Information for the Feed Industry

amino acids and more.

Special Issue from Volume 04 / Number 01 / March 2003

The Ideal Protein Concept in broiler nutrition 1. Methodological aspects opportunities and limitations by Dr. Andreas Lemme

Volume 04/ Number 01 March 2003

amino acids and more.

Poultry

The Ideal Protein Concept in broiler nutrition 1. Methodological aspects opportunities and limitations
Key information

The idea of the Ideal Protein Concept

One important progress in animal nutrition in recent years and thus also in broiler nutrition research has been the development of the concept of an "Ideal Protein". One basic idea of this concept is that birds need amino acids in a certain balance to ensure optimum performance. Any absorbed amino acid which is in relative excess compared to the first limiting amino acid will be oxidised and nitrogen will be excreted. Therefore, adjusting the dietary amino acid supply according to the Ideal Protein Concept helps to maximise nitrogen utilisation. One practical application resulting from this is the establishment of low protein diets allowing for the same animal performance if compared to unbalanced high protein diets. A second basic idea of this concept is that while the bird's requirement for essential amino acids may vary between various practical situa-

The Ideal Protein Concept has been an

important progress in animal nutrition because it enables the nutritionist to immediately adjust amino acid supply to changing production conditions. Lysine is the reference amino acid and all remaining essential amino acids are related to it. They can be adjusted to changing lysine requirements by simple calculation. Although different approaches to determine the Ideal Protein pattern have been applied, the published ideal amino acid profiles are fairly consistent. Ideal ratios between single amino acids and lysine might be affected by factors like age, environmental conditions such as climate or stress, and genetic factors such as strain or sex.

tions, the ratios between these amino acids remain fairly stable at the same time. Therefore, the responses of only one single reference amino acid -usually lysine- to changing production conditions have to be evaluated while the remaining amino acids are then to be adjusted simply by calculation. Only essential amino acids are considered in this concept. However, there is still a scientific discussion about the adequate ratio between essential and non-essential amino acids especially in low protein diets.

Lysine is the reference amino acid

In swine nutrition lysine has been established as reference amino acid mainly because lysine is the first-limiting amino acid in swine diets. Although this is not the case in common broiler diets here methionine and cystine (Met+Cys) are first-limiting in most cases lysine

Figure 1: Daily net Lys (left) and Met+Cys (right) requirement for broilers from 7 to 63 days of age, expressed as a total and differentiated between requirements for body protein deposition, feather growth and maintenance

Net Lys requirement (mg/d)

1600 1600

Net Met+Cys requirement (mg/d)

Total Met+Cys requirement Met+Cys for body protein Met+Cys for feather protein Met+Cys for maintenance

1200

1200

800
Total Lys requirement Lys for body protein Lys for feather protein Lys for maintenance

800

400 0 0 7 14 21 28

400

0 35 42 49 56 63 0 7 14 21 28 35 42 49 56 63

Days of age

Days of age

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(Lys) is taken as the reference amino acid as well supported by the following three arguments: Lys is almost exclusively utilised for body protein accretion and thus requirement is only affected very little by other metabolic functions (maintenance requirement) or feathering as this is the case for Met+Cys (Figure 1). There are no metabolic interactions between Lys and other amino acids. In contrast, Met can be converted to Cys by the bird but the opposite way is not possible. From the analytical perspective, it is easier to analyse Lys than Met and particularly Cys.

