Effect of Protein and Lysine Levels in the Diet on Body Gain Composition and Energy Utilization in Growing Pigs

J. Noblet, Y. Henry and S. Dubois J Anim Sci 1987. 65:717-726.

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EFFECT OF PROTEIN AND LYSINE LEVELS IN THE DIET ON BODY GAIN COMPOSITION AND ENERGY rUTILIZATION IN GROWING PIGS 1
J. Noblet, Y. Henry and S. Dubois 2 I n s t i t u t National de la Recherche A g r o n o m i q u e St. Gilles 35590 l'Hermitage, France
ABSTRACT

Eight replicates of four Large White littermate female pigs were used to evaluate the effect of protein and lysine levels in the diet on the efficiency of protein and energy utilization. In each replicate, one pig was slaughtered at about 20 kg live weight and the others received three diets that contained (per Meal digestible energy) 37.5 and 2.00 g (diet pl), 37.5 and 2.35 g (diet pL) or 45.0 and 2.35 g (diet PL) of digestible protein and lysine, respectively. Pigs were slaughtered after a 7-wk period. Tissue and chemical composition of the gain and energy and nitrogen gain were determined by using the comparative slaughter technique. Metabolizable energy (ME) intakes were similar in the treatments. Pigs fed the pl diet had a smaller body weight and muscle gain and retained less nitrogen and more lipids than pigs fed pL and PL diets. The decrease in the level of nonessential nitrogen in the diet (pL vs PL) did not affect body weight and muscle gain and the amount of nitrogen retained in muscle tissues. However, pigs given the PL diet had a higher total nitrogen retention and a lower fat deposition and exhibited a higher heat production. For each gram of additional protein catabolized for energy purposes (PL vs pL), heat production was increased by 1.8 kcal. The amount of lysine per unit of muscle gain (38 g/kg) or protein deposited (120 g/kg) was independent of protein and lysine levels in the diet. Estimates of energy (indirect calorimetry) and nitrogen (balance technique) retention were also obtained on the same animals; results were comparable with data obtained by direct measurements. (Key Words: Pigs, Lysine, Protein Intake, Energy Balance, Protein Balance, Body Composition.)

I ntroduction

Feed conversion efficiency and growth rate of pigs fed to appetite are n o t significantly affected b y dietary protein or nonessential n i t r o g e n c o n t e n t when the levels of essential a m i n o acids are m a i n t a i n e d adequate to meet the requirements (Wahlstrom and Libal, 1974; Bereskin et al., 1976; Sharda et al., 1976; Easter and Baker, 1980; Noblet et al., 1980; Stahly et al., 1981; Asche et al., 1985). However, most authors have observed a t e n d e n c y for increased b o d y fatness with low protein diets. The r e d u c t i o n of non-essential nitrogen level in the diet is also associated with a decreased nitrogen loss in urine (Sharda et al.,

1Appreciation is expressed to EUROLYSINE S.A., 16, rue Ballu, PARIS for providing L-lysine'HCl and DL-tryptophan and for partial funding of this study. a The authors gratefully acknowledge P. Ecolan, M. Fillaut, J. Lebost and A. Roger for technical assistance and Dr. A. J. Lewis (Univ. of Nebraska) for critical evaluation of the manuscript. Received November 5, 1986. Accepted April 21, 1987.

1976; Russell et al., 1983) and a s u b s e q u e n t lower c o n t r i b u t i o n of d e a m i n a t e d protein to ME supply. In addition, the efficiency of utilization of digestible n u t r i e n t s for maintenance or fat deposition is lower for d e a m i n a t e d protein t h a n for carbohydrate or fat ( H o f f m a n n and Schieman, 1971; Schulz, 1975; Just, 1982). A decrease in the dietary protein level and the s u b s e q u e n t e n r i c h m e n t in carbohydates or fat for the same level o f limiting essential a m i n o acid (i.e., lysine) would allow an improved efficiency of ME utilization, resulting in a t e n d e n c y for increased carcass fatness. The objective of the present e x p e r i m e n t was to study the effects of a r e d u c t i o n in protein level with or w i t h o u t lysine s u p p l e m e n t a t i o n on energy and nitrogen balance in growing pigs. The m e t h o d s used were comparative slaughter and balance techniques (indirect calorimetry and nitrogen balance). In addition, the effects of protein and a m i n o acid supply o n tissue a n d chemical c o m p o s i t i o n of weight gain also were investigated.
Experimental Procedure

