The Effect of Dietary Lysine and Valine Fed During Lactation on Sow and Litter Performance 1,2

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1 The Effect of Dietary Lysine and Valine Fed During Lactation on Sow and Litter Performance 1,2 B. T. Richert*,3, M. D. Tokach*, R. D. Goodband*,4, J. L. Nelssen*, R. G. Campbell, and S. Kershaw *Department of Animal Sciences and Industry, Kansas State University, Manhattan and Bunge Meats Ltd., New South Wales, Australia ABSTRACT: Sows (98 first parity and 104 second parity) were used to determine the effects of dietary lysine and valine on lactation performance. Treatments were arranged in a 2 3 factorial with two levels of lysine (.80 or 1.20%) and three valine:lysine ratios (80, 100, or 120% of lysine). For all sows, increasing dietary lysine increased litter weaning weight ( P <.001) and litter weight gain ( P <.002) and reduced sow weight loss ( P <.001). Litter weight gain tended ( P =.22) to increase with increasing dietary valine, but the increase was not significant. Data were separated into two groups: sows that weaned 10 or more pigs and sows that weaned fewer than 10 pigs. For sows that weaned 10 or more pigs, litter weaning weight ( P <.001) and litter weight gain ( P <.001) increased and sow BW loss decreased ( P <.001) when dietary lysine increased from.80 to 1.20%. For sows that weaned fewer than 10 pigs, increasing lysine had no effect ( P <.77) on litter growth rate. For sows weaning 10 or more pigs, litter weaning weights (linear, P <.04; quadratic, P <.06) and litter weight gain increased (linear, P <.04; quadratic, P <.02) as dietary valine increased. For sows that weaned fewer than 10 pigs, maximum litter weight gain was observed at a valine:lysine ratio of 100% (quadratic, P <.13). These results demonstrate the need to increase dietary lysine and valine as litter weaning weights increase. High-producing sows that wean 10 or more pigs require increased dietary lysine and valine to maximize litter growth rate and minimize sow weight loss compared with sows weaning fewer than 10 pigs. The independent increases in litter weaning weights from adding lysine and valine suggest separate modes of action for these amino acids in high-producing sows. Key Words: Lysine, Valine, Lactation, Sows J. Anim. Sci : Introduction Johnston et al. (1993) demonstrated increased litter weaning weights with increased lysine and protein in lactation diets for high-producing sows with an estimated lysine requirement of 54 g/d. Several studies with lactating sows suggested that approximately 26 g of lysine was required for a kilogram of litter weight gain per day (see Pettigrew, 1993). These results suggest that the lysine requirement of today s high-producing sows far exceeds the current 1 Contribution no J from the Kansas Agric. Exp. Station, Manhattan. 2 Appreciation is expressed to Lonza, Inc., Fair Lawn, NJ, for partial financial support for this research project. 3 Present address: Department of Animal Science. Purdue University, West Lafayette, IN To whom correspondence should be addressed. Received February 26, Accepted March 3, NRC (1988) estimate of 31 g/d. Rousselow and Speer (1980) determined the valine requirement of lactating sows to be.68% in a diet containing.58% lysine. However, they estimated a valine requirement of.53% using N balance as the response criterion. Tokach et al. (1993) and Richert et al. (1996) demonstrated that the valine requirement for high-producing lactating sows also is higher than the ARC (1981) and NRC (1988) estimates. Research using mammary cannulation in sows (Trottier et al., 1994) and dairy cows (Hanigan et al., 1991) has indicated that valine is taken up by the mammary gland in approximately 30% greater amounts than it is secreted in milk protein. This high extraction rate indicates a greater requirement for valine than can be accounted for by estimates of milk protein composition and the current estimated allowance for sow maintenance requirements, which has been the method to estimate amino acid requirements of lactating sows (ARC, 1981). However, no research has been conducted to evaluate whether the valine requirement changes at different 1853

