Specific Effects of Lysine on Broiler Production: Comparison with Threonine and Valine

BERNARD LECLERCQ INRA, Station de Recherches Avicoles, 37380 Nouzilly, France ABSTRACT Lysine exerts specific effects on body composition at dietary levels higher than that required for maximum growth rate, which also results in an improved feed conversion ratio (FCR). As a consequence of these differences, it becomes apparent that there is a hierarchy of requirements. The requirement for maximum gain is lower than that for breast meat (BM) yield, which is lower than the requirement for FCR; and lastly, the requirement for minimum abdominal fat (AF) percentage is the highest of all. Threonine, on the other hand, does not appear to exhibit so pronounced an effect on body composition, whereas, in the case of valine, these effects seem to be absent. These different effects of lysine, threonine, and valine can induce different amino acid requirement profiles according to the criteria used for determining the requirement. Moreover the particular mathematical model being used to calculate amino acid requirements can also influence amino acid requirement profiles.

(Key words: lysine, broiler, growth, body composition, feed conversion ratio) 1998 Poultry Science 77:118123

INTRODUCTION
Lysine and the sulfur amino acids (SAA), are known to exhibit specific effects on carcass composition. This accounts for the higher requirement when breast muscle (BM) yield is used as the basis for setting the lysine requirement as compared to weight gain (Jensen et al., 1980; Hickling et al., 1990; Moran and Bilgili, 1990; Schutte and Pack, 1995). However experiments performed to investigate this issue have not always been precise, as the number of lysine levels are often limited and do not allow accurate estimation of amino acid requirements. This paper deals with some recent results of lysine effects and lysine requirements in broiler chicken production. Moreover, because threonine and valine were studied simultaneously with lysine in one experiment, comparison between these amino acids can provide further information on the specific effects of each amino acid and the relative ratios between threonine or valine and lysine to be adopted in feed formulation.

EFFECT OF LYSINE ON BODY COMPOSITION

Several authors have recently shown that lysine supplementation, at levels above the requirement for

Received for publication March 18, 1997. Accepted for publication August 5, 1997.

maximal growth rate, results in specific and significant effects on body composition. That is the case for breast meat (BM) yield (Hickling et al., 1990; Moran and Bilgili, 1990). Similarly lysine can modify fat deposition, this was very well demonstrated by Grisoni (1991) by using eight levels of lysine supplementation. It can be seen in Table 1 that lysine supplementation reduced abdominal fat (AF) percentage. From this experiment it can be concluded, using a monomolecular model, that the total lysine requirement (0.99 asymptote) for gain, feed conversion ratio (FCR), and AF were, respectively, 9.10, 10.07, and 14.3 g/kg. Unfortunately BM yield was not measured in this experiment. Using seven levels of lysine supplementation Han and Baker (1994) studied the effect of lysine on gain, FCR, BM yield, and AF percentage between 3 to 6 wk of age. By recalculating the digestible lysine requirement from their data on male boilers and using the monomolecular model, requirements for reaching the 0.99 asymptote were 10.14 g/kg for gain and 10.6 g/kg for AF percentage. More recently, Tesseraud et al. (1996) reinvestigated the effects of lysine level on the growth of different muscles and protein renewal. It is clearly shown in Table 2 that a lysine deficiency specifically reduces Pectoralis major growth; these muscles having Type IIb fibers. In contrast, proportions of Sartorius and Anterior latissimus dorsi, which contain both Type I and IIb fibers, were less influenced by lysine (10.1 g/kg vs 7.7 g/kg); this observation was made at 2, 3, and 4 wk of age. Renewal of muscle proteins was studied in the same experiment (Table 3). Lysine deficiency increased 118

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TABLE 1. Effects of lysine supplementation on growth performances and abdominal fat in 49-d-old broilers1 Total lysine content (g/kg) 7 8 9 10 11 12 13 14
1Data

