Seventy-two barrows (initial weight = 57.1 kg) were used to determine the interrelationship between porcine somatotropin (pST) and dietary lysine and their effects on growth performance and carcass characteristics. Pigs were injected daily in the extensor muscle of the neck with either 4 or 8 mg of pST and fed a pelleted corn-soybean meal-sesame meal diet (.8% Lysine; 17.8% CP) or diets containing 1.0, 1.2, or 1.4% lysine provided by additions of L-lysine HCl (2 x 4 factorial arrangement). Control pigs (placebo injection) received the .8% lysine diet. All diets were formulated to contain |is greater than or equal to~ 200% of current recommendations for other amino acids, vitamins, and minerals. A tendency for a pST x lysine interaction was observed for cumulative ADG (P |is less than~ .15) and feed conversion (G/F; P |is less than~ .05). Average daily gain and G/F were improved by increasing dietary lysine level in pigs injected with 4 mg/d of pST; however, pigs injected with 8 mg/d of pST had greater improvements in cumulative ADG and G/F with added lysine. Feed intake was reduced (quadratic, P |is less than~ .10) as dietary lysine level and pST dosage increased. Increasing pST dosage and dietary lysine increased (linear, P |is less than~ .05) longissimus muscle area and decreased backfat thickness. Trimmed ham and loin weights were increased (linear, P |is less than~. 10) by pST dosage. Chemical composition of samples taken from the loin, ham, and belly indicated increased moisture and CP and decreased lipid content as pST dosage and dietary lysine level increased (quadratic, P |is less than~ .05). Shear force values from loin and semimembranosus increased with increasing lysine level (quadratic, P |is less than~ .01) and pST dosage (linear, P |is less than~ .05); however, these differences were not detected by sensory analysis (P |is greater than~ .20). Plasma urea concentrations on d 28 decreased with increasing lysine level (quadratic, P |is less than~ .05), and plasma lysine concentrations increased (linear, P |is less than~ .01). Based on the pST x lysine interaction for ADG and G/F, these data suggest that the lysine level needed to maximize growth performance and carcass characteristics may be proportional to the pST dosage provided. Growth and carcass characteristics were maximized by dietary lysine intakes of 27 to 32 and |is greater than or equal to~ 36 g/d for pigs injected with 4 and 8 of mg/d of pST, respectively. Introduction The feasibility of using porcine somatotropin (pST) as a growth promotant in swine has raised many questions about the effect it will have on the swine industry. One area that is beginning to be defined is the effect of diet and nutrient density on lean tissue deposition and adipose tissue growth in finishing pigs injected with pST. Recent results indicate that the lysine requirement of finishing pigs injected with 4 mg/d of pST is approximately 25 to 30 g/d (Goodband et al., 1990). However, the lysine requirement of finishing pigs may be proportional to their potential for lean tissue deposition, which can be affected by the quantity of pST administered. Consequently, the amount of pST administered may influence the lysine requirement. It would seem that some previous experiments that have evaluated the effects of pST on pig performance may have been confounded by inadequate intakes of amino acids (Kraft et al., 1986; Wolfrom et al., 1986; Smith et al., 1987). Thus, the asymptotic response in growth performance of finishing pigs to increasing quantities of exogenous pST may be a result of inadequate dietary amino acid levels. Therefore, the objectives of this study were to evaluate the performance of finishing pigs treated with different dosages of pST in combination with various levels of dietary lysine and to determine whether responses to increasing pST dosage were affected by dietary lysine level. Experimental Procedures Animals. Seventy-two barrows, averaging 57 kg, were allotted on the basis of initial weight an ancestry in a randomized complete block design to one of nine experimental treatments. Treatments were arranged in a 2 x 4 factorial with a control treatment. Main effects were pST dosage and dietary lysine level. There were two pigs per pen providing four observations per treatment (pen was considered the experimental unit). Pigs were housed in 4.6-m x 1.2-m pens in a modified open-front building with solid concrete floors and allowed ad libitum access to feed and water. Experimental treatments consisted of daily injections of either 4 or 8 mg of recombinant pST in combination with a corn-soybean meal-sesame meal diet (.8% lysine, 17.8% CP; Table 1) or diets containing 1.0, 1.2, or 1.4% lysine provided by additions of L-lysine HCl. The chemical composition of the .8% lysine diet (control) was determined by AOAC (1984) procedures (Table 2). Amino acid