The factorial approach a mathematical method

The factorial approach is based on the concept that the requirement for an amino acid can be divided into three compartments: requirement for body protein accretion, for feather growth and for maintenance. To obtain the requirement for digestible amino acids, the net requirement for body and feather protein deposition, which can simply be determined by chemical analysis of these body compartments, has to be divided by the utilisation coefficient of the absorbed amino acids. Maintenance requirement is usually defined to be that amount of amino acids needed for body weight equilibrium. Having all information for net amino acid retention in body protein and feathers, utilisation of absorbed amino acids, and maintenance requirement together, overall requirement for each digestible amino acid can then be calculated. In Figure 1, the daily net requirement of Lys (left) and Met+Cys (right) for broilers over a period of eight weeks is shown. The overall requirement for Lys and Met+Cys is not that different but

the contributions of protein accretion, feather growth, and maintenance to total requirement differ markedly between both amino acids. Lys is almost exclusively used for protein accretion, whereas relatively high proportions of Met+Cys are required for feather growth and maintenance. Since the latter is a function of body weight, it increases with age. This underlines the advantage of Lys vs. Met+Cys as the reference amino acid. The overall requirement for digestible Met+Cys and Lys as well as their respective ratios are presented in Figure 2. Coefficients related to the utilisation of the absorbed amino acids are taken from Baker (1991). The value for Lys has been reported to be 80 % while it was found to be 76 % for Met+Cys. These factors are assumed to be constant which has been criticised sometimes because the closer the amount of absorbed amino acids gets to the requirement, the more of them will be oxidised. Thus, linear dose-response relationships between retention and intake of digestible amino acids are rather unlikely especially when considering a population of animals. Furthermore, under

There are various ways to determine an Ideal Amino Acid Profile

Different scientific approaches to obtain ideal amino acid profiles have been described. One method is simply to review literature: The results of all available experiments designed to determine the requirement for individual amino acids are compiled and the average optimum amino acid levels are then presented as ideal amino acid profile. However, due to differences regarding the experimental conditions, e.g. environment, climate, housing system, stocking density, diet composition, a number of possible factors and inaccuracies affecting amino acid ratios are not considered properly. Although not explicitly presented as ideal ratios to Lys, the amino acid recommendations for broilers by NRC (1994) might serve as an example for this method. The factorial approach, multiple dose-response experiments investigating the full range of essential amino acids simultaneously, or the deletion method are more sophisticated methods to obtain ideal amino acid profiles. These shall be introduced in more detail in the following.

Figure 2: Overall requirements of broilers 7 to 63 days of age for digestible Lys and Met+Cys and the respective Met+Cys:Lys ratios

Dig. Lys and Met+Cys requirement (mg/d)

2000

Met+Cys:Lys ratio (%)

100

1600

80

1200

60

800
Lys requirement

40

400

Met+Cys requirement Met+Cys:Lys ratio

20

0 0 7 14 21 28 35 42 49 56 63

Days of age

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practical conditions, amino acid levels are very close or even above the requirement. Not only the amount of digestible Met+Cys and Lys changes with age, but also their ideal ratio (Figure 2). The calculated digestible ideal Met+Cys:Lys ratio increases from 65 % at day 7 to 99 % at day 63, mainly due to the increasing Met+Cys demand for maintenance. It is worth noting that NRC (1994) recommendations suggest quite the opposite trend by stating an ideal Met+Cys:Lys ratio of 82 % in the starter but only 72 % in the grower phase, showing again the limitations of the literature survey method. Changing ideal ratios with age can be expected also for other essential amino acids. Examples for ideal amino acid profiles obtained by the factorial approach are the Illinois Ideal Chicken Protein published by Baker (1994) and GfE (1999) (Table 1). It should be noted that Baker (1994) related the profile to digestible amino acids, whereas GfE (1999) gave ratios on a total amino acid basis. Ideal amino acid ratios should be based on digestible rather than total amino acids because digestibility can interfere with the ideal ratios. This becomes important when raw materials with different amino acid digestibilities are used. The amino acid profile arriving at the respective tissues is crucial, and it can be different from the dietary amino acid composition due to losses during digestion (and oxidation as discussed above).