119

TABLE 2. Effect of lysine deficiency on different muscle weights (lysine-deficient diet/control diet)1 Variable Body weight Pectoralis major Sartorius Anterior longissimus dorsi
1Data

Daily gain 53.1 56.6 58.8 59.4 58.7 59.0 58.8 58.8

Feed conversion ratio (21 to 49 d) 2.28 2.15 2.10 2.03 2.04 2.05 2.05 2.04

Abdominal fat (g/kg) 39.7 34.1 31.3 29.0 26.3 26.8 26.7 27.1

2 wk 0.50 0.37 0.72 0.59

3 wk 0.50 0.34 0.61 0.56

4 wk 0.56 0.34 0.62 0.56

from Tesseraud et al. (1996).

from M. L. Grisoni (1991).

both relative protein synthesis and degradation (percentage per day) in the three muscles studied. However, the P. major exhibited the highest increase in rate of renewal. Pectoralis major, therefore, seems to be more sensitive to lysine intake than other muscles. It is still not clear whether this is due to the type of fibers (IIb) or to the late development of these muscles (Acar et al., 1993). The specific effect of lysine intake on body composition was recently reinvestigated in a large collaborative research program, involving TNO-ILOB-Holland (Dr. Schutte), the Rijksstation voor Kleinveeteelt-Belgium (Dr. De Groote), INRA-Station de Recherches Avicoles (Dr. Leclercq), and Degussa and Eurolysine companies. This study dealt with requirements of male broiler chickens for lysine, SAA, threonine, arginine, valine, isoleucine, and tryptophan during the finishing period (20 to 40 d of age). Details of this experiment are still to be published. This paper will give a summary of the lysine, threonine, and valine results observed in that study. Comparison between these three amino acids is particularly powerful because six levels of each amino acid were compared and all three amino acids were studied in the same experiment (same time, same poultry house, same genotype of broiler, same basal diet, same dissection procedure). The lysine responses are presented in Table 4. The lowest level of lysine was the only treatment that significantly reduced growth rate. However BM yield and AF were influenced by higher concentrations of lysine. The effects of threonine on growth and body composition are given in Table 5. Again, the lowest level of threonine was the only treatment exhibiting a significant effect on body mass gain. Breast meat yield was reduced only at the lowest level of threonine. Abdominal fat was not significantly modified by threonine supplementation, even though there was a trend for AF to decrease as the dietary threonine increased. Moreover, the effect of a threonine deficiency on BM yield was less pronounced than that of a lysine deficiency. Table 6 shows that a valine deficiency was reached at the two lowest valine levels. In contrast to the effect of

lysine and threonine, a valine deficiency in this range, did not influence BM yield or AF percentage. This specific effect of lysine, as compared to threonine and valine, can be confirmed if we compare linear regression lines between BM yield or AF and live weight at 40 d of age (slaughter age) for birds fed the lowest level of each amino acid. As is evident from Table 7, if we compare broiler chickens from the treatments containing the lowest levels of lysine, threonine, or valine, mean live weights (LW) were very similar, which shows that the degree of deficiency were similar. However BM yield and AF percentage were different. Covariance analysis shows significant differences between the three amino acids for the BM to LW regression. In the case of the AF to LW regression, the regression line for lysine was higher than those of the other two amino acids. These results clearly demonstrate that amino acid deficiencies that result in a similar reduction of growth rate affect body composition differently. It is also clear that lysine exerts an apparent partitioning effect in the range around the requirement for daily gain.

EFFECT OF LYSINE ON FEED CONVERSION

As the lysine deficiency decreases growth rate, FCR increases. However there seems to be an additional effect that is not accounted for the effect on gain. It is

TABLE 3. Effects of lysine deficiency on turnover constants of proteins from different muscles Control diet (10.1 g/kg lysine) Lysine-deficient diet (7.7 g/kg lysine) (%/d) Pectoralis Synthesis Degradation Sartorius Synthesis Degradation Anterior longissimus dorsi Synthesis Degradation
1Data

Protein

18.3 8.2 14.7 6.2 26.3 18.6

29.7 20.5 17.3 10.0 32.1 26.7

from Tesseraud et al. (1996).