Experimental Design. Eight replicates of

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NOBLET ET AL. levels that met or exceeded requirements (45 and 2.35 g/Mcal DE for digestible protein and lysine, respectively). After a 7-wk period, animals were slaughtered. The comparative slaughter m e t h o d was used to measure the a m o u n t , nature and localization o f weight gain and retention. In addition, energy (indirect calorimetry) and nitrogen (N) balances were carried o u t on six of the eight replicates during three, 2-wk periods (wk 1 and wk 4, wk 2 a n d wk 5 and wk 3 and wk 6). Week 0 was considered as the first week of the experiment. Each animal was measured during one of these three periods, the three periods for each t r e a t m e n t (pl, pL or PL) being represented once in a series of three replicates. One respiration chamber was used for the e x p e r i m e n t with successive replicates. In addition to the effect of dietary treatment, the results of the balance measurements provided

four Large White littermate female pigs were used in the experiment. In each replicate, pigs were chosen at weaning ( a b o u t 25 d of age), and fed individually a standard diet to equalize b o d y weights during the post-weaning phase. At a b o u t 20 kg live weight (mean: 19.5 + .5 kg), one pig was slaughtered (control) and the three others were given three different diets composed of corn and soybean meal (table 1). Two diets (low protein - low lysine, or pl; and low protein-high lysine, or pL) supplied less protein than r e c o m m e n d e d [37.5 g digestible protein/Mcal digestible energy (DE); INRA, 1 9 8 4 ] . Diet pl was inadequate in lysine (2.00 g/Mcal DE) while diet pL was fortified with Llysine HC1 and D L - t r y p t o p h a n to meet lysine and other essential a m i n o acid requirements (2.35 g lysine/Mcal DE; INRA, 1984). The third diet (high protein-high lysine, or PL) had protein, lysine and other essential amino acid

TABLE 1. COMPOSITION AND ANALYSIS OF DIETS Diet pL 37.5 a 2.35 b

pl Item Composition (%) Yellow corn Soybean meal Corn gluten meal Beet molasses Dicalcium phosphate Calcium carbonate Salt Trace minerals, vitamins and amino acids mixture c Analyzed levels (as fed) Gross energy, kcal/kg Crude protein, % Ash, % Crude fiber, % Neutral detergent fiber, % Acid detergent fiber, % Lipid, % Lysine, % Threonine, % Tryptophan, % (calculated) aDigestible protein, g/Mcal DE. bLysine, g/Mcal DE. 37.5 a 2.00 b

PL 45.0 a 2.35 b

75.35 16.40 3.00 2.20 1.20 .50 .35 3,827 15.3 5.1 2.0 8.2 2.2 3.2 .67 .52 .17

76.35 16.40 3.00 2.20 1.20 .50 .35 3,833 15.3 5.1 2.0 8.1 2.3 3.2 .80 .52 .17

69.75 21.50 1.50 3.00 2.20 1.20 .50 .35 3,849 17.8 5.4 2.3 8.1 2.3 2.9 .81 .63 .17

Cprovided per kg of diet: Zn, 120 rag: Fe, 44 rag; Cu, 5 nag; Co, .08 mg; Mn, 15 mg; Se, .18 mg; I, .8 mg; vitamin A, 5,000 IU; vitamin D, 1,000 IU; vitamin E, 10 IU; riboflavin, 4 mg; panthotenic acid, 10 mg; choline, 5 rag; vitamin B,2, 20 ~g; biotin, 200 #g; DL - tryptophan, .3 g (diets pl and pL); glycine, 1.5 g (diet pl); lysine, 1.5 g (1.9 g L-lysine-HCl; diet pL).