conducting simultaneous or multiple dose-response studies. Thus, the growth and body composition responses of birds can be used to derive amino acid requirements including utilisation of absorbed amino acids. Moreover, experimental conditions like feed composition, feed ingredients or environmental aspects can be kept as uniform as possible. The only difference between trials should be therefore the dietary levels of the respective amino acid under test. Additionally, other criteria such as feed conversion or breast meat yield can be investigated and assessed at the same time. Such a project, conducted with growing male chicken (20 to 40 days of age), has been realised by Mack et al. (1999). The response of broilers to increasing levels of either Lys (2 trials), Met+Cys, Thr (2 trials), Trp, Arg, Ile, or Val was investigated. Since Lys is the reference amino acid, two trials were performed. Also Thr was examined twice because of the unexpected low ideal Thr:Lys ratio of 63 in the first trial. However, the second experiment exactly confirmed the results of the first trial.

As shown in Figures 3 and 4 for Lys and Met+Cys, birds clearly responded to increasing dietary Lys and Met+Cys levels. These responses were non-linear; suggesting data analysis by exponential regression rather than by a linear (broken-line) model. The latter model has a defined breakpoint but often underestimates the optimum dietary amino acid level. Moreover, a linear-plateau relationship is difficult to explain especially when the response of a population of animals is investigated (Morris, 1989). In case of the exponential model, the optimum amino acid level is sometimes defined as 95 % of the asymptotic (= maximum) response. However, 95 % is arbitrarily chosen and any change to higher values will have a strong impact on the final recommendation. This would mean for the example shown in Figure 3, that with rising from 95 % to 99 % of the asymptotic response, the respective digestible Lys level shifts from 1.02 % to 1.17 % for the Ross 208 broilers. Due to the fact that a certain point is defined and that these points represent marginal amino acid levels rather than

Figure 3: Response of male Ross 208 broilers to graded levels of true fecal digestible Lys (Mack et al., 1999) Weight gain (g)
1800

1700

Ideal amino acid composition obtained by dose-response studies an experimental approach

As outlined above, the weakness of the factorial approach is the assumption of linear dose-response relationships regarding the utilisation of absorbed amino acids. The weakness of the literature survey is based on the fact that differences in experimental design and conditions are not considered properly. These downsides can be overcome by

1600

True fecal digestible Lys

1500
1.02 % at 95 % of the asymptotic response 1.17 % at 99 % of the asymptotic response

1400

0.87 % with the broken line-model

1300 0.70 0.80 0.90 1.00 1.10 1.20

True fecal digestible Lys (% of diet)

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excess levels, the ideal amino acid to Lys ratios in the project of Mack et al. (1999) were determined by the brokenline model. Subsequently, the Lys requirement was determined by exponential regression analysis. In Figure 4, the response of Ross 208 broilers to graded Met+Cys levels is shown. Relating the Met+Cys breakpoint (0.65) to the Lys breakpoint (0.86, average of two trials), the ideal ratio is 75 %. This result is pretty similar to the ratio suggested by Baker (1994). The complete ideal protein profile obtained by Mack et al. (1999) is listed in Table 1. The data provided by Baker et al. (2002) were also obtained by multiple experiments (Table 1). In the project of Mack et al. (1999), an optimum true fecal digestible Lys level of 1.15 % was found. This figure reflects the level required to minimise feed conversion in both Lys experiments, and it represents 95 % of the as-

ymptotic response. However, using the exponential equation, any percentage of the asymptotic response can be calculated. In addition, the exponential formulas can easily be combined with economic parameters to estimate the most profitable dietary amino acid level as described by Pack and Schutte (1995) or Hhler (2000).