2 1854 RICHERT ET AL. Table 1. Diet composition (as-fed basis) a Dietary lysine, % Item.80 b 1.20 c Wheat Barley Millmix d Wheat starch Expelled soybean meal (44% CP) Tallow Salt.5.5 Limestone Dicalcium phosphate L-Lysine-HCl DL-Methionine L-Threonine L-Isoleucine L-Tryptophan L-Histidine Premix e a Diets were fed for a 23.7-d lactation period. b Valine levels of 80, 100, and 120% of lysine corresponding to.65,.80, and.95% dietary valine were created by replacing wheat starch in the basal diet in.15% increments with L-valine. All three diets were formulated to 14.4% CP, 1.0% Ca,.8% P. c Valine levels of 80, 100, and 120% of lysine corresponding to.96, 1.20, and 1.44% dietary valine were created by replacing wheat starch in the basal diet in.24% increments with L-valine. All three diets were formulated to 20.6% CP, 1.05% Ca,.8% P. d Consisted of mostly wheat middlings with an analyzed concentration of 15.4% CP and.69% lysine. e Provided (per kg of diet): 16.5 IU vitamin A, 3.3 IU vitamin D3, 88 IU vitamin E, 3.9 IU vitamin B 2, 2.2 IU vitamin B 6,.022 IU vitamin B 12,.22 mg biotin, 11 mg d-pantothenic acid,.55 mg folic acid, 16.5 mg niacin, 220 mg choline, 95 mg betaine, 1.1 mg iodine,.22 mg selenium, 22 mg copper, 88 mg iron, 60 mg manganese, 83 mg zinc, 110 mg endox (antioxidant). levels of milk production or lysine intakes. Therefore, our objectives were to evaluate the effects of increasing lysine and valine on sow and litter performance during lactation and to determine whether the response to valine changes as dietary lysine increases, indicating a similar or additive effect of these two amino acids on milk production. Materials and Methods Animals and Diets. Maternal white-line sows (Large White Landrace Yorkshire; 98 first parity and 104 second parity) were used to evaluate the interactive effects of increasing lysine and valine on sow and litter performance. Treatments were arranged in a 2 3 factorial including two levels of lysine (.80 and 1.20%) and three valine:lysine ratios (80, 100, and 120% of lysine). Two basal diets (.80 and 1.20% lysine) were formulated (Table 1). The.80% lysine diet contained.65% valine (80% valine: lysine). The 100 and 120% valine:lysine diets were formulated by replacing wheat starch in the basal diet with.15 and.30% L-valine, respectively. The 1.20% lysine diet contained.96% valine (80% valine:lysine). The 100 and 120% valine:lysine diets were formulated by replacing wheat starch with.24 and.48% L-valine, respectively. All.80% lysine diets were formulated to 14.4% CP, 1.0% Ca, and.8% P. All 1.20% lysine diets were formulated to 20.6% CP, 1.05% Ca, and.8% P. Diet samples were collected at the beginning of the experiment and stored at 20 C until amino acids were analyzed by HPLC (AOAC, 1990) (Table 2). Husbandry. The experiment was conducted on a research farm in New South Wales, Australia (Bunge Meats Ltd.). The lactation portion of the experiment was conducted from January to March Sow and litter weights were recorded on d 2 postfarrowing, after cross-fostering was completed to equalize litter size across dietary treatments, and at weaning. Sows were fed three times daily (0630, 1130, and 1600) in an attempt to provide ad libitum access to feed and minimize any possible confounding effects of synthetic amino acid utilization and feeding frequency. Feed intake was recorded for the entire lactation period. The average lactation length was 23.7 ± 2 d. Sows were evaluated using real time ultrasound (Corometrics Medical Systems, Wallinford, CT) for backfat ( BF) thickness at approximately 6 cm off the midline on both sides of the body at the 10th rib and last lumbar vertebra. Eight farrowing groups of sows were used, with five litters per treatment in each group. Thirty-eight sows failed to complete the study or their data were removed from the data set before analysis. Twenty-two sows were removed from the study for complete refusal to consume the experimental diets or structural unsoundness. Data for eight sows were removed from the data set because of chronically low feed intake, which averaged less than 1.5 kg/d, and data for eight sows were removed because sows Table 2. Chemical analysis of basal diets (as-fed basis) Basal diet, % lysine Item, %.80 a 1.20 b DM CP Fat Arginine Histidine Isoleucine Leucine Lysine Methionine Cystine Phenylalanine Threonine Tryptophan Valine a Analyzed valine concentrations at 100 and 120% of lysine were.77 and.98%, respectively. b Analyzed valine concentrations at 100 and 120% of lysine were 1.17 and 1.37%, respectively.

