AMINO ACID REQUIREMENTS FOR BREEDING SOWS Sung Woo Kim1; and Guoyao Wu2 1 Texas Tech University;

2Texas A&M University ABSTRACT The current feeding program for lactating and gestating pigs is not based on actual amino acid needs for highlean type sows. The findings of our recent studies indicate ideal amino acid patterns for high-lean type sows during lactation and gestation periods. A single diet feeding of gestation sows with fixed daily allowance does not reflect actual their daily amino acid needs. Varying daily allowance may better satisfy amino acid needs but this will be associated with excess provision of energy. A phase feeding program for gestating sows during the early (d 0-70) and late (d 70-farrowing) gestation is an ideal one if individual farms have the resources for this application. A parity feeding regimen should be adopted to satisfy accurate amino acid needs for both primiparous and multiparous sows during lactation, because they exhibit differences in appetite and body weight loss. A proper balance among amino acids in diet is necessary to support the maximal productivity of breeding sows. INTRODUCTION Proteins are the building block of animal body and have additional important functions as enzymes, peptide hormones, antibodies, and regulators of gene expression (Wu and Self, 2005a). Proteins consist of specific sequences of amino acids and have dimentional structures. Amino acids are essential nutrients required for tissue protein synthesis and other vital metabolic functions in animals (Wu and Self, 2005b). Because excess amino acids are usually not stored in the body and subject to irreversible loss via oxidation and production of nitrogenous metabolites (e.g., ammonia, urea, and nitrate), amino acids must be supplied at least daily from diet. Pigs receive amino acids from various protein sources in feedstuffs and from supplements. Feed ingredients vary greatly with the content of amino acids. Available evidence shows that amino acid profiles in dietary proteins are an important determinant of rates of tissue protein synthesis. Because amino acids are needed not only for maintenance and growth but also for lactation and reproduction, gestating and lactating pigs have high requirements for dietary protein for supporting fetal growth and milk production. There are published reports of requirements for dietary protein to support reproduction in pregnant pigs. However, the concept of ensuring a high quality of dietary amino acids has received lesser attention from animal nutritionists. Identifying ideal profiles of amino acids for efficient protein synthesis in breeding pigs is an important goal of swine production and warrants intensive research efforts. In this paper, we review new information about amino acid and protein nutrition in breeding pigs. Additionally, we propose new concepts about feeding amino acids to breeding pigs. PROTEIN AND AMINO ACID REQUIREMENTS FOR PREGNANT SOWS Pregnant pigs need nutrients to support the growth of the conceptus (the fetus and associated membranes) as well as maternal maintenance and growth. Thus, dietary amino acids should be formulated to balance amino acid needs for various components such as conceptus gains, mammary tissue gain, maternal maintenance needs, and maternal tissue growth. Amino acid requirements for maternal tissue are greater for gilts and young sows compared with multiparous sows. The traditional feeding programs for modern pregnant sows do not provide accurate quantity and quality of amino acids, because significant genetic improvement has affected sow performances during the last two decades (MLC 1979 and 1999). On the basis of the growth of fetus, mammary glands, maternal maintenance, and maternal gain, we have proposed new requirements of protein and amino acids by gestating sows. Protein and amino acids for fetal growth As noted above, amino acids are essential nutrients for synthesizing molecules with enormous biological importance (Reeds and Hutchens, 1994) and fetal protein deposition. Amino acids are also important energy source for fetal growth (Bell et al., 1989). McPherson et al. (2004) characterized the development and growth of fetuses obtained from high-lean type sows. The amounts of proteins in various fetal tissues were measured

