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Optimal dietary arginine levels in modern broiler chickens

Published: June 18, 2019
By: I.C. Ospina-Rojas, R.J.B. Rodrigueiro and L. Otani / CJ do Brasil. Av. Engenheiro Luís Carlos Berrini, 105 - Monsões, 04571-010, São Paulo - SP, Brazil.

Arginine has a vast variety of effects on metabolism, including protein synthesis, immunity, wound healing, hormone secretion, neurotransmission, normal cell differentiation, and growth, vasodilation, and antioxidant activity. Several of these effects are mediated by nitric oxide that elicits a surprisingly wide range of physiological functions and arginine is the only effective precursor in birds. Broilers are usually exposed to various stressors during their rearing period (vaccination, cold or heat stress, immune challenge, and high stocking density among others) that results in economic expenses by decreased growth and increased susceptibility to diseases. Because dietary arginine levels could be insufficient for a broiler’s metabolic needs during stress periods, arginine supplementation has been considered important to attenuate reduced performance of broilers in these conditions. Dietary arginine:lysine ratio was estimated at 114 and 115 for optimal performance of broilers during the starter and grower phases, respectively. However, these ratios may be greater to maximize immune system function and skin breaking strength. Higher arginine levels may be an interesting strategy to minimize carcass condemnation issue in the poultry industry, reducing the infected scratches and improving wound healing as well as carcass quality. 