3 weaned six pigs or fewer. In addition, rebreeding data from nine sows were not used because of health problems. A wheat-barley-lupin-based gestation diet formulated to 13.9% CP and.60% lysine was fed to all sows during gestation at 2.3, 2.5, and 2.7 kg/d during the first, second, and last third of gestation, respectively. Following weaning, sows were fed 2.8 to 3.0 kg/d of the gestation diet in two equal feedings until breeding. Days to estrus, removal rate, and subsequent litter size were also recorded. Data for sows not bred by 21 d postweaning were removed from the subsequent litter size data. Statistical Analysis. The GLM procedure of SAS (1988) was used to determine treatment effects of lysine, valine, and their interaction (Peterson, 1985). Days of lactation and litter size after cross-fostering were used as covariates for all data. Initial sow BW and initial sow BF thickness were used as covariates for changes in these characteristics. Because of the greater increase in litter weaning weight in response to increasing valine by high-producing sows (i.e., those weaning greater than 10 pigs per litter; Tokach et al., 1993), the data set was split into two sow groups: sows that weaned 10 or more pigs and sows that weaned fewer than 10 pigs. The data also were divided into first and second parity sows, to evaluate any effect of lysine and valine on litter growth rate and sow performance between parities. Chi-square analysis was used to determine differences in distribution of days to estrus and percentages of sows returning to estrus, being rebred, and farrowing. LYSINE:VALINE RATIO FOR LACTATING SOWS 1855 Results All Sows. No interactions ( P >.10) were observed between valine and lysine for litter growth rate or sow BW and BF loss, indicating that their effects are independent. Lysine intake increased from 35.7 to 55.7 g/d as dietary lysine increased from.80% to 1.20% ( P <.001, Table 3). Increased lysine intake resulted in increased litter weaning weights ( P <.001), litter weight gain ( P <.002), litter weight gain per day ( P <.002), and average weanling pig weight ( P <.006) and reduced sow BW loss ( P <.001). Valine intake ranged from 28.9 g/d for sows fed.80% lysine and.64% valine to 67.3 g/d for sows fed 1.20% lysine and 1.44% valine (linear, P <.001). Increasing dietary valine had no effect ( P >.10) on litter weaning weight, litter weight gain, or average weaned pig weight. Dietary lysine and(or) valine had no effect ( P >.10) on number of pigs weaned, BF loss, or days to estrus. Sows Weaning Ten or More Pigs. Two thirds of the sows weaned 10 or more pigs (Table 4). For this group, litter weaning weight, litter weight gain, and pig weaning weights increased as lysine increased ( P <.001). Average daily feed intake increased ( P <.02) and sow weight loss decreased ( P <.001) for sows fed the high-lysine diet. Litter weaning weight, litter weight gain, and pig weaning weights increased as dietary valine increased (linear, P <.04; quadratic, P <.08). Dietary valine had no effect ( P >.10) on sow ADFI or weight loss. Increasing valine tended ( P <.07) to increase days to estrus from 5 to 6.3 d. Neither lysine nor valine had an effect on sow BF loss ( P >.10). Table 3. Effects of dietary lysine and valine on sow and litter performance during lactation: All sows a a Days of lactation and litter size after cross-fostering were used as a covariate. No. of sows Days of lactation No. pigs weaned Litter weaning weight, kg Litter weight gain, kg Litter weight gain/d, kg Avg. pig weaning weight, kg Initial sow BW, kg Sow weight loss, kg b Initial 10th rib backfat, mm Initial last lumbar backfat, mm th Rib backfat loss, mm c Last lumbar backfat loss, mm c ADFI, kg Lysine intake, g/d Valine intake, g/d Days to estrus