The deletion method a further experimental approach

Recently, an ideal protein for starting broiler chicken has been published which was determined by the deletion method (Gruber, 1999, partly published by Gruber et al., 2000), a method originally established in swine research. The deletion method is based on the concept that the reduction of a non-limiting amino acid has no effect on nitrogen retention. The change in nitrogen retention (as % of N-intake) due to a reduced

proportion of each amino acid is used to determine a dietary amino acid profile in which all the amino acids tested were equally limiting - an ideal protein. This principle is shown in Figure 5 and explained below using the Lys and Met+Cys data reported by Gruber et al. (2000, Expt. 2). Male eight day-old broilers of a Ross strain were housed individually in cages. After twenty days of feeding the experimental diets, birds were slaughtered in order to determine nitrogen (N) retention by comparative slaughter technique. The final calculations of the deletion method are based on the bird's responses to feeding the control diet on the one hand and the test diets on the other hand. In contrast to the control diet of the multiple dose-response trial approach, the control diet of the deletion method is a positive control and is not deficient in any amino acid. The content of each of these amino acids is set as

Table 1: Selected Ideal Protein profiles (Lys = 100 %) for broilers taken from different references Phase Source Method NRC (1994)a Literature survey total 1.10 e 46 82 73 18 114 73 109 82 122 32 GfE (1999)b Factorial approach total 1.09 e,f 36 71 67 16 108 69 112 96 118 32 Starter Baker (1994)c Factorial approach Baker et al. (2002) Doseresponse study Gruber (1999) Deletion method Starter/ Grower Schutte (1996)d Literature survey NRC (1994)a Literature survey total 1.00 e 38 72 74 18 110 73 109 82 122 32 Grower GfE (1999)b Factorial approach total 0.92 e,f 38 87 76 17 117 78 125 109 137 33 Baker (1994)c Factorial approach Mack et al. (1999) Doseresponse study

Based on ... amino acids Lys (%) of diet Met Met+Cys Thr Trp Arg Ile Leu Val Phe+Tyr His
a b c

true true true apparent digestible digestible digestible digestible 1.12 g 36 72 67 16 105 67 109 77 105 35 1.03 g 56 17 61 78 not stated 1.05 /1.02 e 37 70 66 14 108 63 108 81 121 38 38 73 65 16 105 66 80 -

true true digestible digestible 0.89 g 37 75 70 17 105 67 109 77 105 32 1.15 g 75 63 19 112 71 81 -

Amino acid ratios to Lys

Ratios calculated, not explicitly given in the recommendations German Society for Nutrition Also known as Illinois Ideal Chicken Protein d Also known as CVB (Dutch Centraal Veevoederbureau) recommendations e Mixed sexes, f Recalculated assuming dry matter content of the diet: 88 % g For male birds

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100 %. The test diets are identical to the control diet except for the respective amino acid under test, which is reduced by a certain percentage. In the example shown in Figure 5 (left), the Lys and Met+Cys levels in the control diet were 0.95 % (=100 %) and 0.81 % (=100 %), while in the test diets the dietary Lys or Met+Cys content was reduced by 20 % or 35 % to 0.76 % or 0.53 %, respectively. The N-retention achieved with the control diet, in this case 56.2 % of Nintake (Gruber et al., 2000), was also set as 100 %. Subsequently, the animals' responses to the test diets were related to the N-retention of the control group. Birds receiving the Lys test diet retained 16.2 % less and those fed the Met+Cys test diet retained 14.4 % less nitrogen compared to the control. That means, per percentage point of Lys removal, Nretention decreased by 0.81 %, whereas this figure for the Met+Cys group was only 0.41 %. The relative effect of Lys reduction was stronger being a prerequisite for taking Lys as reference. A crucial point of the concept is that the effects of the test diets on N-retention are related to the slope of the regression line between the Lys test and the control group. It is assumed that the slopes of the individual regression lines are

Figure 4: Response of Ross 208 broilers to graded levels of true fecal digestible Met+Cys (Mack et al., 1999) Weight gain (g)
1900

1800

1700

True fecal digestible Met+Cys 0.73 % at 95 % of the asymptotic response 0.65 % with the broken line-model

1600

Ideal ratio to Lys 75 % according to break points

1500 0.50 0.60 0.70 0.80 0.90

True fecal digestible Met+Cys (% of diet)

identical for all essential amino acids. According to this assumption, a point on the x-axis can be calculated for each amino acid - other than Lys - down to which a reduction of an amino acid would not have affected N-retention. Accordingly, the Met+Cys level of the control diet could theoretically have been reduced to 83 % without an effect on N-retention (Figure 5). Eighty-three percent of 0.81 % Met+Cys in the control diet is 0.67 %. Therefore, with