during different days of gestation (Table 1). Results of this study indicate that protein gains in various fetal tissues occur at different stages during fetal growth. Protein gains in fetal carcass, liver, GIT, heart, lung, kidney, and placenta were combined to obtain protein needs for the fetus during different days of gestation. Fetal protein gain was 0.25 g/d until d 68.5 of gestation but it was 4.63 g/d after d 68.5 of gestation indicating that protein needs for fetal tissue growth increases significantly (18.5-fold) as gestation progressed (Figure 1). Table 1 - Protein contents (g) in fetal tissues on different days of gestation. Data were adopted from McPherson et al. (2004). Data from heart, kidney, brain, and placenta were excluded in this table Day of gestation 45 60 75 90 102 110 SEMa b c cd d e f g Carcass 2.48 17.39 33.46 109.01 156.93 164.09 0.001 GITb 0.078c 0.602cd 1.505d 5.941e 7.082f 15.305g 0.00001 b c cd d e e Liver 0.391 1.864 2.386 6.377 4.992 7.567f 0.0001 b c cd d e f Lung 0.097 0.704 1.695 3.482 4.171 9.297g 0.0001 a Pooled standard error of the mean. b Within a row, protein content increased cubically as gestation progressed (P < 0.001). cdefg Within a row, means lacking a common superscript letter differ (P < 0.05).

Figure 1 - Fetal protein accretion (g) increases at d 68.5 of gestation (R2 = 0.99 and P < 0.001). The regression equation for before d 68.5 is 0.249 x (x 68.5) + 17.078, and for after d 68.5 is 4.629 x (x 68.5) + 17.078, where x is day of gestation. Wu et al. (1999) measured amino acid contents in fetal tissues during different days of the gestation period. These authors found that the profiles among some major essential amino acids (i.e., lys, thr, val, leu, ile, and met) in fetal pigs varied with the stage of gestation. Based on the data from McPherson et al. (2004) and Wu et al. (1999), protein and amino acid needs for fetal growth were calculated (Table 2). If a sow has 14 fetuses, she would need 0.27 g/d or 4.00 g/d lysine just for fetal tissue gain before or after d 70 of gestation, respectively. Table 2 - Protein and amino acid gain in McPherson et al. (2004) CP Lys D 0 to 70 of gestation % in protein 7.79 Gain, g/d/fetus 0.25 0.019 D 70 of gestation to farrowing % in protein 6.11 3.51 Gain, g/d/fetus 4.63 0.283 fetal tissues. Data were adopted from Wu et al. (1999) and Thr Trp Met Val Leu Ile Arg

4.04 1.22 2.23 5.30 8.12 3.56 6.45 0.010 0.003 0.006 0.013 0.020 0.009 0.016 1.20 1.99 4.55 7.16 3.07 6.84 0.162 0.056 0.092 0.211 0.332 0.142 0.317

Placental development is an important factor for fetal growth because a fetus receives nutrients through placental blood vessels (Ford, 1995). Knight et al. (1977) demonstrated that the placental growth was nearly maximized by d 60 of gestation and this supports the findings of McPherson et al. (2004) that the fetal needs