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LECLERCQ
TABLE 4. Effect of lysine supplementation (20 to 40 d of age) on growth performance and body composition of male broilers True digestible lysine (g/kg) 8.0 8.9 9.8 10.7 11.6 12.5
adMeans

Total lysine

Daily gain (g/d) 69.2a 75.1b 77.1b 77.0b 77.8b 77.5b

Feed conversion ratio (g:g) 1.982a 1.846b 1.755cd 1.796c 1.719cd 1.710d

Breast muscle 148.7a 159.5b 169.0c 171.3c 168.1c 170.3c

Abdominal fat (g/kg) 25.4c 24.3bc 22.4ab 20.4a 20.4a 19.4a

7.28 8.18 9.08 9.98 10.88 11.78

in a column with no common superscript differ significantly (P 0.05). TABLE 5. Effect of threonine deficiency (20 to 40 d of age) on growth performances and body composition of male broilers True digestible threonine Feed conversion ratio (g:g) 1.939b 1.764a 1.730a 1.731a 1.724a 1.730a

Total threonine (g/kg) 5.50 6.10 6.70 7.30 7.90 8.50

a,bMeans

Daily gain (g/d) 69.8a 77.9b 78.1b 78.6b 78.3b 77.1b

Breast muscle 160.5a 169.8b 170.5b 169.9b 169.8b 170.0b

Abdominal fat (g/kg) 22.9 21.1 21.5 20.3 20.7 19.8

4.79 5.39 5.99 6.59 7.19 7.79

in a column with no common superscript differ significantly (P 0.05).

TABLE 6. Effects of valine deficiency (20 to 40 d of age) on growth performances and body composition of male broilers True digestible valine (g/kg) 6.80 7.34 7.88 8.42 8.96 9.50
acMeans

Total valine

Daily gain (g/d) 67.2a 73.8b 77.6c 77.8c 77.2c 78.1c

Feed conversion ratio (g:g) 1.937a 1.770b 1.754b 1.720b 1.719b 1.710b

Breast muscle (g/kg) 167.8 169.8 172.1 172.7 170.1 170.9

Abdominal fat 21.7 21.2 21.7 21.1 21.2 21.8

6.12 6.66 7.20 7.74 8.28 8.82

in column with no common superscript differ significantly (P 0.05).

TABLE 7. Comparison of linear regression equations between breast muscle (BM) or abdominal fat (AF) and live weight (LW) from male broilers fed the lowest levels of lysine, threonine, or valine at 40 d of age Mean BM weight 314.2 343.9 354.3

Amino acid Lysine (8 g/kg) Threonine (5.5 g/kg) Valine (6.8 g/kg)

Mean LW 2,212 2,145 2,115

Mean AF weight (g) 54.0 49.2 46.1

Breast muscle vs live weight Lysine BM = 0.1651 LW + 34.4 (r = 0.741) Threonine BM = 0.1378 LW + 48.3 (r = 0.723) Valine BM = 0.1214 LW + 97.6 (r = 0.564) Covariance analysis: lysine < threonine < valine Abdominal fat vs live weight Lysine AF = 0.0491 LW 50.0 (r = 0.646) Threonine AF = 0.0312 LW 17.7 (r = 0.434) Valine AF = 0.0365 LW 31.1 (r = 0.480) Covariance analysis: lysine > threonine valine