0.67 % Met+Cys and 0.95 % Lys both amino acids are in the proper ratio (Met+Cys:Lys = 71 %) to maintain performance and to avoid a relative excess of dietary Met+Cys. The ideal protein profile derived by Gruber et al. (2000) is listed in Table 1. Since the effects of the test diets are related to the slope of the Lys dose-response, special attention has to be given to the latter. The linear regression is

Figure 5: Principle of determining ideal amino acid to Lys ratio taking Met+Cys (Gruber et al., 2000) as example (left) and influence of the amino acid level of the control diet on the slope of the Lys dose-response line if only two data points are used (right) N-retention (% of control treatment)
110 100 90
Amino acids in control diet Digestible Lys content 0.95 % Digestible Met+Cys content 0.81 % Ideal Met+Cys:Lys ratio y = (0.81 % Met+Cys x 0.83)/0.95 % Lys = 71 %

N-retention (% of linear control)

110

100

80 83 %

Response to Met+Cys reduction Response to Lys reduction

90

Lys - linear range Lys - flattening part Lys - plateau

70 60 65 70 75 80 85 90 95 100 105

80 75 85 95 105 115 125 135

Amino acids in the diet (% of control diet)

Digestible Lys content (% of linear control diet)

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done using only two data points which means once the dietary Lys exceeds the broiler's requirement the slope of the line will decrease, thus distorting all conclusions drawn regarding the ideal protein. This risk is indicated in Figure 5 (right). Therefore, the N-retention level derived by the control diet has still to be kept in the linear range of the non-linear response curve. The assumption that the slopes of the dose-response relationships are identical for all amino acids might be a matter for criticism since as shown in Figure 1 the three compartments defining the total requirement contribute to different proportions depending on the amino acid.

Figure 6: Response of fat and lean line broilers to dietary threonine (data obtained by Alleman et al. (1999), recalculated) Weight gain (g)
1100

900

lean line

fat line

700

0.57 % True fecal digestible Thr for the lean line at 95 % of the asymptotic response 0.48 % True fecal digestible Thr for the fat line at 95 % of the asymptotic response

500 0.35 0.40 0.45 0.50 0.55 0.60

True fecal digestible Thr (% of diet)

Ideal amino acid ratios might be affected by environmental issues

Comparing the outcome of all the different approaches above to determine ideal amino acid ratios, the obtained amino acid profiles are surprisingly similar - disregarding their advantages and limitations. Thus, it can be stated that we have a reasonably clear picture of the ideal amino acid profile needs of current broiler strains. However, there are some indications that the ideal ratios might be affected under certain circumstances. One factor is, as described above, the age or body weight of the birds. Other factors, which might interfere with ideal amino acid ratios can be due to environmental conditions. Prominent examples are the Arg:Lys and Thr:Lys ratios. general performance level in these trials was lower at high temperature and the authors further reported that in vitro experiments revealed a significantly reduced Arg uptake by the intestinal epithelium under heat stress conditions. From these statements it could be concluded that the lower absorption rate at high temperature might partly be compensated by increased Arg levels which in turn would result in higher ideal Arg:Lys ratios. In addition, it seems that this interaction is further complicated by interactions between sodium (chloride) and Arg. More research is needed in this field. er dietary Thr levels to achieve maximum performance or minimum feed conversion under dirty production conditions compared to those raised in a clean environment. This finding matches results reported for swine, where increased physiological or digestive stress, respectively, produced by increased dietary pectin levels resulted in an enhanced Thr requirement (Zhu and de Lange, 2001). Increased mucin secretions into the gut as a response to increasing pectin levels might serve as one explanation for this phenomenon because of its extraordinary high Thr content of 16.4 % (Lien et al, 1997). In addition, immune globulins also contain relatively high amounts of Thr. Therefore, the low Thr:Lys ratios of 63 % (Mack et al., 1999) and even 56 % Baker et al., 2002) might be the result of excellent experimental conditions and the high digestibility of the experimental diets. Thus, it is likely that under practical conditions optimum Thr:Lys ratio is higher than 63 %. However, this issue is currently a matter of investigation. Taking Arg and Thr as examples, it appears that ratios between certain amino acids can be affected by certain exogenous circumstances.