for amino acids increased after d 70 of gestation to support accelerated fetal growth fetal. Wu et al. (1998) showed that severe protein restriction to pregnant gilts decreased placental amino acid contents, which will in turn reduce fetal growth. Thus, maternal amino acid supply should match the needs for fetal growth. If a sow has 14 fetuses, her needs for lysine solely for the fetal tissue gain would be 4.0 g/d after d 70 of gestation. The traditional feeding of pregnant sows provides an average of 8.50 to 9.00 g true digestible lysine/d during the entire gestation period (NRC, 1998). It can be estimated that at least 45% of absorbed lysine would be needed only to support fetal tissue growth during late gestation for sows under this feeding program. Arginine (an essential amino acid for the fetus) plays important roles in fetal growth. This amino acid is particularly rich in porcine allantoic fluid (4-6 mM) during early gestation (Wu et al., 196). Tate of arginine deposition to fetal tissue proteins was one of the greatest among amino acids (Wu et al., 1999). Arginine is metabolized to nitric oxide (NO) in animal cells that functions as an endothelium-derived relaxing factor, a neurotransmitter, and a modulator of the immune response (Wu and Morris, 1998), indicating the significant metabolic roles of arginine in fetal development. Wu et al. (1998) have also demonstrated that reducing maternal dietary protein level decreased placental concentrations of free arginine and, therefore, reduced NO synthesis. The latter may contribute to impaired angiogenesis and growth of placental and endometrial tissues. Reduced arginine concentration in placenta likely leads to reduction in placental-fetal blood flow, nutrient supply from mother to fetus, and ultimately fetal growth retardation (Wu et al, 2004). Conventional corn- and soybean-meal based diets for gestating sows normally contain 0.8% to 1.0% arginine. However, given extensive catabolism of dietary arginine by intestinal mucosal enterocytes (Wu, 1998), it is not known whether feed-derived arginine can efficiently be used to support the needs of gestating sows. Research is needed to address this potentially important question. Protein and amino acids for mammary tissue development Protein content in mammary parenchymal tissue is highly correlated to DNA content (an indicator of functional mammary cell numbers) (Kim et al., 1999a), and therefore is a key determinant of the growth of nursing pigs suckling those mammary glands (Kim et al., 2000a). Improving the growth of the mammary gland during pregnancy, therefore, will be beneficial to litter weight gain after parturition. Mammary glands undergo physiological and morphological changes at the onset of pregnancy and grow substantially during gestation. Ji (2004) measured protein gain in mammary parenchymal tissues during different days of pregnancy. Protein gain in individual mammary glands was 0.14 g/d before d 80 of gestation and 3.41 g/d after d 80 of gestation (Figure 2). The pattern of protein accretion in the mammary gland obtained from the current study was similar to that reported by Whittemore (1996).

Figure 2 - Protein contents in individual mammary glands during gestation. Breakpoint of the mammary protein content (g) occurs at d 81.6 of gestation (R2 = 0.99 and P < 0.001) showing that the mammary protein accretion mainly occurred after d 81.6 of gestation; the regression equation before d 81.6 is 11.18 + 0.137 x (d - 81.6), and after d 81.6 is 11.18 + 3.409 x (d - 81.60). Amino acid profiles in mammary parenchymal tissues were measured by Kim et al. (1999a) and were adopted