LYSINE SYMPOSIUM
TABLE 8. Amino acid requirements using the broken line model Variable Gain mg/g gain g/kg diet Feed conversion ratio mg/g gain g/kg diet Breast muscle proportion mg/g gain g/kg diet Abdominal fat proportion mg/g gain g/kg diet Lysine 16.64 9.24 20.08 10.10 17.02 9.75 21.78 11.88 Threonine 10.75 6.10 10.75 6.10 10.75 6.10 9.54 5.50 Valine 12.90 7.66 12.34 7.34 12.34 7.34 11.40 6.80 Variable Gain mg/g gain g/kg diet Feed conversion ratio mg/g gain g/kg diet Breast muscle proportion mg/g gain g/kg diet Abdominal fat proportion mg/g gain g/kg diet TABLE 9. Amino acid requirements using a monomolecular model1 Lysine 16.71 9.69 18.33 11.84 17.21 10.63 44.46 25.78 Threonine 10.31 5.91 11.08 6.35 10.14 5.81 15.65 8.97

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Valine 14.12 8.14 13.70 7.90 13.91 7.11 11.79 6.80

1y = a b ecx y = performance; a = asymptote; and x = amino acid. Requirements were calculated for the 0.99 asymptote (Gain and BM) or the 1.01 asymptote (FCR and AF).

likely that the effect on body composition leads to a better FCR, because lipid gain is reduced by high levels of lysine. This effect on FCR has been observed by a number of authors. The experiment of Grisoni (1991) shows that the requirement for FCR, using the monomolecular model and the performance of 1.01 asymptote, was 10.07 g total lysine/kg feed, whereas the requirement for gain was 9.10 g/kg. In the experiment from Han and Baker (1994) the requirement for FCR (1.01 asymptote) was 11.55 g digestible lysine/kg of feed, which was very close to the requirement for gain (10.14 g/kg). In the collaborative work presented in Table 4, the requirement for FCR was always higher than that for gain, whatever the mathematical model used. The broken line model, see Table 8, resulted in 9.24 and 10.10 g total lysine/kg feed, as requirements for gain and FCR respectively. Table 9 gives the results of the monomolecular model. Requirements needed to reach the 0.99 asymptote were 9.69 and 11.84 total lysine/kg feed for gain and FCR, respectively. It appears, therefore, that the lysine requirement for FCR is higher than the requirement for gain. Threonine and valine, when compared to lysine in the same trial, did not exhibit the same effects. Using the broken line model, requirements for gain and FCR were similar (6.10 g total threonine/kg feed). Using the monomolecular model requirements were 5.91 and 6.35 g total threonine/kg respectively. In the case of valine, the broken line model led to requirements of 7.66 and 7.44 g total valine/kg feed for gain and FCR, respec-

tively, although the monomolecular model gave requirements (at 0.99 asymptote) of 8.14 and 7.90 g total valine/ kg. Based on these results, the valine requirement for FCR was lower than the requirement for gain.

CONSEQUENCES FOR AMINO ACID REQUIREMENTS

These studies indicate that requirements for a given amino acid depend on the criteria being measured and the mathematical model applied to estimate them. We have observed that in most of the experiments the lysine requirement for gain is lower than the requirements for FCR, BM percentage, or AF percentage; the lysine requirement for lowest AF percentage being the highest. In the Grisoni experiment, the requirement for reaching the 1.01 asymptote of AF percentage was 14.32 g/kg, 9.10 for gain. In the experiment of Han and Baker (1994), it was not easy to calculate the lysine requirement for lowest AF percentage, because the asymptote was near 0 and the requirement higher than 100 g/kg, which is unlikely. In our recent collaborative work (Table 6 and 7), the lysine requirement for the lowest AF percentage was estimated as 11.88 g/kg (compared to 9.24 for gain) using the broken line model, and 25.78 g/kg (instead of 9.69 for gain) using the monomolecular model; this last value is also unlikely. If we consider BM yield, the lysine requirement generally falls between the requirement for gain and the

TABLE 10. Calculated ratios between threonine or valine and lysine according to the criterion1 and the mathematical model (calculations based on dietary amino acid contents) Broken line model Amino acid Lysine Threonine Valine
1FCR

Monomolecular model AF% 100 46.3 57.2 Gain 100 61.0 84.0 FCR 100 53.6 66.7 BM% 100 54.7 66.9 AF% 100 34.8 26.4

Gain 100 66.0 82.9

FCR 100 60.4 72.7

BM% 100 62.6 75.3

= feed conversion ratio; BM% = breast meat percentage; AF% = abdominal fat percentage.