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EFFECT OF DIETARY PROTEIN AND LYSINE ON PIGS information about the effect of week of experiment and interaction between week of experiment and dietary treatment. Housing and Feeding. Pigs were housed individually in fiat-deck cages in a temperaturecontrolled room (20 + 2 C). During energy and N balances, the pigs were kept individually in metabolism cages located in respiration chambers (Noblet et al., 1985). The temperature within the chamber was 22 C, this temperature being at or above the critical temperature (Close and Mount, 1978). Food was offered in dry pellets, in two meals (0900 and 1500) and adjusted each day on the basis of 120 g of feed/kg metabolic body weight (w'TS). This feeding level corresponded to 80 to 100% of ad libitum intake. Body weight was the mean of the three individual weights within a replicate. The pigs were weighed each week. Feed refusals and spillage were collected. Water was available ad libitum. For each treatment, samples of feed were collected each week and analyzed for moisture content. For each replicate, an aliquot of the feed given during the whole experiment was also prepared for analysis.

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Nutrient Digestibility, N Retention and Respiratory Exchanges. Pigs were kept in the
respiratory chamber for a 7-d period. Feces were collected daily, pooled and at the end of the period, weighed, mixed, subsampled and freeze-dried for analysis. Similarly, urine was collected daily, pooled and, at the end of the period, weighed and sampled for analysis. Water that condensed in the chamber was weighed and sampled at the end of the period. Finally, an aliquot of the outgoing air was bubbled through a sulfuric acid solution. Determination of N content in condensed water and outgoing air allowed determination of N losses in the air. Oxygen consumption and carbon dioxide production were measured daily. Heat production was calculated from gas exchanges and urinary N losses according to the formula proposed by Brouwer (1965). In each replicate, feed and feces were analyzed for moisture, ash, N and fat according to AOAC (1975) methods. Gross energy was measured using an adiabatic bomb calorimeter. Nitrogen in urine, condensed water and outgoing air was measured on fresh material, whereas energy content of urine was obtained after freeze-drying approximately 50 ml in small polyethylene bags. A composite sample of each diet, obtained from the combination of samples

of the eight successive replicates was analyzed for crude fiber, cell wall components (Van Soest and Wine, 1967), amino acids and N (AOAC, 1975). Metabolizable energy (ME) intakes were calculated as the difference between energy values of feed intake and those of feces, urine and methane. Methane production was estimated from previous experiments (Noblet et al., 1985) as .4% of gross energy intake. Nitrogen retention was calculated as the difference between N intake and N in feces, urine, condensed water and outgoing air. Retained energy and its partition between protein and fat were calculated according to the method described by Noblet et al. (1985). Comparative Slaughter Method. After a 16 h-fast, body weight (BW) was determined and animals were sacrificed by electrical stunning and exsanguination. At slaughter, blood, bristles and visceral organs were collected separately and weighed. The digestive tract was weighed after emptying. A fraction including head, feet and tail was also constituted and weighed. Each half of the eviscerated carcass was weighed immediately after slaughter and after a 20-h chilling. On the day after slaughter, one-half of each carcass was dissected in four fractions: muscles (including intermuscular fat), adipose tissues, skin and bones. These different fractions were weighed and placed in plastic bags and stored at - 2 0 C. For each pig, the different fractions were finely ground and homogenized separately. From homogenates of these fractions, aliquots of the following compartments were prepared: muscles, carcass (muscles, fatty tissues, skin and bones), non-carcass (blood, bristles, organs and head + feet + tail) and empty body weight (EBW: carcass + non-carcass). Each compartment was analyzed for dry matter, ash, N, ether extract (referred to as lipid) and gross energy according to AOAC (1975) methods. Protein content was estimated as 6.25 times the N content. Finally, an aliquot of freeze-dried samples of empty body was constituted for each dietary treatment (eight pigs/sample) and analyzed for amino acid content. Anatomical and chemical composition of experimental animals at the beginning of the experiment were calculated for each replicate from composition of the control littermate. It was assumed that the composition of EBW and the ratio EBW: BW were identical for littermates. The weight and amount of nutrients and energy