to calculate amino acid needs for mammary parenchymal tissue growth during pregnancy (Table 3). Table 3 - Protein and amino acid gains in the porcine mammary gland. Data were adopted from Ji (2004) and Kim et al. (1999a) CP Lys Thr Trp Met Val Leu Ile Arg % in protein 7.50 4.25 1.17 1.98 5.70 8.37 4.13 6.13 Day of gestation Gain, g/d/gland 0 to 80 0.14 0.011 0.006 0.002 0.003 0.008 0.012 0.006 0.009 80 to farrowing 4.63 0.347 0.197 0.054 0.092 0.264 0.388 0.191 0.284 If a sow has 16 mammary glands, she would need 0.17 g/d lysine for mammary tissue gain until d 80 of gestation but 5.56 g/d lysine after d 80 of gestation. This is because protein or amino acid needs increase significantly (33-fold) as gestation progressed. The traditional feeding of pregnant sows provides an average of 8.50 to 9.00 g true digestible lysine/d during the entire gestation period (NRC, 1998). It can be estimated that at least 60% of absorbed lysine would be needed only to support mammary tissue growth during late gestation for sows under this feeding program. Protein and amino acids for maternal tissue gain and maintenance Amino acids are continuously needed for maintaining normal body functions during pregnancy. In the case of pregnant gilts and young sows, additional amino acids are needed for their own tissue growth. NRC (1998) provides the amount and profile of amino acids needed for body maintenance that are correlated to the body weight. Ji et al. (2005a) measured the amount of proteins in various maternal tissues from high-lean type gilts during different days of pregnancy and calculated the changes in protein gains as gestation progressed (Table 4). Protein gain in maternal carcass increased during mid pregnancy but decelerated as gestation progressed (cubical changes). Protein gain in maternal carcass, maternal liver, GIT, and remaining viscera did not follow the patterns for the gains of the fetus and mammary glands. Kim and Easter (2001) measured amino acid profiles in maternal tissues, including carcass, liver, GIT, remaining viscera, and uterus from first- parity sows and these measurements were used to calculate the amount of amino acids needed to support maternal tissue gain from Ji et al. (2005a). Table 4 - Protein contents (g) in maternal tissues on different days of gestation. Data were adopted from Ji et al. (2005a) Day of gestation CP, kg 0 45 60 75 90 102 112 PSDa b Carcass soft tissue 14.88 16.47 16.10 18.36 18.20 18.46 18.97 1.13 Liverc 0.44 0.37 0.38 0.41 0.37 0.41 0.37 0.03 GITbd 0.57 0.57 0.56 0.59 0.51 0.48 0.49 0.04 Remaining viscerae 0.48 0.48 0.45 0.54 0.48 0.45 0.51 0.05 Uterus + placentac 0.14 0.37 0.60 0.96 0.85 0.98 0.89 0.15 a Pooled standard deviation. b Cubic response (P < 0.01). c Linear response (P < 0.05). d Gastrointestinal tract (GIT) is composed of the empty stomach, empty small and large intestine, cecum, and rectum . e Remaining viscera is the remaining parts of organs and tissues after the removal of carcass, liver, pancreas, kidney, spleen, GIT, and reproductive tract. Concept of providing protein and amino acids during gestation Feeding pregnant sows should be based on accurate protein requirement and ideal amino acid pattern. It is well documented that gestation nutrition is closely correlated to the performance of milk production during lactation. Obese pigs at farrowing possess a low voluntary feed intake during lactation (Sinclair et al., 2001), which will result in an excessive loss of body tissues at weaning, delay days for returning to estrus after weaning, and even cause reproduction failure (Jones and Stahly, 1995). Thus, restricting nutrient intake