4 1856 RICHERT ET AL. Table 4. Effects of dietary lysine and valine on sow and litter performance during lactation: 10 or more pigs weaned per litter a a Days of lactation and litter size after cross-fostering were used as a covariate. No. of sows Days of lactation No. pigs weaned Litter weaning weight, kg Litter weight gain, kg Litter weight gain/d, kg Avg. pig weaning weight, kg Initial sow BW, kg Sow BW loss, kg b Initial 10th rib backfat, mm Initial last lumbar backfat, mm th Rib backfat loss, mm c Last lumbar backfat loss, mm c ADFI, kg Lysine intake, g/d Valine intake, g/d Days to estrus Sows Weaning Fewer Than 10 Pigs. In sows that weaned fewer than 10 pigs, increasing dietary lysine had no affect ( P >.10) on litter growth rate, number of pigs weaned, ADFI, or sow BF and weight loss (Table 5). Dietary lysine and valine intake increased ( P <.001) with increasing lysine. Litter weight gain and number of pigs weaned tended to numerically increase and then decrease (quadratic, P =.13) with increasing dietary valine, with maximum litter weight gain and number of pigs weaned observed at a 1:1 lysine to valine ratio. As expected, increasing dietary valine increased valine intake ( P <.001). Dietary Table 5. Effects of dietary lysine and valine on sow and litter performance during lactation: Less than 10 pigs weaned/litter a a Days of lactation and litter size after cross-fostering were used as a covariate. No. sows per treatment Days of lactation No. pigs weaned Litter weaning weight, kg Litter weight gain, kg Litter weight gain/d, kg Avg. pig weaning weight, kg Initial sow BW, kg Sow BW loss, kg b Initial 10th rib backfat, mm Initial last lumbar backfat, mm th Rib backfat loss, mm c Last lumbar backfat loss, mm c ADFI, kg Lysine intake, g/d Valine intake, g/d Days to estrus

5 LYSINE:VALINE RATIO FOR LACTATING SOWS 1857 Table 6. Effect of dietary lysine and valine on sow and litter performance during lactation: First parity sows a a Days of lactation and litter size after cross-fostering were used as a covariate. No. sows per treatment Days of lactation No. pigs weaned Litter weaning weight, kg Litter weight gain, kg Litter weight gain/d, kg Avg. pig weaning weight, kg Initial sow BW, kg Sow BW loss, kg b Initial 10th rib backfat, mm Initial last lumbar backfat, mm th Rib backfat loss, mm c Last lumbar backfat loss, mm c ADFI, kg Lysine intake, g/d Valine intake, g/d Days to estrus valine had no effect ( P >.10) on sow BW or BF loss or on ADFI. Parity Effects. For first parity sows, litter weaning weight ( P <.005), litter weight gain ( P <.002), average pig weaning weight ( P <.005), and feed intake ( P <.001) increased as dietary lysine increased (Table 6). Litter weaning weight (linear, P <.09) and litter weight gain (linear, P <.08) also tended to increase as dietary valine increased. For second parity sows, a lysine valine interaction was observed for litter weaning weight ( P <.08) and average weanling pig weight ( P <.05) (Table 7). Linear increases occurred in litter and pig weaning weights with increasing valine when sows were fed the.80% lysine diet. However, linear decreases in litter and pig weaning weights were observed with increasing valine when sows were fed 1.20% lysine. First and second parity sows had reduced weight loss ( P <.006) with increased dietary lysine. Backfat loss was not affected ( P >.10) by dietary treatment. Subsequent Litter Size. Subsequent litter size was not affected ( P >.10) by dietary lysine or valine (Table 8). Previous lactation litter size (<10 or 10 pigs) had no effect on subsequent litter size. However, 7 to 15% more sows nursing <10 pigs farrowed in the subsequent lactation. Discussion Tokach et al. (1993) and Richert et al. (1996, 1997) demonstrated the need for increased dietary valine for high-producing lactating sows. Considering all sows (Table 3), increasing dietary lysine increased litter weaning weight by 4.2 kg, litter weight gain by 3.7 kg, and average weaned piglet weight by.3 kg. Increased dietary lysine also reduced sow weight loss by 8.4 kg. Similar responses to increased dietary lysine (35.7 to 55.7 g/d) have been documented (Stahley et al., 1990; Johnston et al., 1993), especially with first parity sows (Tokach et al., 1992; King et al., 1993). Increasing dietary valine from 80 to 120% of lysine resulted in 50% (1.8 kg) of the increased litter weaning weights and litter weight gains observed with lysine. The valine response was not as great as that in previous research (Tokach et al., 1993; Richert et al., 1996, 1997); however, the valine and lysine requirements seem to be dependent on the milk production of the sow. Furthermore, the litter weaning weight and weight gain responses to valine in the present experiment were curvilinear; they tended to increase and then decrease in sows weaning fewer than 10 pigs per litter but decrease and then increase in sows weaning 10 or more pigs per litter. This is contrary to the findings of our previous research (Richert et al., 1996, 1997) in which we observed linear increases in litter weaning weight and litter weight gain with increasing dietary valine. We have no explanation for this variation in response between experiments. The response of lactating sows to these two amino acids seems to be unique in light of the traditionally held concept of protein synthesis being limited by the first-limiting amino acid. We observed a tendency to increased litter weaning weights with increasing