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NOBLET ET AL. increased (P<.01) from wk 1 to wk 6 : 8 7 . 5 to 90.0% and 81.3 to 86.6% for energy and nitrogen, respectively. In the absence (P>.10) of an interaction between treatment and week number, the pooled data for each dietary treatment are presented in table 2. Digestibility coefficients of energy and N were higher for diet PL, the difference being significant only for N. This resulted in higher DE (P<.10) and ME (not significant) contents of that diet. Because of the higher excretion of N in urine, the ratio of ME:DE was lower (P<.05) for the PL diet. Reduction of protein and essential amino acid supply in the pl diet, compared to the PL diet, resulted in a significant decrease in average daily gain of BW, EBW, carcass and muscles (table 3). On the other hand, adipose tissue gain was higher in pigs fed the pl diet. The decrease in the level of nonessential N in the diet (pL vs PL) did not affect BW gain, EBW gain or muscle growth, but weight gain in organs + blood was reduced (P<.05) and gain in adipose tissues (P<.10) was increased. Lysine addition to the lysine deficient diet (pL vs pl) resulted in an increase of BW, EBW, carcass and muscles gains, and a reduction of body fatness. EBW gain:BW gain ratio was not affected (P>.10) by dietary treatments and averaged 95.8 (-+ 1.0%). The efficiency of utilization of ME for weight gain was lower (P<.01) for the pl diet than the pL and PL diets. Reduction of nonessential N level resulted in a better efficiency of protein conversion to weight or muscle gain (P<.01).

in the different compartment gains were obtained as the differences between the values measured on experimental animals at the end of the experiment and those predicted at the beginning of the experiment. Two feces and urine collections were carried out on each animal during the respiration chamber trials. Digestible energy and ME content of each diet over the 7-wk experimental period were calculated for each pig as the mean of both values thus obtained. Heat production (comparative slaughter method) was then computed as the difference between ME intake and retained energy in the empty body. Results are expressed either as kcal/d or k c a l . d - t . k g W-'Ts . Metabolic body weight corresponded to the mean of weekly metabolic body weights. Statistical Analysis. Within each replicate, there were differences in ME intake between animals due to feed spillage or refusals and to the slightly higher ME content of PL diet. The data were therefore adjusted and analyzed by covariance with ME intake (kcal/d or kcal-d - 1 -kg W-'Ts ) as a covariate. Main effects considered in the analysis were dietary treatments and replicates. The effect of stage of growth (i.e., week number) was tested in the balance technique data. Newman-Keuls' multiple range test was used to partition treatment means.

R esu Its

Apparent digestibility of energy and

TABLE 2. DIGESTIBILITY AND NUTRITIONAL CHARACTERISTICSOF THE DIETS Diet Item Apparent digestibility (%)b Energy Nitrogen Nutritional characteristics b Apparent DE, kcal/kg Apparent ME, kcal/kg ME/DE ( 100) Apparent digestible Protein, g/Meal DE Lysine, g/Mcal DE pl pL PL SDa

88.5 83.2 c 3,395 c 3,277 96.5 c 37.6 1.98

88.4 82.7 c 3,399 c 3,283 96.6 c 37.5 2.36

89.2 86.Od 3,446 d 3,305 95.9 d 44.8 2.35

1.1 1.8 57 57 .5

astandard deviation. bAs measured: 12 values per treatment. C'dMeans on the same line with superscripts that do not have a common letter differ (P<.05).

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The a m o u n t of lysine per kg muscle gain was n o t affected by dietary t r e a t m e n t s and ranged f r o m 37 to 39 g. At similar daily ME intakes, daily retention of dry matter and ash, as d e t e r m i n e d by the comparative slaughter technique, were n o t affected (P>.10) by dietary t r e a t m e n t (table 4). A d d i t i o n of lysine to the pl diet resulted in an increase in protein deposition ( P < . 0 1 ) , and a r e d u c t i o n in fat deposition (P<.05). Similarly, an increase in nonessential N level (PL vs pL) was associated with e n h a n c e d protein deposition (P<.10), a lower (P<.05) N retention coefficient (table 5) and a t e n d e n c y for a lower fat deposition. However, the daily a m o u n t s of protein retained in muscular tissues and in the carcass were comparable in pL and PL t r e a t m e n t s (table 5). A higher p r o p o r t i o n of retained protein was t h e r e f o r e located in the non-carcass c o m p a r t m e n t in pigs given the PL diet. Dry matter, fat and energy c o n t e n t of EBW or carcass gain were higher (P<.05) and protein c o n t e n t was lower in the pl t r e a t m e n t , in comparison with the pL and PL t r e a t m e n t s