during gestation to prevent excessive fat deposition is the most popular practice. However, severe protein restriction would not be beneficial for the performance or health of gestating mammals (Wu et al., 2004). Obtaining accurate protein and amino acid requirements, therefore, is a key factor for the successful management of pregnant pigs. Amino acid profile in the diet is another important factor to consider because feeding diets with unbalanced amino acid pattern causes a reduction in protein synthesis and an increase in amino acid oxidation in the body (Voet and Voet, 1990). These excessive amino acids due to decreased protein synthesis are instead used as a source of energy and precursors for fat synthesis, which will increase fat deposition during pregnancy and nitrogen excretion to the environment. Thus, an ideal dietary amino acid pattern for feeding pregnant sows can increase the efficiency of protein synthesis, reduce unnecessary loss of essential amino acids, and reduce unnecessary fat deposition. When protein gains in maternal tissues and the conceptus were combined, amino acid requirements can be calculated for different stages of pregnancy (Ji, 2004). Due to a significant increase in protein gains in mammary glands and fetuses during late pregnancy, overall protein gain in pregnant sows also increased after d 70 of gestation. When the amounts of protein (or lysine) needed for maternal maintenance, maternal tissue growth, fetal growth, and mammary gland growth are all combined, pregnant gilts accumulate 148 g protein/d (or 6.83 g lysine/day) from the conception to d 70 of gestation and 330 g protein/d (or 15.26 g lysine/day) from d 70 of gestation to parturition. This finding clearly indicates that sows require different amounts of amino acids depending on the stage of gestation to support growth and maintenance (Table 5). In a recent study, Ji et al. (2005a) found that pregnant gilts require an increased amount of amino acids to support maternal carcass gain. Multiparous sows or mature sows may not need additional amino acids to support their own body growth. However, multiparous sows will have increased amino acid needs for maintenance as their body weights increase during pregnancy. Table 5 - Amino acid needs for tissue gains and maintenance of pregnant gilts before and after d 70 of gestation. Adopted from Ji et al. (2005a), Ji (2004), and NRC (1998) Lys Thr Val Leu Ile Arg D 0 to 70 of gestation (g/d) AA accretion 3.93 1.94 2.52 3.77 2.03 3.74 AA needs for accretion 5.13 2.53 3.29 4.92 2.65 4.88 AA needs for maintenance 1.64 2.48 1.10 1.15 1.23 1.23 AA needs for A+M 6.83 5.42 4.44 6.03 4.00 6.09 D 70 to 112 of gestation (g/d) AA accretion 7.00 3.56 4.64 7.11 3.53 7.25 AA needs for accretion 13.34 6.78 8.84 13.55 6.73 13.82 AA needs for maintenance 1.78 2.69 1.19 1.25 1.34 1.34 AA needs for A+M 15.26 10.86 10.13 14.53 8.46 14.93 Ideal amino acid patterns were obtained for pregnant gilts to support their maintenance and growth, including maternal tissues, fetuses, and mammary glands, using the information from Kim et al. (1999a), Kim and Easter (2001), McPherson et al. (2004), and Ji et al. (2005a). It was interesting to observe that the ideal amino acid patterns vary with the stage of gestation (Ji, 2004). Ideal amino acid patterns for pregnant gilts were different between before and after d 70 of gestation. This further indicates that optimal amino acid profiles in diet depend on the stage of gestation to support both growth and maintenance needs (Table 6). Current feeding strategy for pregnant sows involves a single diet feeding with fixed or adjusted daily allowance. As shown in Table 5, the sows require nearly 100% more amino acids after d 70 of gestation compared with before d 70 of gestation. Adjusting daily feed allowance can be used to support more accurate amino acid requirements. However, dietary energy should not be provided in excess by simply increasing daily feed allowance to match amino acid requirements during late pregnancy. It would be, therefore, ideal to have different types of gestation diets with different amino acid profiles to reflect accurate amino acid needs as gestation progresses without increasing energy content in diet. The phase feeding of pregnant pigs can be adopted if it is practically feasible on individual farms.

Table 6 - Estimate of ideal amino acid patterns for protein accretion and maintenance in pregnant gilts Lys Thr Val Leu Ile Arg D 0 to 70 of gestation For accretion 100 49.4 64.1 95.9 51.8 95.3 For maintenance 100 151.0 67.0 70.0 75.0 75.0 Accretion + maintenance 100 79.4 65.0 88.3 58.6 89.3 D 70 to 112 of gestation For accretion 100 50.8 66.3 101.6 50.4 103.6 For maintenance 100 151.0 67.0 70.0 75.0 75.0 Accretion + maintenance 100 71.2 66.4 95.3 55.5 97.9 Ji et al. (2005b) measured blood urea nitrogen in the pregnant gilts fed either corn-soybean meal based diets or diets with the estimated ideal amino acid patterns. We found that the gilts fed a diet with an ideal amino acid pattern had a decreased level of blood urea nitrogen, indicating that amino acid oxidation was decreased (or amino acid utilization for tissue protein synthesis was improved) (Figure 3). It remains to be determined whether pregnant sows receiving diets with both adjusted amounts and ideal amino acid profiles perform better than sows on traditional single-diet feeding.