6 1858 RICHERT ET AL. Table 7. Effect of dietary lysine and valine on sow and litter performance during lactation: Second parity sows a a Days of lactation and litter size after cross-fostering used as a covariate. No. sows per treatment Days of lactation No. pigs weaned Litter weaning weight, kg Litter weight gain, kg Litter weight gain/d, kg Avg. pig weaning weight, kg Initial sow BW, kg Sow BW loss, kg b Initial 10th rib backfat, mm Initial last lumbar backfat, mm th Rib backfat loss, mm c Last lumbar backfat loss, mm c ADFI, kg Lysine intake, g/d Valine intake, g/d Days to estrus valine in sows fed.80% lysine, as well as a response to increasing lysine in sows fed low valine (80% of lysine). We also have observed this type of response in a similar study evaluating the effects of valine and isoleucine on sow and litter performance (Richert et al., 1997). One possible explanation is that valine is used by the mammary gland for purposes other than milk protein synthesis. At this time, we have no conclusive evidence what this non-milk protein synthesis role is; however, the possibilities include an energy source (Roets et al., 1979) or a source of N for nonessential amino acid synthesis, and(or) the ketoacid derivative may have a role in stimulating milk production (Nissen et al., 1994). Tokach et al. (1993) reported that the valine requirement is related to the number of pigs nursed and the related change in milk production. Similarly, in a review article, Pettigrew (1993) reported that the lysine requirement increases as daily milk production (litter size) increases. This observation was evident in this experiment; sows that had a greater demand for milk production (nursing 10 or more pigs) were responsible for most of the lysine and valine responses. For sows that weaned 10 or more pigs, litter weight gain increased by 4.6 kg with increased lysine and by 3.1 kg when valine was increased to 120% of lysine. This demonstrates greater lysine and valine responses in sows nursing 10 or more pigs and having higher milk production (as measured by litter weaning weights) than for sows weaning fewer than 10 pigs. For sows weaning 10 or more pigs and fed.80% lysine, valine increased litter weight gain by 2.2 kg at 120% of lysine compared with sows fed the 80% valine. For sows fed diets containing 1.20% lysine, litter weight gain increased by 4.4 kg at the 120% valine level compared with the 80% valine level in the high-lysine diets. In high-producing sows, the valine response in the high-lysine diet equaled the increase in litter weight gain from increasing dietary lysine and was additive to the lysine response. Thus, a 9-kg increase (i.e., 4.6 kg from lysine and 4.4 kg from valine) in litter weight gain resulted from feeding high-producing sows a diet containing high levels of lysine (1.20%) and valine (120% of lysine) compared with sows fed low levels of lysine (.80%) and valine (80 or 100% of lysine). The increased demand for milk production by sows nursing larger litters (10 or more pigs) likely increases the extraction rate of amino acids (Hanigan et al., 1991) by the mammary gland. This demand would increase the total requirement for amino acids (Pettigrew, 1993). Sows that weaned fewer than 10 pigs had only a.2 kg increase in litter weight gain as dietary lysine increased. The response to valine in sows weaning fewer than 10 pigs tended to be quadratic, with maximum litter weight gain at a 1:1 valine:lysine ratio. Providing a 1:1 valine:lysine ratio in the diet gave a greater response in litter weaning weights than simply increasing lysine in the diet. These results indicate that valine and lysine are required for maximum litter weight gain, and their ratio likely changes as milk production demand increases. Weight loss was 2.4 kg (1.20% lysine) or 9.6 kg (.80% lysine) greater for sows nursing at least 10 pigs than for sows nursing fewer than 10 pigs. Sow weight loss serves as an additional indicator of the difference in the demand