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TABLE 11. Calculated ratios between threonine or valine and lysine according to the criterion1 and the mathematical model2 Broken line model Amino acid Lysine Threonine Valine
1FCR 2Calculations

Monomolecular model AF% 100 43.8 52.3 Gain 100 61.7 84.5 FCR 100 60.5 74.7 BM% 100 58.9 80.8 AF% 100 35.2 26.5

Gain 100 64.6 77.5

FCR 100 53.5 61.5

BM% 100 63.2 72.5

= feed conversion ratio; BM% = breast meat percentage; AF% = abdominal fat percentage. based on dietary amino acid intakes.

requirement for FCR. In the experiment of Han and Baker (1994) it was not possible to estimate this requirement because some of the data were erratic which resulted an unlikely equation. However, if we discard the higher lysine level, it seems that the requirement for BM yield is similar to the requirement for gain. In our recent experiment, the requirements for BM yield were 9.75 and 10.63 g/kg, respectively, for the broken line model and the monomolecular model (respective values for gain: 9.24 and 9.69). Therefore, it appears that the lysine requirement for BM yield is slightly higher than the requirement for gain, but lower than the requirements for FCR and AF percentage. Different conclusions must be drawn for threonine and valine. Using the broken line model to estimate the threonine requirements, it was found that the requirements for gain, FCR, and BM yield were similar. The requirement for the lowest AF percentage was lower than for LW gain. This is probably due to the variability of lipid deposition, which makes it difficult to measure very small differences between treatments. When the monomolecular model is applied, the requirement for BM yield is slightly lower than that for gain. The requirement for the lowest FCR is higher, and the requirement for the lowest AF percentage is higher still. These conclusions agree closely with those of RangelLugo et al. (1994), who suggested that threonine requirement for FCR is slightly higher than the requirement for maximum gain. In the case of valine, the broken line model gave the lowest requirement for AF percentage, then FCR and BM yield. The highest requirement was seen for LW gain. Similar conclusions were observed using the monomolecular model. It, therefore, appears that the hierarchy of requirements for the different criteria depends on the amino acids involved. These results have direct consequences on the relationship of other amino acids to lysine. If we consider only threonine and valine, the relative ratios between them and lysine depend on the criteria and the mathematical model being used. Table 10 shows that the relative ratio between threonine and lysine, or valine and lysine are lower if the monomolecular model is used than if the broken line model is used. These results show the direct consequences of the different effects of lysine, threonine, and valine on body composition and FCR; however, in all cases, the threonine to lysine ratio

was lower than that proposed by Baker (1994), whereas the valine to lysine ratio tended to be higher. Similar conclusions can be drawn from calculations based on amino acid requirements expressed as milligrams per gram of gain, rather than grams per kilogram of feed. Expressing amino acid requirements on gram per unit feed mass basis is inaccurate, as feed consumption is dependent on ME content of the feed, environmental temperatures, and genotype of broiler. Expressing amino acid requirements as unit intake per unit gain (milligrams per gram of gain), for a given age, minimizes the error due to these factors. If we use this method for expressing requirements, relative ratios of threonine to lysine or valine to lysine change slightly. Table 11 illustrates the relative ratios for the different growth parameters and the mathematical model used. The question is how to select the correct lysine intake and how to express the requirement for other amino acids in reference to lysine. This can be determined by calculating an economic response. Using our own data, presented in Table 4, and the monomolecular model, it is possible to combine several curvilinear equations in order to minimize the feeding costs of either 1 kg of LW gain or BM yield. The results of this exercise are presented in Table 12. Using our data and prices given in this table, we observed that the economical requirement for gain was 10.87 g total lysine/kg diet, which is very close to the requirement at the 1.01 asymptote of FCR (11.84 g/kg). The requirement for BM yield was