Figure 3 - Serum urea nitrogen contents in pregnant gilts fed either corn-soybean meal diet or corn-soybean meal diet with crystalline amino acids to match the ideal dietary amino acid pattern. Protein and Amino Acids for Lactating Sows Amino acid metabolism during lactation is directed favorably to milk protein synthesis. Even during a catabolic status, sows synthesize milk protein by mobilizing their body proteins to provide free amino acids, which renders the lactation period physiologically and nutritionally unique and distinct to other stages of the pigs life. Lactating sows often lose their body proteins to support milk production and an excessive protein loss will result in a nutritional deficiency, ultimately impairing normal reproductive performance for subsequent parities, including delayed return to estrus, reduced litter size, or even reproduction failure (Kirkwood et al., 1987). Thus, a feeding strategy for lactating sows should focus on minimizing body protein loss by providing high quality dietary proteins. Modern sows are highly prolific but have a low appetite. It is, therefore, a challenge to animal nutritionists to increase sows feed intake during lactation. Increasing dietary protein content and improving the quality of dietary protein are two practical feeding strategies for lactating sows. The amino acid requirements of modern high-lean type sows have been investigated in recent years, which reflect the significant genetic improvement of sows (Kim and Easter, 2003; Kim et al., 2001a). Several studies were also conducted to investigate amino acid needs for mammary gland growth (Kim et al., 1999abc; Kim et al., 2000b; Nielsen et al., 2002), because improving mammary gland growth is directly related to increased litter weight gain (Kim et al., 2000a).

Protein and amino acids for mammary gland growth The mammary gland is a functionally major tissue for lactating sows because of its metabolic importance in synthesizing and secreting milk. Milk production is directly related to litter weight gain and litter weaning weight. Of note, lactating mammary glands continue to grow as lactation progresses. We have reported that 1.0 g/d of lysine (or 7.0 g/d of essential amino acids) is incorporated into mammary tissue protein for sows with 10 nursing pigs (Kim et al., 1999a). However, non-suckled extraneous mammary glands undergo substantial involution during the first 7 to 10 d of lactation (Kim et al., 2001b). Trottier et al. (1997) measured the amount of essential amino acids taken up by mammary glands (188.5 g/d), secreted as milk proteins (139.5 g/d), and retained in mammary gland (49.0 g/d). Among the 49.0 g/d retained essential amino acids, 7.0 g/d (14%) were accreted as a mammary tissue and the remaining 76% may be either transformed to other nonessential amino acids or oxidized as energy (Richert et al., 1998) (Table 7). Table 7 - Amino acids retained or accumulated in mammary gland of lactating sowsa Lys Thr Trp Met Val Leu Ile Retained, g/d 3.2 4.8 0.8 1.7 6.9 12.9 7.4 Tissue, g/d 1.01 0.56 0.15 0.28 0.79 1.16 0.58 Disapperance, % 68.2 88.4 80.0 83.2 88.5 91.0 92.2 a Adopted from Trottier et al. (1997) and Kim et al. (1999a). Kim et al. (1999b) demonstrated that the mammary gland growth can be affected by dietary amino acid and energy intakes during lactation. Mammary gland growth in lactating sows is maximized when a sow consumed 55 g total lysine and 16.9 Mcal ME per day during lactation (Figure 4). The amount of amino acids accreted in mammary tissue is affected by the litter size. Lysine (or essential amino acids) accretion was increased by 0.13 g/d (or 1.20 g/d) for an increase in one pig in a litter during 21-d lactation (Kim et al. 1999c). Nielsen et al. (2002) further measured the amount of lysine (or essential amino acids) taken up by mammary glands of the sows with different litter sizes (1.92 g lysine/d (or 7.65 g essential amino acids/d). Considering Nielsens data, amino acid needs for an additional pig in litter size would be higher than actual tissue accumulation (Table 8).

Figure 4 - Protein contents in individual mammary glands (g/gland) from sows received different amounts of energy and protein during the lactation (Adopted from Kim et al., 1999b). Table 8 - Amino acid use by mammary gland as litter size increased (g/d/pig increase)a Lys Thr Trp Met Val Leu Ile Retained, g/d 1.92 1.00 0.47 1.35 2.08 0.83 Tissue, g/d 0.13 0.18 0.05 0.08 0.24 0.35 0.17 Disappearance, % 6.8 18.0 17.0 17.8 16.8 20.5 a Adopted from Kim et al. (1999c) and Nielsen et al. (2002).