7 LYSINE:VALINE RATIO FOR LACTATING SOWS 1859 Table 8. Effects of dietary lysine and valine on sows subsequent litter size Item CV Lys Lin. Quad. Val Lys All Sows Total number of sows No. of sows removed No. of sows rebred Subsequent litter size No. of sows Born alive Still born Total born Farrowing rate, % pigs weaned Total no. of sows No. of sows removed No. of sows rebred Subsequent litter size No. of sows Born alive Still born Total born Farrowing rate, % <10 pigs weaned a Total no of sows No. of sows removed No. of sows rebred Subsequent litter size No. of sows Born alive Still born Total born Farrowing rate, % a Valine affected number of sows removed, rebred, and farrowed (chi-square; P <.11). for milk production between these two groups of sows. This finding supports previous research suggesting that increasing dietary lysine decreases sow weight loss during lactation (Stahley et al., 1990; Tokach et al., 1992; King et al., 1993). Valine had no effect on sow weight loss. Valine and lysine likely increase litter growth rate through separate mechanisms. The valine metabolite isobutyrate has been shown to increase the production of 12:0, 14:0, and 16:0 in goat s milk (Massart-Leen et al., 1986). This is similar to the response of the leucine metabolite b-hydroxy-b-methyl butyrate, which increases milk fat in lactating sows (Nissen et al., 1994). Another explanation for the increased litter growth rate with increasing valine is the possibility of increased prolactin or somatotropin levels. When increased levels of another branched-chain amino acid, leucine, were provided in the diet, plasma prolactin increased in lactating rats over time (Chugh et al., 1991). Also, Daniel et al. (1991) reported increased somatotropin levels when all three-branched-chain amino acids were infused in the lactating cow mammary gland. For first parity sows, significant increases occurred in litter weight gain with increased dietary lysine (5.2 kg) and valine (3.4 kg). However, for second parity sows, litter growth rate showed no response to increasing lysine and valine. The difference may have been due to second parity sows being 40.8 kg heavier than first parity sows, providing more muscle mass from which to draw amino acids for milk production. An additional factor leading to the different responses to lysine and valine may be that litters of first parity sows had 3.0 kg greater litter weight gain on the highlysine diet and 4.1 kg greater litter weight gain on the 120% valine:lysine diet than litters of second parity sows. First and second parity sows responded similarly to increased lysine with reduced sow weight loss. Valine had no effect on sow weight loss of first or second parity sows. The similarity in BW loss between parities indicates that lysine was limiting for both parities. The lack of response to valine by second parity sows could be related to their lower milk production rates and(or) the feeding and handling of the sows in their previous lactation (Hall et al., 1985; King et al., 1993). Contrary to our results, Tritton et al. (1993) observed an increase in subsequent litter size in gilts fed high-lysine diets during lactation. We observed a

8 1860 RICHERT ET AL. numerical increase (.8 pigs) in the number born live for sows that previously weaned fewer than 10 pigs and were fed increased lysine. However, for sows weaning more than 10 pigs, a reduction in pigs born live (.7) may indicate that their previous lactation milk production required even greater dietary lysine to achieve an increase in subsequent litter size. Tritton et al. (1993) observed a 1.2 pig increase in subsequent litter size for first parity sows fed 61 g/d lysine. However, for the lower producing sows in this experiment (weaning fewer than 10 pigs), 55 g/d of lysine was sufficient to increase subsequent litter size by.8 pigs. An additional factor in this experiment is that the last third of the sows developed swine dysentery between weaning and rebreeding and subsequently were medicated. This disease factor may have played an additional role in the subsequent litter size data. The numerical trend, with few sows per treatment, for increased subsequent litter size for lower producing sows serves as a potential indicator of being able to alter subsequent performance by previous dietary treatment and