(table 4). No differences (P>.10) b e t w e e n pL and PL t r e a t m e n t s were observed for chemical c o m p o s i t i o n of EBW or carcass gain (table 4). Pigs fed the PL diet tended to retain less energy and p r o d u c e m o r e heat than those given the pL diet (at similar ME intakes: kcal/d or kcal.d-l-kg W - ' T s ) . Energy r e t e n t i o n and h e a t p r o d u c t i o n (kcal/d) were similar in pl and pL t r e a t m e n t s (table 5). However, when the data were expressed per unit o f m e t a b o l i c size, heat p r o d u c t i o n was higher in the pl t r e a t m e n t and c o m p a r a b l e to the value o b t a i n e d with the PL diet. Differences in nitrogen balance and heat p r o d u c t i o n b e t w e e n dietary treatments, as measured by balance procedure and indirect calorimetry, respectively, were c o m p a r a b l e to those o b t a i n e d with the comparative slaughter technique. No interaction b e t w e e n week of e x p e r i m e n t and dietary t r e a t m e n t was observed. Therefore, data in table 6 represent p o o l e d values of the different t r e a t m e n t s and e x t r a p o l a t e d to t h e 7-wk period (see table 6). However, at similar ME intakes

TABLE 3. EFFECT OF PROTEIN AND LYS1NE LEVEL 1N THE DIET ON GROWTH PERFORMANCE, TISSUE GAIN AND FEED EFFICIENCY IN GROWING PIGS a Diet Item Daily intake ME, kcal Crude protein, Lysine, g Daily gain (g) BW EBW Carcass c Organs d + blood Muscle Adipose tissues Efficiency of feed conversion kcal ME/kg EBW gain g protein/kg EBW gain g protein/kg muscle gain g lyslne/kg EBW gain g lysine/kg muscle gain pl pL PL sDb

5,264 246 e 10.8 e 649 e 625 e 485 e 83 e 294 e 132 e 8,391 e 393 e 836 e 17.3 e 36.7

5,264 246 e 12.8 f 699 f 675 f 527 f 85 e 337 f 124 ef 7,765 f 364 f 727 f 19.0 f 37.9

5,264 285 f 12.9 f 700 f 680 f 523 f 93 f 338 f 118 f 7,781 f 420g 855 e 19.1 f 38.8

4 .2 29 21 15 6 14 7 274 15 45 .6 2.1

aBody weight (BW) at the beginning of the experiment was 19.5 -+ .5 kg; the duration of the experiment was 7 wk. bstandard deviation. CEmpty body weight - (blood + organs + head + feet + bristles), (after chilling). dorgans = stomach + intestine + bladder + heart + lungs + liver + spleen + kidneys. e'f'gMeans on the same line with superscripts that do not have a common letter differ (P<.05).

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NOBLET ET AL. TABLE 4. EFFECT OF DIETARY PROTEIN AND LYSINE LEVELS ON NUTRIENT BALANCE AND CHEMICAL COMPOSITION OF GAIN IN GROWINGPIGS (COMPARATIVE SLAUGHTER TECHNIQUE) Diet

Item Daily retention Dry matter, g Protein, g Ash, g Lipid, g Energy, kcal Empty body gain, composition Dry matter, % Protein, % Ash, % Lipid, % Energy, kcal/kg Carcass gain, composition b Dry matter, % Protein, % Ash, % Lipid, % Energy, kcal/kg astandard deviation. bchilled carcass.

pl

pL

PL

SDa

297.3 90.9 c 17.6 186.8 c 2,254 47.40 c 14.52 c 2.81 29.72 c 3,589 e 51.55 c 14.72c 2.83 34.18 c 4,046 c

307.4 103.9 d 20.3 175.3 d 2,257 44.75 d 15.50 d 3.04 25.65 d 3,315cd 48.68 d 16.24 d 2.90 29.20 d 3,693 d

300.9 110.1 e 20.0 167.7 d 2,191 44.59 d 16.20 d 2.90 25.12 d 3,267 d 49.19 d 16.34 d 2.81 29.40 d 3,710 d

4.9 4.7 2.3 9.0 75 1.92 .62 .34 2.30 218 1.90 .78 .20 2.50 21

c'd'eMeans on the same line with superscripts that do not have a common letter differ (P<.05).