analysis was determined by ion-exchange chromatography after acid hydrolysis. Cystine and methionine were determined after oxidation with performic acid (Moore, 1963). The control diet was calculated to meet the estimated requirement for lysine but to contain twice the requirements (NRC, 1988) of all other amino acids. Amino acid analysis indicated that all amino acids (other than lysine) exceeded the pig's estimated requirement by 200%. Table 1. Composition of control diet(a) Ingredient Perce ntage Corn 60.98 Soybean meal (44% CP) 18.30 Sesame meal 10.00 Soybean oil 5.00 Monocalcium phosphate 2.77 Limestone .78 Salt .50 Trace mineral premix(b) .20 Vitamin premix(c) .5 0 Selenium premix(d) .05 Threonine .10 Methionine .05 Sucrose .77 Total 100.00 a Dietary lysine l evels 1.0, 1.2, and 1.4% were formulated by adding .26, .52, and .77 L-lysine HC l at the expense of sucrose. b Provided the following per kilogram of complete d iet: Ca, 220 mg; Fe, 440 mg; Zn, 440 mg; Mn, 440 rag; Cu, 44 rag; I, 13.2 mg; an d Co, 4.4 mg. c Provided the following per kilogram of complete diet: vitamin A, 11,023 IU; vitamin D, 1,102 IU; vitamin E, 44 IU; menadione, 4.4 mg; riboflavin , 11 mg; D-panthothenic acid, 27.6 mg; niacin 60.6 mg; choline, 1,102 mg; and vi tamin |B.sub.12~, .55 mg. d Provided .3 mg of Se per kilogram of complete diet. Table 2. Chemical analyses of control diet(a) Component Percentage DM 89.07 CP 17.80 Ether extract 9.93 Crude fiber 5.60 Ash 5.60 Ca 1.10 P 1.00 Essential and se miessential amino acids Arginine 1.32 Cyst ine .31 Histidine .45 Isoleucine .90 Leucine 1.64 Lysine .80 Methionine .45 Phenylalanin e .95 Tyrosine .74 Threonine .90 Tryptophan .22 Valine .90 a Values are expressed on an as-fed basis. Porcine somatotropin was dissolved in buffer 14 d before initiation of the study and stored frozen to ensure stability. Before use, the daily allotment of pST was allowed to thaw overnight at 4|degrees~C. Pigs were injected daily between 0600 and 0700 in the extensor muscle of the neck. The contralateral muscle was injected on alternate days. Pigs and feeders were weighed on d 14 and 28 and then weekly until the mean weight of the two pigs in a pen reached 105 |+ or -~ 2 kg. Daily injections were terminated at this time, and a 7-d withdrawal period was initiated. Blood Sampling. Serum and plasma samples were obtained on d 14 and 28 and on the last day of pST administration. Analyses for glucose, urea, free fatty acids (FFA), creatinine, insulin, somatotropin, and triiodothyronine (|T.sub.3~) were conducted as described previously (Goodband et al., 1990). Interassay CV were .6, 1.3, and 6.6% for somatotropin, insulin, and |T.sub.3~, respectively. Carcass Data Collection. On the last day of pST administration, all pigs were ultrasonically scanned (Technicare 210 DX, Johnson & Johnson, Englewood, CO) for backfat thickness adjacent to the first and last ribs and last lumbar vertebra. The three measurements were then averaged and adjusted to a constant weight (104 kg) using NPPC (1988) procedures. In addition, backfat depth was also measured at the 10th rib approximately 6 cm from the midline. Backfat was measured at this time because pigs were skinned at slaughter. Pigs were slaughtered in six groups of 12 pigs each over a 26-d period. Carcass measurements, muscle dissection, sampling procedures, shear force determination, and sensory panel evaluations were conducted as previously described (Goodband et al., 1990). In the sensory analysis, an incomplete block design (Snedecor and Cochran, 1980) was used to assign daily sampling order to treatments to allow panelists to evaluate six chops per day. Statistical Analysis. Data were analyzed using the GLM procedure of SAS (1985). Analysis was conducted according to procedures described by Milliken and Johnson (1984) as a 2 x 4 factorial with a control treatment. Linear, quadratic, and cubic comparisons of treatment means for lysine level were evaluated among pST-treated pigs, whereas linear and quadratic caparisons were made among control pigs and pigs treated with 4 or 8 mg of pST. Sensory analysis and carcass measurement adjustments were described previously (Goodband et al., 1990). Results Growth Performance. A pST x lysine interaction was observed for d-28 ADG (P |is less than~ .05); similar numeric trends were observed (P |is less than~ .15) on d 14 and overall (Table 3). This seemed to be the result of the relatively small increase in ADG for pigs injected with 4 mg/d of pST compared with the larger increase to added lysine observed in pigs injected with 8 mg/d. Overall ADFI was decreased by increasing pST dosage (quadratic, P |is less than~ .10) and increasing lysine level (quadratic, P |is less than~ .05). A pST x lysine interaction was observed for d-28 and overall G/F (P |is less than~ .05); injection of 8 mg/d of pST resulted in a greater improvement