Protein and amino acids for milk production and litter growth Milk production is relatively unaffected by a modest dietary protein restriction because a sow has remarkable capacity to buffer by mobilizing her body protein to support amino acids needed for milk protein synthesis (Revell et al., 1998). However, severe protein restriction during lactation decreased milk production (Jones and Stahly, 1999; Knabe et al., 1996). Milk production can respond to a high protein diet with an increase in milk production (King et al., 1993). In an attempt to increase milk production through feeding a high protein diet, it is important to consider the balance of amino acids especially among limiting essential amino acids (Copper et al., 2001). Dietary protein fortification through stomach cannula did not increase milk production but decreased maternal tissue loss during lactation (Pluske et al., 1998). The concentrations of protein-bound amino acids are relatively in sows milk between Days 7 and 21 of lactation (Wu and Knabe, 1994). Mature sow milk contains 5.2% of crude protein (Tilton et al., 1999; Renaudeau and Noblet, 2001). However, this crude protein consists of significant amounts of urea and ammonia nitrogen (Wu and Knabe, 1994). Thus, true protein levels in milk should be determined by quantifying individual amino acids. To estimate the intake of amino acids from milk, it is necessary to properly express the values of amino acid concentrations in the milk (e.g., g/L whole milk). Available evidence shows that the profiles of amino acids in plasma (Wu et al., 1999) differs remarkably from the patterns of amino acids taken up by mammary glands or those in sows milk (Trottier et al., 1997) (Table 9). These differences in amino acid patterns reflect different rates of transport for amino acids by mammary tissue and extensive transformation of amino acids (synthesis and catabolism) in mammary tissue (OQuinn et al., 2002). Table 9 - Amino acid patterns relative to lysine from tissue mobilization, milk protein, mammary tissue gain, and corn-soybean meal based lactation diet. Adopted from Kim et al. (1999a; 2001a) and King et al. (1993)a Tissue mobilization Milk production Mammary growth Dietary sourceb Lys 100 100 100 100 Thr 42 59 58 70 Val 77 77 78 90 Leu 101 114 116 176 Ile 59 60 58 78 Arg 124 65 89 125
a b

Numbers are relative ratios to lysine. Amino acid ratio relative to lysine that is provided in a conventional corn-soybean meal diet (71.1% corn and 22.8% soybean meal containing 17% CP and 0.87% lysine) based on true ileal amino acid digestibility values for lactating sows (Stein et al., 1999, 2001).

Protein and amino acids for optimal body conditions When sows do not receive adequate amounts of dietary amino acids, maternal tissue proteins (particularly skeletal muscle proteins) are mobilized to support milk production. Excessive maternal protein mobilization often results in reproduction failure for the next parity (Jones and Stahly, 1995). Establishing nutrient requirements for lactating sows, therefore, is not limited to maximizing milk yield for nursing pigs, but extends also to maintaining an optimum body condition for subsequent parities (NRC, 1998; Kim and Easter, 2003). Amino acid mobilization occurs from various tissues of the sows with different rates (Escobar, 1998; Kim and Easter, 2001). Muscle is the major amino acid donor during food deprivation or inadequate provision of dietary protein, whereas the reproductive tract contributes the largest portion of its own amino acids (Kim and Easter, 2001). Dourmad et al. (1998) reported that high-producing sows need at least 55 g/d of dietary lysine for the minimal weight loss and this requirement is closed to the lysine need for the maximal mammary gland growth (55 g/d) suggested by Kim et al. (1999b). However, maintaining sows in an anabolic status during lactation alone may not be sufficient for improving sows fertility (Zak et al., 1998). Thus, the primary target should be minimizing body protein loss. Concept of providing amino acids during lactation: Ideal protein Kim et al. (2001a) identified an ideal dietary amino acid pattern for lactating sows with a concept that different amino acid patterns among tissue protein, milk protein, and dietary protein would affect the final amino acid pattern needed for sows during lactation. The content of some essential amino acids in milk