(kcal.kg w - ' T S . d - 1 ) , heat production was lower than measured by the indirect calorimetry technique, the difference between both methods representing 6.4% of the value obtained with the comparative slaughter technique. N retention was lower when measured by the comparative slaughter technique as compared with the balance technique (table 6). The difference in N retention amounted to 12.1% of the N retention measured by the comparative slaughter method.

Discussion
Most research has shown that a reduction of the level of protein and essential amino acids in the diet, from an optimum level for growth, is associated with a decreased growth rate and efficiency of feed utilization and a concomitant increase in body fatness (Wahlstrom and Libal, 1974; Noblet and Henry, 1977; Russell et al., 1983). However, our data indicate that the gross efficiency of ME for energy gain (retained energy: ME intake) is higher with such diets (table 5). This is due to the smaller part of ME

used for maintenance requirements (lower body weight) and the reduced amount of protein deposited with low protein diets. On the other hand, a reduction of the level of non-essential N has little effect on growth rate or feed efficiency and tends to increase body fatness (Sharda et al., 1976; Bereskin et al., 1976; Noblet et al., 1980; Stahly et al., 1981; Asche et al., 1985; present experiment). Under the conditions of this study, fat deposition, energy retention, and gross efficiency of ME for energy gain were increased and heat production reduced, with the reduction of nonessential N (tables 4 and 5). From N balance data, it can be calculated that pigs fed the PL diet excreted in the urine about 5.8 g more N per day than those fed the pL diet. After relating the a m o u n t of additional protein catabolized for energy purposes to the increased heat production observed in pigs fed the PL diet, it appears that each gram of protein deaminated produced, in comparison with carbohydrates, an additional 1.8 kcal of heat. This is comparable to the value (1.6 kcal) reported by Just (1982) for growing pigs. The

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values o b t a i n e d in m a t u r e rats or pigs (Hoffm a n n a n d S c h i e m a n n , 1 9 7 1 ) or c a l c u l a t e d f r o m b i o c h e m i c a l m o d e l s (Schulz, 1 9 7 5 ) , c o n f i r m t h a t t h e e f f i c i e n c y o f u t i l i z a t i o n o f ME f r o m p r o t e i n for m a i n t e n a n c e or f a t d e p o s i t i o n is lower than from carbohydrates. However, the a d d i t i o n a l h e a t loss varies b e t w e e n .6 a n d .9 kcal/g. T h e d i f f e r e n c e b e t w e e n b o t h sets o f d a t a suggests t h a t t h e m e t a b o l i s m o f a n i m a l s is also a f f e c t e d b y t h e level o f n o n - e s s e n t i a l N in t h e diet. This h y p o t h e s i s is c o n s i s t e n t w i t h t h e h i g h e r p r o t e i n t u r n o v e r associated w i t h an increased p r o t e i n s u p p l y ( R e e d s e t al., 1981). In a d d i t i o n , t h e mass o f visceral organs a n d b l o o d seems t o b e h i g h e r in pigs f e d a high p r o t e i n diet ( S t a h l y e t al., 1 9 7 9 ; p r e s e n t results). C o n s i d e r i n g t h a t h e a t loss associated with maintenance requirements is highly c o r r e l a t e d w i t h t h e i m p o r t a n c e o f such m e t a -

bolically active tissues ( K o o n g et al., 1 9 8 2 ; Tess et al., 1 9 8 4 ) , pigs fed t h e PL diet w o u l d t h e n exhibit a higher heat production related to maintenance requirements. O u r d a t a s h o w t h a t in o r d e r t o achieve similar ME i n t a k e s , pigs f e d t h e PL diet m u s t b e given 4 1 kcal m o r e D E p e r d a y t h a n pigs f e d t h e p L diet. This d i f f e r e n c e c o r r e s p o n d s t o t h e e n e r g y c o n t e n t o f a d d i t i o n a l loss o f a m m o n i a in u r i n e (i.e., e q u i v a l e n t t o 36 g o f p r o t e i n ) . In o t h e r words, e n e r g y loss in u r i n e was i n c r e a s e d b y a b o u t 1.1 kcal f o r each a d d i t i o n a l gram o f p r o t e i n d e a m i n a t e d . J u s t ( 1 9 8 2 ) a n d J. M. Perez ( p e r s o n a l c o m m u n i c a t i o n ) f o u n d similar values in g r o w i n g pigs. D e g r a d a t i o n o f digestible p r o t e i n in excess o f r e q u i r e m e n t s f o r e n e r g y p u r p o s e s results, t h e r e f o r e , in a n i n c r e a s e d e n e r g y loss in u r i n e a n d a n e l e v a t e d h e a t loss a s s o c i a t e d w i t h