deserves further investigation. Implications Dietary lysine and valine levels should be based on the milk production of the sow. High-producing sows weaning 10 or more pigs and first parity sows require increased lysine (56 vs 36 g/d) and dietary valine (120% of lysine) to maximize litter growth rate. However, lower-producing sows require less lysine (.8%; 37 g/d) and valine (100% of lysine) to maximize litter growth rate. This trial suggests that valine and lysine act through different mechanisms to increase litter weaning weights. Literature Cited AOAC Official Methods of Analysis (15th Ed.). Association of Official Analytical Chemists, Arlington, VA. ARC The Nutrient Requirements of Pigs. Commonwealth Agricultural Bureaux, Slough, U.K. Chugh, K., V. Shaker, J. Kaur, A. S. Saini, and H. Lal Plasma prolactin and insulin changes on feeding leucine supplemented diet in rats. Horm. Metab. Res. 23:141. Daniel, T. E., A. H. Rakes, C. A. Zimmerman, and B. A. Hopkins Effect of arginine or branched chain amino acids in early lactation Holstein dairy cows fed corn silage. J. Dairy Sci. 74: 182 (Abstr.). Hall, D. D., G. L. Cromwell, T. J. Prince, D. A. Knabe, A. J. Clawson, C. V. Maxwell, P. R. Noland, D. E. Orr, and G. E. Combs Depletion of body fat reserves: An explanation for decreased second parity prolificacy in sows. J. Anim. Sci. 61(Suppl. 1): 316 (Abstr.). Hanigan, M. D., C. C. Calvert, E. J. DePeters, B. L. Reis, and R. L. Baldwin Whole blood and plasma amino acid uptakes by lactating bovine mammary glands. J. Dairy Sci. 74:2484. Johnston, L. J., J. E. Pettigrew, and J. W. Rust Response of maternal-line sows to dietary protein concentration during lactation. J. Anim. Sci. 71:2151. King, R. H., M. S. Toner, H. Dove, C. S. Atwood, and W. G. Brown The response of first-litter sows to dietary protein level during lactation. J. Anim. Sci. 71:2457. Massart-Leen, A. M., G. Peeters, G. Vandeputte-Van-Messom, E. Roets, and C. Burvenich Effects of valerate and isobutyrate on fatty acid secretion by the isolated perfused mammary gland of the lactating gland. Reprod. Nutr. Dev. 26: 801. Nissen, S., T. D. Faidley, D. R. Zimmerman, R. Izard, and C. T. Fisher Colostral milk fat percentage and pig performance are enhanced by feeding the leucine metabolite b- hydroxy-b-methyl butyrate to sows. J. Anim. Sci. 72:2331. NRC Nutrient Requirements of Swine, 9th rev. ed. National Academy Press, Washington, DC. Peterson, R. G Design and Analysis of Experiments. Marcel Dekker, New York. Pettigrew, J. E Amino acid nutrition of gestating and lactating sows. Biokyowa Technical Review #5. Nutri-Quest, Inc., Chesterfield, MO. Richert, B. T., R. D. Goodband, M. D. Tokach, and J. L. Nelssen Increasing valine, isoleucine, and total branched-chain amino acids for lactating sows. J. Anim. Sci. (In press). Richert, B. T., M. D. Tokach, R. D. Goodband, J. L. Nelssen, J. E. Pettigrew, R. D. Walker, and L. J. Johnston Valine requirement the high-producing lactating sow. J. Anim. Sci. 74: Roets, E., A. M. Massart-Leen, R. Verbeke, and G. Peeters Metabolism of [U- 14 C; 2,3-3 H]-L-valine by the isolated perfused goat udder. J. Dairy Res. 46:47. Rousselow, D. L., and V. C. Speer Valine requirement of the lactating sow. J. Anim. Sci. 50:472. SAS SAS/STAT User s Guide (Release 6.03). SAS Inst. Inc., Cary, NC. Stahley, T. S., G. L. Cromwell, and H. J. Monegue Lactational responses of sows nursing large litters to dietary lysine levels. J. Anim. Sci. 68(Suppl. 1):369 (Abstr.). Tokach, M. D., R. D. Goodband, J. L. Nelssen, and L. J. Kats Valine a deficient amino acid in high lysine diets for the lactating sow. J. Anim. Sci. 71(Suppl. 1):68 (Abstr.). Tokach, M. D., J. E. Pettigrew, B. A. Crooker, G. D. Dial, and A. F. Sower Quantitative influence of lysine and energy intake on yield of milk components in the primiparous sow. J. Anim. Sci. 70:1864. Tritton, S. M., R. H. King, R. H. Campbell, and A. C. King The effects of dietary protein on the lactation performance of first-litter sows. In: E. S. Batterham (Ed.) Manipulating Pig Production IV. Australasian Pig Science Association, Attwood, Victoria, Australia. Trottier, N. L., C. F. Shipley, and R. A. Easter Arteriovenous differences for amino acids, urea nitrogen, ammonia and glucose across the mammary gland of the lactating sow. J. Anim. Sci. 72(Suppl. 1):332 (Abstr.).

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