in G/F than injection of 4 mg/d. The administration of pST (both 4 and 8 mg/d) improved G/F (quadratic, P |is less than~ .05) throughout the study compared with the placebo injections. Blood Metabolites. No pST x lysine interactions (P |is greater than~ .20) were observed for plasma urea, glucose, FFA, or creatinine concentrations (Table 4). Plasma urea concentrations decreased as pST dosage increased (quadratic, P |is less than~ .05). Plasma urea concentrations (d 28 and final bleeding) also were decreased by increasing dietary lysine level (quadratic, P |is less than~ .05). In pigs injected with 4 mg/d of pST, urea concentration on d 28 decreased and then increased as dietary lysine increased. However, in pigs injected with 8 mg/d of pST, urea concentrations reached a plateau at 1.0% dietary lysine and remained unchanged at either 1.2 or 1.4% lysine. Glucose concentrations were elevated in pST-treated pigs at all sampling times (quadratic, P |is less than~ .05) but were unaffected by dietary lysine level. TABULAR DATA OMITTED Plasma FFA concentrations increased (linear, P |is less than~ .05) as dietary lysine level increased and tended to be increased by pST dosage (d 28, linear, P |is less than~ .05). Creatinine concentrations were reduced (quadratic, P |is less than~ .05) in the serum of pST-treated pigs compared with that of control pigs. TABULAR DATA OMITTED Plasma lysine concentrations on d 28 increased (linear, P |is less than~ .05) as dietary lysine increased (Table 5). Histidine was the only other amino acid to be affected by dietary lysine level (linear, P |is less than~ .05). Leucine, threonine, and valine concentrations decreased as pST dosage increased (linear, P |is less than~ .05), whereas methionine concentration decreased then increased (quadratic, P |is less than~ .10). Plasma somatotropin concentrations increased with increasing pST dosage (linear, P |is less than~ .05; Table 6). Increasing dietary lysine level tended (linear, P |is less than~ .10) to lower somatotropin concentrations in serum samples collected at termination of daily injections. Insulin concentrations were elevated by increasing TABULAR DATA OMITTED pST dosage (linear, P |is less than~ .05; quadratic, P |is less than~ .10) and also increased (linear, P |is less than~ .10) as dietary lysine level increased. Triiodothyronine concentrations increased numerically (P |is greater than~ .15) as lysine level increased. Increasing pST dosage increased (linear, P |is less than~ .05) |T.sub.3~ concentrations in serum. Carcass Data. Although pigs were slaughtered at a constant weight, hot carcass weight and chilled carcass weight were decreased (quadratic, P |is less than~ .05) by increasing pST dosage (Table 7). Dietary lysine level had no effect on carcass weight. Control pigs had the heaviest hot and chilled carcass weights. This resulted in decreased dressing percentage of pigs injected with either 4 or 8 mg/d of pST (linear, P |is less than~ .05; quadratic, P |is less than~ .10). Control pigs had greater average backfat and 10th rib fat depth than pigs injected with either 4 or 8 mg of pST (linear, P |is less than~ .05). Increasing dietary lysine reduced average and 10th rib backfat (linear, P |is less than~ .05). Longissimus muscle area increased with increasing pST dosage (linear, P |is less than~ .05) and tended to increase with dietary lysine level (linear, P |is less than~ .15). Carcass length was unaffected by experimental treatments. Kidney fat weight decreased at pST dosage increased (quadratic, P |is less than~ .05) and also tended to decrease with increasing dietary lysine level (linear, P |is less than~ .15). Trimmed weights of the ham and loin tended to TABULAR DATA OMITTED increase with increasing pST dosage (linear, P |is less than~ .10) and dietary lysine level (linear, P | is less than~ .15). Weights of the semitendinosus, adductor, and quadriceps femoris muscles and lean trim were increased (linear, P |is less than~ .05) by pST dosage (Table 8). Color scores from the ham increased as pST dosage increased (linear, P |is less than~ .05); however, a pST x lysine interaction was observed for longissimus color scores (P |is less than~ .05). This seemed to be a result of a greater change in color score in pigs treated with 4 mg/d of pST than observed in pigs treated with 8 mg/d of pST. Ham firmness increased with increasing pST dosage (linear, P |is less than~ .10) and dietary lysine (linear, P |is less than~ .01), whereas longissimus firmness increased then decreased as lysine level increased (quadratic, P |is less than~ .05). These changes in longissimus color and firmness correspond with the pST x lysine interaction (P |is less than~ .05) observed for longissimus 24-h pH. As with backfat thickness, marbling scores decreased with increasing pST dosage (quadratic, P |is less than~ .10) and dietary lysine level (linear, P |is less than~ .10). TABULAR DATA OMITTED Composition of longissimus, belly, and individual ham muscles (semimembranosus, SM; semitendinosus, ST; and biceps femoris, BF) were affected by pST dosage and dietary lysine level (Table 9). Moisture content increased as pST dosage (quadratic, P |is less than~ .05) and dietary lysine level increased (linear, P |is less than~ .10) in all muscles except the SM. The increase in moisture was associated with a trend toward increased CP content, which was increased by lysine level (linear, P |is less than~ .05) and pST dosage (linear and quadratic, P |is less than~ .05), but only in the longissimus and belly. Lipid content was decreased in the longissimus, ST, BF, and belly samples by increasing pST dosage (quadratic, P |is less than~ .05). Lipid content of all muscles also was substantially reduced by increasing dietary lysine level (linear, P |is greater than~ .05). Ash content increased with pST and dietary lysine, but only in SM and belly samples. In general, the greatest response to pST and dietary lysine level was observed in belly samples, in which there was an increase of approximately 30% in CP and a reduction of 50% in lipid content for pigs treated with 8 mg/d of pST and fed 1.4% lysine compared with control pigs. Weights of the heart, liver, kidney, and spleen were increased (linear, P |is less than~ .05) by pST dosage (Table 10). Thaw losses were decreased in longissimus and SM samples by increasing pST dosage (linear, P |is less than~ .05; quadratic, P |is less than~ .10; Table 11). Drip cooking losses were decreased in longissimus samples by increasing pST dosage (quadratic, P |is less than~ .10), whereas evaporative TABULAR DATA OMITTED cooking losses were decreased in SM samples by increased lysine level (quadratic, P |is less than~ .10). This resulted in a decrease in total cooking losses in SM samples (quadratic, P |is less than~ .10). However, when combined, total cooking and thaw losses were unaffected by experimental treatment, although there were tendencies toward decreased total losses among pST-treated pigs as dietary lysine level increased. Shear force values increased with increasing pST dosage in longissimus and SM samples (linear, P |is less than~ .05) and increased as lysine level increased (quadratic, P |is less than~ .10; Table 12). However, these changes in shear force were not detected by sensory analysis, because scores for juiciness, flavor, and tenderness were unaffected by either pST dosage or lysine level (P |is greater than~ .25). TABULAR DATA OMITTED TABULAR DATA OMITTED Discussion These data confirm previous results indicating an improvement in growth performance and carcass characteristics of pST-treated pigs in response to additional lysine (Goodband et al., 1990). As in our previous experiment, performance of pigs injected with 4 mg/d of pST was maximized at a daily lysine intake of approximately 27 g. However, these data also indicate an interactive effect between pST and dietary lysine on gain and feed efficiency (pST x lysine interaction, P |is less than~ .05). When pST dosage was increased to 8 mg/d, pig performance continued to improve up to a lysine level of 1.4% (approximately 36 g/d). Based on the linear (P |is less than~ .05) nature of the response to added lysine, it would seem that pigs injected with 8 mg/d of pST may require more dietary lysine than the 1.4% provided in this experiment. Inadequate amino acid levels may have limited performance in other experiments and could account for some of the variation in response of pigs in pST administration (Kraft et al., 1986; Wolfrom et al., 1986; Smith et al., 1987). TABULAR DATA OMITTED Administration of pST decreased ADFI in a dose-dependent fashion. This response was similar to results from previous experiments (Etherton et al., 1986, 1987; Campbell et al., 1988). Feed intake was decreased by increasing dietary lysine level. This response may have been an attempt by pigs fed the .8% lysine diets to compensate for the inadequate lysine level by eating more feed (Baker, 1986). The elevations in plasma glucose concentrations are consistent with other experiments (Etherton et al., 1987; Campbell et al., 1988; Goodband et al., 1990) and are a result of altered hepatic glucose output and clearance (Gopinath and Etherton, 1989). Elevations in FFA and |T.sub.3~ concentrations paralleled changes observed in our previous experiment and support the current hypothesis that FFA uptake by adipose tissue is attenuated by the antagonism between pST and insulin in adipose tissue (Walton and Etherton, 1986). Thus, fat may be utilized to a greater extent in pST-treated pigs as an energy substrate. This would necessitate greater demand for oxygen, which would be supplied by |T.sub.3~-stimulated dissociation of oxygen from hemoglobin. Changes in plasma urea concentrations are