protein is relatively higher than that of amino acids released from tissue mobilization and in a common cornsoybean meal based diet. Amino acids needed for mammary gland growth may also affect ideal dietary amino acid pattern. Considering these factors, the ideal dietary amino acid patterns for lactating sows likely change dynamically, while responding to the expected maternal protein loss during lactation (Table 10). Table 10 - Ideal dietary amino acid patterns and the order of limiting amino acids for lactating sows (Adopted from Kim et al., 2001a) Level of tissue mobilization Kim et al. (2001a) NRC (1998) a 100%b 80% 40% 10% 0% Ideal amino acid pattern relative to lysine Lys 100 100 100 100 100 100 Thr 75 69 63 60 59 62 Val 78 78 78 77 77 85 Leu 128 123 118 115 115 114 Ile 60 59 59 59 59 56 Arg 22 38 59 69 72 56 Order of limiting amino acids 1st Thr Lys Lys Lys Lys Lys 2nd Lys Thr Thr Val Val Val 3rd Val Val Val Thr Thr Thr
Ideal amino acid pattern from NRC (1998) does not respond to the level of tissue mobilization. 100% level means when about 50% of amino acids in milk were quantitatively equivalent to the amount of amino acids from tissue protein mobilization. c Based on assumption that sows were provided corn (71%) and soybean meal (23%) based diet.
b a

Figure 5 - Litter weight gain of first parity sows and weight loss of second parity sows fed either cornsoybean meal diet or corn-soybean meal diet with crystalline amino acids to match the ideal dietary amino acid pattern. The body condition and expected levels of amino acid mobilization are important factors that must be considered in designing diets for lactating sows. This dynamic ideal protein concept would allow for a more precise estimation of amino acid needs for lactating sows. For sows possessing a low voluntary feed intake

and substantial tissue mobilization during lactation (i.e., primiparous and second parity sows), threonine is a critical amino acid, whereas valine becomes increasingly important for sows having a high feed intake and limited tissue mobilization (i.e., multiparous sows) during lactation (Kim et al., 2001a). However, lysine is the primary limiting amino acid in both cases. To apply the dynamic ideal protein pattern and the order of limiting amino acids, lactation diets can be designed for individual sows based on their expected levels of tissue mobilization during lactation. Recently, Kim et al. (2004) demonstrated that the first parity sows fed diets with amino acids that are ideally balanced for their needs had improved lactation performance by increasing litter weight gain or decreased body weight loss as compared to those fed common corn-soybean meal diets (Figure 5). CONCLUSION Establishing optimal requirements of amino acids (both quality and quantity) is critically important for maximizing the production performance of gestating and lactating sows. Recent research demonstrates that the current feeding strategy for these sows may not accurately provide amino acid needs. Gestation feeding program should be revised to include increased provision of amino acids after d 70 of gestation to support increased protein use for fetal and mammary gland growth. Ideal patterns of dietary amino acids dynamically change after d 60 of gestation, which can be applied to feeding gestating sows. If a two-phase-feeding strategy is adopted, it will provide ideally balanced amino acids for fetal and mammary gland growth while minimizing unnecessary fat gain during gestation. In addition, lactation feeding program needs to be modified to better reflect changes in feed intake or weight loss of sows and to enhance milk production. If a parity feeding is practically feasible, first and second parity sows should be fed different diets than those for multiparous sows, because of their differences in ideal dietary amino acid patterns, voluntary feed intake, and body weight loss. Diets for breeding pigs should be formulated according to accurate amino acid requirements and ideal dietary amino acid patterns. This knowledge can be gained through a better understanding of the metabolism and biology of pigs, which will ultimately lead to improved sow performance and overall efficiency of swine production. ACKNOWLEDGMENTS
The work in our laboratories was supported in part by funds from Texas Tech University, Texas Agricultural Experiment Station, Texas A&M University, and United States Department of Agriculture. We thank our students and technicians for their important contribution to our research programs.

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