mST x P
~~

aPooled standard error of the mean. ~ N S not statistically significant. =

aPooled standard error of the mean. ~ N S not statistically significant. =

or percentage of fiber area (all three fiber types). The lack of muscle fiber hypertrophy in response to pST administration reported by Whipple et al. ( 1992) is rather unusual. The majority of pST studies involving muscle fiber evaluation have found pST administration to enlarge muscle fiber size. As much as a 26% increase in ST fiber size (all three fiber types) was observed when pigs were injected with rpST and fed 19% CP. The other protein levels evaluated did not provide for hypertrophic responses as consistent as those provided for by the 19% CP diet. Beermann et al. (1990) using the ST muscle found 120 pgkg of pST dose to increase Type I and I1 fiber size by 16% for the deep medial portion of the ST, which corresponded to the same portion of the ST we used. In the TB muscle, the administration of rpST at the 11% protein level resulted in muscle fibers larger than those in controls. At the three higher protein levels, rpST tended to increase the size of only aR and aW fibers. OR fibers were less responsive t o rpST when protein seemed to be adequate in the diet. The aR and CYWfibers accounted for the largest part of muscle hypertrophy. Periodically, giant muscle fibers were present (one or two per template field) in the LM, ST, and TB (not in tabular form) from pigs receiving rpST but were not present in the SM muscle or in excipient pigs. These abnormally large muscle fibers were not used in quantifying muscle fiber population or area. Shear force of the LM removed at 24 h (delayed) was increased 13%(pooled all dietary protein levels) with administration of rpST (Table 9). As much as a

22% increase in shear force was observed with 19%' dietary protein and rpST. Consistent with these results are the findings from our previous studies (Solomon et al., 1988, 1990, 1991), in which pST administration to barrows increased shear force of the LM. These results are also in agreement with those of Evock et al. (1988), Prusa et al. (19891, and

Table 8. Effects of dietary protein level and of recombinant porcine somatotropin (rpST) administration on triceps brachii muscle fiber area
Dietary protein, %
1 1 1 1

rpST, pgkg-'.d-'
0 100
0 100 0 100
0 100

Fiber area, pm2

PR
1,828 2,393 2,279 2,371 2,174 2,191 2,176 2,189 2,157 2,255 214

CR Y
1,983 2,810 2,618 2,686 1,936 2,654 2,429 2,731 2,380 2,608 205

lW Y
3,152 3,728 3,982 3,210 2,836 3,665 3,715 3,741 3,125 3,829 303 .05

15 15 19 19 23 23 27 27

0 100

SEMa Significance, P <: 0 rpST vs 100 rpST Dietary protein ( P ) rpST x P

NSb NS

.01 NS
.05

NS

NS .05

aPooled standard error of the mean. ~ N S not statistically significant. =

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Table 9. Effects of dietary protein level, administration of recombinant porcine somatotropin (rpST),and time of sampling on longissimus shear-force values
Dietary protein, %
11 11 15 15 19 19 23 23 27 27

rpST, pg.kg-'.d-'
0 100 0 100 0 100 0 100

Shear force, kg11.27 cm Delayeda

4.96 5.58 5.11 5.66 5.62 6.88 5.49 6.01 5.11 5.67 .36 .01 .05 NSd

Earlyb
4.09 3.94 4.07 4.27 4.69 5.85 4.79 4.62 4.46 4.14 .24

tein concentrations in the diet constrained the muscle fiber hypertrophy response to rpST. High protein levels (above 19%) were not required for significant increases in carcass protein accretion in pigs treated with rpST. Exogenous rpST increased loin eye area through hypertrophy of individual muscle fibers without altering the distribution (population) of fiber types. The negative effects of rpST treatment on shear force were overcome when meat was frozen within 1.5 h postmortem (pre-rigor) rather than post-rigor.