Thr:Lys ratio Comparing the ideal protein profiles published by Mack et al. (1999) and Baker et al. (2002) with the other ones presented in Table 1, particularly the low Thr:Lys ratios of 63 % or 56 % are remarkable. This is especially the case since both Thr experiments performed by Mack et al. (1999) revealed the same ideal ratio. However, recent research indicates interactions between Thr requirement and environmental conditions. Kidd et al. (2002) reported that 42 to 56 days old broilers required high-

Arg:Lys ratio According to the ideal protein profiles cited herein, ideal Arg:Lys ratios based on digestible amino acids vary between 105 % and 112 % (Table 1). However, Brake et al. (1998) reported that particularly weight gain but also feed conversion was improved with Arg:Lys ratios up to 139 % at high temperature while this effect could not be observed under moderate temperature conditions. The

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The impact of genetics on optimum amino acid ratios

There is evidence that genetics in terms of sex, strain, or genetic progress over the years might influence the optimum amino acid ratio. Generally, any shift in proportion of body protein, feathers, and maintenance requirement possibly results in changes in ideal amino acid ratios. This principle is illustrated in Figures 1 and 2 for the Met+Cys to Lys ratio with age. Based on this principle, Fisher (2002) recently showed how ideal Met+Cys:Lys ratios differ between male and female broilers. Accordingly, the ideal ratio for male broilers increased to a maximum at about five weeks of age and maintained there until eight weeks. Regarding female birds, maximum Met+Cys:Lys ratio was already achieved at three to four weeks but at a clearly higher level and dropped then until eight weeks of age to the same level as reported for the males. These findings reflect the differences in genetic potential between genders. Based on further simulations Fisher (2002) pointed out that especially feathering rate is a potential source for the variation of the ideal Met+Cys:Lys ratio.

A broiler experiment conducted with extreme genetic differences has been reported by Alleman et al. (1999). They compared the responses of male genetically lean or fat line broilers to graded levels of dietary Thr fed from 28 to 49 days of age. The study revealed that the magnitude of the response starting from a Thr deficient situation was clearly stronger in the lean line compared to the fat line broilers (Figure 6). The data were re-analysed by an exponential regression model and at 95 % of the asymptotic response, optimum digestible Thr level would be 0.57 % and 0.48 % for the lean and fat line birds, respectively. Keeping in mind that a big portion of the dietary Thr is required by the gut itself as reported for piglets by Law et al. (2000), and that the genetic progress in terms of weight gain or feed conversion is finally the result of altered organ capacity and efficiency, the conclusion that ideal dietary amino acid proportion will change with genetic progress can easily be drawn.

Current suggestions for an ideal protein are fairly consistent

In summary, it can be stated that although there are different methods to determine an ideal protein available results are fairly similar. Apparent differences could be attributed to the method applied for ideal protein determination but also to interactions between amino acid requirements and other animal related and/or environmental factors. The most important factors are age, climate, management as well as genetic factors like sex or strain. Applying the ideal protein concept regardless of small variations has been shown to be effective. This will be the subject of a separate paper, which will be published soon in this magazine.