typical of those observed in experiments to determine amino acid requirements of pigs (Lewis et al., 1980; Goodband et al., 1990). As dietary lysine increased and approached the pig's requirement, protein synthesis became more efficient, and other amino acids were incorporated into protein rather than being deaminated with the formation of urea. Plasma lysine concentrations were decreased in pST-treated pigs fed .8% lysine compared with control pigs, suggesting insufficient dietary lysine intake. However, plasma lysine concentrations increased in pST-treated pigs as dietary lysine increased above 1.0%, suggesting that the pig's lysine requirement was met or exceeded. Surprisingly, high plasma lysine concentrations were observed in pigs injected with 8 mg/d of pST and fed the 1.4% lysine diet despite increasing ADG and G/F. Plasma lysine values would be expected to increase sharply only after growth performance criteria had leveled off (Lewis et al., 1980; Asche et al., 1985; Baker, 1986). Furthermore, the range of plasma lysine concentrations, although similar to those observed in a previous study (Goodband et al., 1990), are much lower than values observed in the young pig (Lewis et al., 1980; Asche et al., 1985). The change in plasma lysine values in response to increasing dietary lysine observed in our study supports the hypothesis that pST administration, in addition to increasing the pig's amino acid requirement, may also alter the efficiency of amino acid utilization (Campbell et al., 1991). Other amino acid concentrations decreased in response to increasing pST dosage, which is similar to the response observed for plasma urea concentrations. Insulin concentrations were elevated as a result of increased glucose concentrations (Gopinath and Etherton, 1989), in agreement with previous experiments (Etherton et al., 1987; Evock et al., 1988; Goodband et al., 1990). Because all pigs were slaughtered at approximately the same weight, differences in carcass weight seemed to be caused by a reduction in skinned dressing percentage of pST-treated pigs. Dressing percentage values have been variable but in general are approximately 0 to 3% lower in pigs treated with pST than in control pigs (Evock et al., 1988; Bryan et al., 1989; Goodband et al., 1990). Dressing percentage would be expected to decrease, because pST-treated pigs were leaner and had increased organ weights. Responses in longissimus muscle area and backfat thickness were consistent with previous studies evaluating the effects of pST dosage on carcass traits (Etherton et al., 1987; Campbell et al., 1988; Evock et al., 1988) and also with studies showing that carcass leanness is increased by increasing dietary lysine (Williams et al., 1984; Asche et al., 1985; Goodband et al., 1990). However, as with ADG and G/F, backfat thickness and longissimus muscle area were improved to a greater extent when high-lysine diets were fed to pigs injected with 8 mg/d. Reductions in kidney fat paralleled those observed for backfat thickness, marbling score, and intramuscular lipid content. The changes observed in color, firmness, and 24-h pH in pST-treated pigs may indicate that a slightly higher 24-h pH resulted in slightly darker and firmer pork compared with that from control pigs. However, because color and firmness scores were still within an acceptable range of normal values, it is unlikely that treatments caused a predisposition toward DFD pork. Chemical composition of selected muscles was similar to that observed previously in pST-treated pigs fed added lysine (Goodband et al., 1990). The belly, one of the less economically important, fatter cuts of pork, seemed to show the greatest response to pST and dietary lysine. Little information is available on the effects of pST on such cuts of pork; however, our data indicate a potential to improve their quality. Cooking losses and sensory properties of pork from pST-treated pigs were unaffected by experimental treatment. These findings are consistent with previous data indicating little adverse effect on pST on pork quality (Beermann et al., 1988; Evock et al., 1988; Goodband et al., 1990). Our sensory panel evaluations seem to contradict those of DeVol et al. (1988), who found decreased tenderness and juiciness of pork containing |is less than~ 2.5% lipid. Longissimus samples from control and pST-treated pigs were on either side of this threshold value and yet no differences were distinguishable by the trained sensory panel. Implications These data further demonstrate that porcine somatotropin administration improves daily gain, feed conversion, and carcass characteristics of finishing pigs without adversely affecting pork quality. The response of finishing pigs to exogenous porcine somatotropin administration is dependent on the dietary lysine level provided. 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