TABLE 5. EFFECT OF DIETARY PROTEIN AND LYSINE LEVEL ON NITROGEN AND ENERGY BALANCE IN GROWING PIGS (COMPARATIVE SLAUGHTER TECHNIQUE) Diet Item Nitrogen balance Lysine intake, g/d Nitrogen intake, g/d Nitrogen retained, g/d % of N digested % of N intake Nitrogen retained In carcass, g/d In muscles, grid Energy balance (kcal/d) DE intake ME intake Retained energy % of DE % of ME As protein b As fat c Heat production Mean BW"~s Energy balance (kcalokg W-'Ts -d - l ) DE intake ME intake Retained energy As protein As fat Heat production astandard deviation. bCalculated as protein gain X 5.7. CCalculated as retained energy - energy retained as protein. d'e'fMeans on the same line with superscripts that do not have a common letter differ (P<.05). pl pL PL SD a

10.8 d 39.4 d 14.5 d 44.4 d 36.9 d 11.4 d 8.2 d 5,456 d 5,264 2,254 41.3 42.8 518 d 1,736 d 3,010 13.96 d 383.4 d 369.8 156.1 37.5 d 118.5 d 213.8

12.8 e 39.4 d 16.7 e 52.0 e 43.1 e 13.6 e 10.1 e 5,450 d 5,264 2,257 41.5 42.9 592 e 1,665 de 3,007 14.34 e 383.0 d 369.8 159.5 41.1 e 118.3 d 210.4

12.9 e 45.6 e 17.8 f 44.8 d 38.6 e 13.7 e 10.2 e 5,491 e 5,264 2,190 39.9 41.7 633 f 1.557 e 3,074 14.40 e 385.8 e 369.8 155.7 43.6 f 112.1 e 214.2

.2 .6 .7 3.2 2.6 .6 .5 16 75 1.4 27 93 75 .16 1.1 5.1 1.7 6.2 5.1

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NOBLET ET AL. TABLE 6. COMPARISON OF ENERGY BALANCE DATA OBTAINED FROM RESPIRATORY CALORIMETRY AND COMPARATIVE SLAUGHTER TECHNIQUES Indirect calorimetrya
15.2

Item Mean BW "Ts, kg

Comparative slaughter technique b 14.3 374.8 158.1 40.9 117.2 216.7 42.0 16.5 25.5

ME intake, kcal.kg W-'Ts -d -1 Retained energy, kcal,kg W-'~s .d -1 As protein As fat Heat production, kcal.kg W-'Ts .d -1 N intake, g/d Nrctained, g/d N losses, g/d

375.3 (374.8) c 174.3 (171.9) c 46.6

127.7

201.0 44.7 19.6 25.1

(202.9) c (42.0) d (18.5) d (23.5) d

aFrom wk 1 to 6; 36 measurements on 18 animals (two per pig). bFrom wk 0 to 6; 18 measurements on 18 animals. CExtrapolated to 7 wk of experiment (+ 4 kcal/kg W"~s per wk of experiment) and 374.8 kcal ME intake (.3 kcal heat production per additional kcal ME). (J. Noblet and C. Karege, unpublished data). dcorrected to 42.0 g/d N intake and assuming that the ratio N retained:N intake is 40% (table 5).