References Alleman, F., J. Michel, A. M. Chagneau and B. Leclercq (1999): Comparative responses of genetically lean and fat broiler chickens to dietary threonine concentration. British Poultry Science 40: 485-490. Baker, D. H. (1991): Partitioning of nutrients for growth and other metabolic functions: efficiency and priority considerations. Poultry Science 70: 1797-1805. Baker, D. H. (1994): Ideal amino acid profile for maximal protein accretion and minimal nitrogen excretion in swine and poultry. Proceedings Cornell Nutrition Conference, 56th Meeting, 18.-20. October, Rochester, New York, USA: 134-139. Baker, D. H., A. B. Batal, T. M. Parr, N.-R. Augspurger and C. M. Parsons (2002): Ideal Ratio (relative to lysine) of tryptophan, threonine, isoleucine, and valine for chicks during the second and third weeks posthatch. Poultry Science 81: 485-494. Brake, J., D. Balnave and J. J. Dibner (1998): Optimum dietary arginine : lysine ratio for broiler chickens is altered during heat stress in association with changes in intestinal uptake and dietary sodium chloride. British Poultry Science 39: 639-647. Fisher, C. (2002): Possibilities and limitations of ideal protein concepts in evaluation of amino acids for fast growing poultry. 11th European Poultry Conference, 6.-10. September, Bremen, Germany. GfE (1999): Empfehlungen zur Energieund Nhrstoffversorgung der Legehennen und Masthhner (Broiler). DLG Verlag, Frankfurt a.M., Germany. Gruber, K. (1999): Experimentelle Untersuchungen zum idealen Aminosurenverhltnis in der Broilerftterung. Doctoral thesis, Institute of Nutrition Physiology of the Technical University of Munich, Germany. Gruber, K., F. X. Roth and M. Kirchgessner (2000): Effect of partial dietary amino acid deductions on growth rate and nitrogen balance in growing chicks. Archiv fr Geflgelkunde 64 (6): 244-250.

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Hoehler, D. (2000): Evaluation of amino acid dose-response data and implications for commercial formulation of broiler diets. AminoNewsTM 1 (1): 9-16. Kidd, M.T., W. A. Dozier III, S. J. Barber, W. S. Virden, D. W. Chamblee and C. Wiernusz (2002): Threonine needs of Cobb male broilers from days 42 to 56. Abstracts International Poultry Scientific Forum, 14.-15. January, Atlanta, Georgia, USA, 5-6. Law, G., A. Adjiri-Awere, P. B. Pencharz and R. O. Ball (2000): Gut mucins in piglets are dependent upon dietary threonine. Advances in Pork Production, Proceedings of the 2000 Banff Pork Seminar, Volume 11, Abstract #10. Lien, K., W. Sauer and M. Fenton (1997): Mucin output in ileal digesta of pigs fed a protein-free diet. Zeitschrift fr Ernhrungswissenschaft 36 (2): 182-190.

Mack, S., D. Bercovici, G. de Groote, B. Leclercq, M. Lippens, M. Pack, J. B. Schutte and S. van Cauwenberghe (1999): Ideal amino acid profile and dietary lysine specification for broiler chickens of 20 to 40 days of age. British Poultry Science 40: 257-265. Morris, T. R. (1989): The Interpretation of response data from animal feeding trials. In: Recent developments in Poultry Nutrition. Butterworths, London, UK. Ed.: Cole, D.J.A. and W. Haresign. 1-11. NRC (1994): Nutrient requirement of poultry - Ninth revised edition. National Academy Press, Washington, D.C., USA. Pack, M. and J. B. Schutte (1995): Sulfur amino acid requirement of broiler chicks from fourteen to thirty-eight days of age 2. Economic evaluation. Poultry Science 74: 488-493.

Schutte, J. B. (1996): Aminozurenbehoefte von leghennen en vleeskuikens. CVB (Centraal Veevoederbureau) documentatiereport nr. 18. Zhu, C. L. and C. F. M. de Lange (2001): Influence of graded levels of pectin on the utilization of dietary threonine and lysine for body protein deposition. Proceedings of the Canadian Society of Animal Science.

Dr. Andreas Lemme email: andreas lemme@ degussa.com

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