TABLE 12. Lysine content required to reach the minimum feeding cost per kilogram of gain or per kilogram of breast muscles using curvilinear responses1 Hypothesis Gain = 1,552 12.34 106 e14.02 lysine FCR = 1.707 + 85.99 e7.195 lysine BM% = 17.09 5,850 e9.824 lysine Where lysine is the total lysine content (percentage) Price of the unsupplemented diet: 0.2 $/kg Lysine content of the unsupplemented diet: 8 g/kg Price of lysine: 0.003 $/g Economical requirements to minimize feeding cost per kilogram of gain (20 to 40 d) 10.87 g total lysine/kg diet to minimize feeding cost of 1 kg of breast muscles 11.42 g total lysine/kg diet
1Monomolecular

model (20 to 40 d).

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higher (11.42 g/kg). From this exercise, it is clear that the economic requirement is much higher than the requirement for gain calculated using the broken line model, which underestimates lysine requirement. As for the relative ratios between threonine and lysine or valine and lysine, it seems more accurate to use those values calculated from the broken line model. Indeed the ratios obtained from the curvilinear model are greatly influenced by the shape of response curve of both lysine and the other amino acids. Furthermore, it seems arbitrary to choose 0.99 asymptote as the level of requirement. Small changes in the position of asymptote of the response curves of some of the growth parameters, may result in large variations in requirement. This situation does not occur with the broken line model.

REFERENCES
Acar, N., E. T. Moran, and D. R. Mulvaney, 1993. Breast muscle development of commercial boilers from hatching to twelve weeks of age. Poultry Sci. 73:317325. Baker, D. H., 1994. Ideal amino acid profile for maximal protein accretion and minimal nitrogen excretion in swine and poultry. Proc. Cornell Nutr. Conf. 134138. Grisoni, M. L., 1991. Role des acides amines alimentaires dans la lipogenese du poulet de chair. Ph.D. thesis. University of Aix-Marseille, France. Han, Y., and D. H. Baker, 1994. Digestible lysine requirement of male and female broiler chicks during the period three to six weeks posthatching. Poultry Sci. 73:17391745. Hickling, D., M. Guenter, and M. E. Jackson, 1990. The effect of dietary methionine and lysine on broiler chicken performance and breast meat yield. Can. J. Anim. Sci. 70: 673678. Jansen, L. S., C. L. Wyatt, and B. I. Fancher, 1989. Sulfur amino acid requirement of broiler chickens from 3 to 6 weeks of age. Poultry Sci. 68:163168. Moran, E. T., and S. F. Bilgili, 1990. Processing losses, carcass quality and meat yields of broiler chickens receiving diets marginally deficient to adequate in lysine prior to marketing. Poultry Sci. 69:702710. Rangel-Lugo, M., C. L. Su, and R. E. Austic, 1994. Threonine requirement and threonine imbalance in broiler chickens, Poultry Sci. 73:670681. Schutte, J. B., and M. Pack, 1995. Effects of sulphur-containing amino acids on performance and breast meat deposition of broiler chicks during the growing and finishing phases. Br. Poult. Sci. 36:747762. Tesseraud, S., N. Maa, R. Peresson, and A. M. Chagneau, 1996. Relative responses of protein turnover in three skeletal muscles to dietary lysine deficiency in chicks. Br. Poult. Sci. 37:641650.

CONCLUSION
From this review, it is apparent that increasing lysine intake above the level required for maximum live weight gain affects body composition by increasing BM yield and decreasing AF percentage. These effects, which can be added to a curvilinear effect on growth rate, induce a beneficial effect on FCR. This leads to increasing the lysine requirement according to the criteria desired: gain < BM < FCR < minimum AF. As threonine and valine do not exhibit such pronounced effects or such a hierarchy of requirements, the amino acid ratios between these amino acids and lysine depend on the desired response.