Literature Cited
Ashmore, C. R., and L. Doerr. 1971. Comparative aspects of muscle fiber types in different species. Exp. Neurol. 31:408. Ashmore, C. R., G. Thompkins, and L. Doerr. 1972. Postnatal development of muscle fiber types in domestic animals. J. Anim. Sci. 34:37. Beecher, G. R., R. G. Cassens, W. G. Hoekstra, and E. J. Briskey. 1965. Red and white fiber content and associated post-mortem properties of seven porcine muscles. J. Food Sci. 30:969. Beermann, D. H., V. K. Fishell, K. Roneker, R. D. Boyd, G. Armbruster, and L. Souza. 1990. Dose-response relationships between porcine somatotropin, muscle composition, muscle fiber characteristics and pork quality. J. Anim. Sci. 68:2690. Boyd, R. D., D. E. Bauman, D. G. Fox, and C. G. Scanes. 1991. Impact of metabolism modifiers on protein accretion and protein and energy requirements of livestock. J. Anim. Sci. 69(Suppl. 2):56. Campbell, R. G., R. J. Johnson, R. H. King, M. R. Taverner, and D. J. Meisinger. 1990. Interaction of dietary protein content and exogenous porcine growth hormone administration on protein and lipid accretion rates in growing pigs. J. Anim. Sci. 68:3217. Campbell, R. G., N. C. Steele, T. J. Caperna, J. P. McMurtry, M. B. Solomon, and A. D. Mitchell. 1988. Interrelationships between energy intake and endogenous porcine growth hormone administration on the performance, body composition and protein and energy metabolism of growing pigs weighing 25 t o 55 kilograms live weight. J. Anim. Sci. 66:1643. Caperna, T. J., N. C. Steele, D. R. Komarek, J. P. McMurtry, R. W. Rosebrough, M. B. Solomon, and A. D. Mitchell. 1990. Influence of dietary protein and recombinant porcine somatotropin administration in young pigs: growth, body composition and hormone status. J. Anim. Sci. 68:4243. Easter, R. A. 1987. Nutritional requirements and repartitioning agents. In: Proc. Univ. of Illinois Pork Industry Conf. Dec. 10, 1987. p 193. Etherton, T. D., J. P. Wiggins, C. M. Evock, C. S. Chung, J. F. Rebhun, P. E. Walton, and N. C. Steele. 1987. Stimulation of pig growth performance by porcine growth hormone: Determination of the dose-response relationship. J. Anim. Sci. 64:433. Evock, C. M., T. D. Etherton, C. S. Chung, and R. E. Ivy. 1988. Pituitary porcine growth hormone ( p G H ) and a recombinant pGH analog stimulate pig growth performance in a similar manner. J. Anim. Sci. 66:1928. Goodband, R. D., J. L. Nelssen, R. H. Hines, D. H. Kropf, R. C. Thaler, B. R. Schricker, G. E. Fitzner, and A. J. Lewis. 1990. The effects of porcine somatotropin and dietary lysine on growth performance and carcass characteristics of finishing swine. J. Anim. Sci. 68:3261. Krausgrill, D. I., R. G. Campbell, and N. M. Tulloh. 1990. Effects of exogenous porcine growth hormone (pGH), sex and strain on semitendinosus muscle fiber in pigs. Proc. Aust. Soc. Anim. Prod. 18:509. Lefaucheur, L., A. Missohou, P. Ecolan, G. Monin, and M. Bonneau. 1992. Performance, plasma hormones, histochemical and biochemical muscle traits, and meat quality of pigs administered

0 100
__

SEMC Significance, P <: 0 rpST vs 100 rpST Dietary protein ( P ) rpST x P

NS .05 NS

aSamples removed at 24 h postmortem and frozen after 5 d postmortem of cooler storage ( 2 "C ) , bSamples excised and frozen ( -20C) within 1.5 h postmortem. CPooled standard error of the mean. ~ N S not statistically significant. =

Goodband et al. (1990), who indicated that meat from pST-treated pigs was slightly less tender than that from controls. Novakofski ( 198 7 ) found that shear forces increased slightly in pST-treated compared with untreated pigs. Other researchers have found no negative effects on tenderness due to pST (Beermann et al., 1990; Nieuwhof et al., 1991). The time postmortem of sampling the LM for subsequent shear-force analysis had a significant effect on shear-force tenderness. Differences in shear force between rpST and control pigs were virtually eliminated when chops were removed and frozen within 1.5 h postmortem (early). However, the effect of rpST was not eliminated at the 19% dietary protein when chops were frozen at 1.5 h postmortem. Some of the inconsistencies reported in the literature for shear force and tenderness as a result of pST may be a result of inconsistencies in the time at which the sample is removed and the time at which the sample is frozen. Compositional differences between supportive and locomotive muscle types have been reported and may explain differences in the response to rpST and dietary protein of the four muscles evaluated in the present study.

Implications
Dietary protein had a marked effect on the magnitude of changes in muscle fiber hypertrophy elicited by recombinant porcine somatotropin (rpST). Low pro-

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