utilization of ME and changes in metabolism. To obtain a similar energy retention in the pL and PL treatments, pigs fed the PL diet would require a daily supplement of 97 and 140 kcal ME and DE, respectively (assuming that the efficiency of ME utilization for energy deposition is 70% in both treatments and that the ME:DE ratio is equivalent to the values presented in table 2). These values represent 1.9 and 2.6% additional ME and DE, respectively, or about .8 and 1.0% more ME and DE, respectively per one percentage increase in protein level. However, the energy sparing effect due to reduced nonessential N level in the diet increases fat deposition (present experiment). Therefore, it will produce small effects on growth rate (about 11 g of fat between treatments pL and PL, table 5). Consequently, even if the reduction of protein level in the diet is associated with a better efficiency of utilization of ME as shown by Just (1982) and as found in our experiment, it does not significantly affect feed efficiency (Russell et al., 1983; present results). A decreased level of nonessential N in the d i e t does not significantly affect growth rate or f e e d efficiency, but the amount of nitrogen retained in the b o d y is lower (Sharda et al., i 9 7 6 ; Russell et al., 1983; present experiment). In fact, muscle growth and N retained in muscles seem to be independent of the level of nonessential N per se. Therefore, N retained in

the non-carcass compartment is lower with low protein diets (table 5). Our results also show that when lysine is the limiting factor in the diet, the amount of lysine per unit of weight gain (19 g/kg), muscle gain (38 g/kg) or protein retained (120 g/kg) is constant and independent of protein and lysine levels in the diet. Similar conclusions were given by Stahly et al. (1979, 1981) and Asche et al. (1985). Lysine content of e m p t y body protein was similar in the three treatments and averaged 7.3%, which is close to values reported in literature (Wiesemuller and Poppe, 1974). Therefore, it can be calculated that lysine retained in the e m p t y body represents about 60% of the lysine intake and 70% of the ileal lysine (85% availability; Laplace et al., 1985) in corn-soybean diets over the growing phase (20 to 55 kg b o d y weight). Finally, free lysine appears to be used as efficiently for growth and protein deposition as the lysine contained in protein. Nevertheless, as shown by Batterham and O'Neil (1978) and Partridge et al. (1985), there is evidence that the efficiency of utilization of free lysine is poorer when feed is given once daily compared with several meals. In agreement with other data (Just et al., 1982), heat production as measured by the indirect calorimetry method is lower than by the comparative slaughter technique. In spite of probable systematic errors in the indirect calorimetry method (Just et al., 1982), one

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EFFECT OF DIETARY PROTEIN AND LYSINE ON PIGS

725

r e a s o n f o r t h e d i s c r e p a n c y w o u l d be t h e l o w e r level o f a c t i v i t y o f t h e pigs w h e n in t h e respirat i o n c h a m b e r in cages, as c o m p a r e d w i t h o u t s i d e c o n d i t i o n s in flat-decks ( p e r s o n a l o b s e r v a t i o n ) . Since h e a t p r o d u c t i o n d i r e c t l y associated w i t h a n o r m a l level o f a c t i v i t y m a y r e p r e s e n t 15% o f fasting h e a t p r o d u c t i o n ( N i e n a b e r et al., 1 9 8 5 ) , a large p r o p o r t i o n o f t h e d i f f e r e n c e in h e a t p r o d u c t i o n o b s e r v e d between both methods could be accounted for b y d i f f e r e n c e s in level o f activity. In a d d i t i o n , a m b i e n t t e m p e r a t u r e was c o n s t a n t a n d p r o b ably slightly h i g h e r w i t h i n t h e r e s p i r a t i o n c h a m b e r (22 C vs 18 t o 22 C). T h e r e f o r e , pigs k e p t in a flat-deck e n v i r o n m e n t were a t o r b e l o w t h e i r l o w e r critical t e m p e r a t u r e (Close a n d M o u n t , 1978), w i t h a s u b s e q u e n t i n c r e a s e d h e a t p r o d u c t i o n . As r e p o r t e d b y J u s t et al. ( 1 9 8 2 ) , e s t i m a t e s o f N r e t e n t i o n are l o w e r when measured by the comparative slaughter t e c h n i q u e , as c o m p a r e d w i t h t h e b a l a n c e t e c h n i q u e ( t a b l e 6). T h e m a i n e x p l a n a t i o n f o r t h e d i s c r e p a n c y is t h a t N losses (feces, u r i n e ) are u s u a l l y u n d e r - e s t i m a t e d , w i t h a s u b s e q u e n t over-estimation of N retention. The difference f o u n d in t h e p r e s e n t e x p e r i m e n t (12.1%) is c o m p a r a b l e t o t h e m e a n l i t e r a t u r e value (16%) given b y J u s t e t al. ( 1 9 8 2 ) .

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Citations

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