Dietary arginine supplementation higher than the recommended levels was shown to improve broiler performance (Murakami et al. 2012; Xu et al., 2018; Zampiga et al., 2018). It suggests that arginine levels needed for maximum performance could be higher in modern broilers, which is probably related to the several functions of arginine on animal metabolism. Besides, arginine enhances the protein synthesis via the activation of the mTOR signaling pathway (Yao et al., 2008; Yuan at., 2015); this amino acid is a precursor for critical molecules such as nitric oxide, creatine (Chamruspollert et al., 2002), polyamines, proline, glutamate, citrulline, and agmatine (Wu and Morris, 1998). Arginine regulates key aspects of animal growth by enhancing insulin, growth hormone, and insulin-like growth release (Cochard et al., 1998; Yu et al., 2018; Xu et al., 2018). However, there is limited research on arginine requirements in broilers, and most research has focused on arginine supplementation in birds receiving some physiological stress (immune challenge, cold, or heat stress), wherein dietary arginine levels could be insufficient to support broiler growth or meet their metabolic requirements during those stress periods. Due to arginine’s role in healing wounds and reducing the carcass incidence of skin scratch infections, arginine supplementation could be a nutritional strategy to reduce the relevant poultry economic losses with carcass condemnation issues.
Optimal dietary arginine levels for broiler performance
A meta-analysis based on 94 observations from eight peer-reviewed papers was conducted to determine the optimal arginine levels in broiler diets. The arginine base data included peer-reviewed papers between 2001 and 2019. It has used data of birds raised in normal conditions and excluded the dose-response data of broilers challenged or raised in suboptimal temperatures. When not given the standardized ileal digestible (SID) amino acid contents of the experimental, they were calculated using digestibility coefficients of ingredients as proposed by Rostagno et al. (2017). Summary references and some experimental conditions used for the meta-analysis are shown in Table 1.
The BW gain distribution of base data for each growth period was described in Figure 1. Arginine requirement during the finisher phase has been seldom studied, data obtained from this period showed a larger variation than the observations from the starter and grower phases where arginine supplementation has shown to increase breast meat yield (Corzo et al., 2003; Jiao et al., 2010) while reducing wooden breast myopathy incidence in modern fast-growing broilers (Bodle et al., 2018; Zampiga et al., 2019).
Optimal dietary arginine levels in modern broiler chickens - Image 1
Optimal dietary arginine levels in modern broiler chickens - Image 2
BW gain data (g/d) plotted against arginine intake (g/d) showed a positive response in BW gain as increasing arginine intake (Figure 2). A significant quadratic regression (P=0.0138) was fitted from the 94 observations to predict the daily SID arginine intake to achieve a desirable BW gain during the different growth phases. For instance, an optimal level of SID arginine of 1.45% would be needed to achieve a daily BW gain of 19.8 g in broilers from 1 to 7 days of age (Table 2).
Optimal dietary arginine levels in modern broiler chickens - Image 3
Optimal dietary arginine levels in modern broiler chickens - Image 4
Considering a SID lysine recommendation of 1.28% and 1.07% for birds from 1 to 21 days and 22 to 42 days according to Rostagno et al. (2017), the ideal SID arginine:lysine ratio could be estimated at 114 and 115 for broilers during the starter and grower periods, respectively.
To standardize and compare performance data from broilers at different ages, dose-response data was expressed as a percentage. The best performance was considered as the reference (100%) and their respective dietary SID arginine level and arginine:lysine ratio were set at zero. The other treatments were expressed relative to the reference treatment. Quadratic regression (QR) and linear response plateau (LRP) models were used to determine the optimum arginine:lysine ratio for the relative BW gain. Also, the first point where the quadratic response curve interested the plateau value was determined (LRP + QR). The SID arginine:lysine ratio was estimated to be at 104, 110, and 127 based on LRP, LRP + QR, and QR models, respectively (Figure 3). The ideal arginine:lysine ratio may be at 114 considering the three approaches. This value is close to the arginine:lysine ratio of 114 and 115 estimated for broilers during the starter and grower periods using a fitted equation in Figure 2.
Optimal dietary arginine levels in modern broiler chickens - Image 5
Recommendations of arginine:lysine ratio range in the literature from 105 to 125, depending on the variable studied, with a mean of 115% (Table 3) (NRC, 1994, Lippens et al., 1997; Baker, 1997; Kidd et al., 2001; Corzo and Kidd, 2003; Jahanian, 2009; Corzo et al., 2012). The SID arginine:lysine ratio to optimize the immune system response is higher than that for optimal feed conversion which is higher than for maximum BW gain (Corzo et., 2003; Jahanian, 2009). Corzo et al. (2012) indicated that broiler chicks optimize their BW gain and feed conversion values at arginine:lysine ratios of 108 and 114, respectively. The SID arginine:lysine ratio estimated from the nutritional recommendations of Jahanian (2009) was 115, 117, and 122 to optimize BW gain, feed conversion, and immune system function, respectively.
Optimal dietary arginine levels in modern broiler chickens - Image 6
Effect of arginine levels on immune response
The higher arginine level needed for maximum immune system function is expected considering the important role of arginine in immunity as it is a substrate for the immune cells. The immune cells use the system y+ cationic amino acid and B0,+ transporters to acquire arginine (Nicholson et al., 2001; Humphrey et al., 2004) where this amino acid promotes lymphocytic proliferation through polyamine synthesis (Ochoa et al., 2001) and improve the phagocytic activity of macrophages by increasing their internal nitric oxide levels (Sung et al., 1991). Consequently, functional aspects of T and B cells are reliant on arginine utilization (D’Amato and Humphrey, 2010).
Dietary arginine supplementation in broilers challenged with Eimeria spp. has shown to reduce mortality (Kidd et al., 2002) and alleviate intestinal villus damage (Tan et al., 2014) as well as attenuate gut injury of broiler chickens infected with Clostridium perfringens (Zhang et al., 2018). Also, Corzo et al. (2012) reported a linear reduction of mortality in broilers supplemented with graded levels of arginine. The positive effects of arginine in challenged broilers (having an increased concentration of CD4+ and CD8+ lymphocytes and B cells, increased antibody titers after vaccination, increased relative weight of the thymus and bursa, and improved T cells response) may be due to improve the general health status (Tayade et al., 2006; Abdukalykova et al., 2008; Jahanian, 2009).
Arginine supplementation as a strategy to reduce broiler carcass condemnation
Interestingly, Corzo et al. (2003) found a linear reduction in infected carcass scratches of broilers fed arginine-supplemented diets (Table 4). Infected skin scratches as cellulitis are considered the first major reason for condemnation at slaughter followed by fractures/contusion (Santana et al., 2008), which is associated with significant economic losses in poultry industry. The reduced infected scratches in arginine-supplemented diets are not only related to the good immunity status as well as to a dermis with greater resistance. The arginine role in enhanced wound healing has been well-documented in mammals by increasing breaking strength and reparative collagen accumulation (Shi et al, 2003; Tong et al., 2004; Stechmiller et al., 2005). For instance, the production of nitric oxide synthase peaks within 24–72 h after wounding (Stechmiller et al., 2005), producing nitric oxide, and citrulline, where nitric oxide plays a major role in antimicrobial activity and improving blood flow to the healing wound (Alexander and Supp, 2014). Thereafter, arginase activity increases with a marked rise in wound ornithine levels which is metabolized to polyamines by ornithine decarboxylase (Shi et al., 2002). The polyamine concentrations increase within 12 h after skin incisional wounding in rats and they are factors necessary for cellular growth involved in tissue repair and wound healing (Maeno et al., 1990). The inhibition of ornithine decarboxylase, an enzyme related to polyamine synthesis, impairs skin wound healing (Lesiewicz and Goldsmith, 1980).
Optimal dietary arginine levels in modern broiler chickens - Image 7
In addition, arginine is a metabolic precursor of proline and hydroxyproline, the major amino acids in the collagen proteins, which are major extracellular components in connective tissues of skin and tendons (Wu et al., 2011). Fernandes (2007) reported a linear increase in broiler dermis thickness as increased dietary arginine levels. It suggests the importance of arginine in skin breaking strength. Scratches with no dermis penetration resulted in a more resistant dermis having less chance to be infected. The effect of arginine supplementation should be further studied considering the huge economic benefits that it may bring to poultry production to reduce carcass condemnation.
Arginine and lysine antagonism effects on broiler performance
When the best performance treatment in each study was considered as the reference (100%) and its respective dietary arginine:lysine ratio was set at zero from dose-response data, birds fed SID arginine:lysine ratios higher than the optimal level did not show pronounced negative effects on broiler performance (Figure 4). The optimal SID arginine:lysine ratios used as reference were above nutritional recommendations in 75% of the peer-reviewed papers used for the meta-analysis. On the contrary, marginal arginine:lysine ratios reduced drastically BW gain and feed conversion. The antagonism between dietary arginine and lysine has long been defined. This effect is clearer with an excess of lysine and marginal arginine level (low arginine:lysine ratio) rather than an excess of arginine (high arginine:lysine ratio) (Balnave and Brake, 2002). Indeed, higher arginine:lysine ratio can improve broiler performance (Murakami et al. 2012; Zampiga et al., 2018).
High lysine levels increase the renal arginase activity and consequently increase arginine degradation (Austic and Scott, 1975). As a result, less arginine is available for protein synthesis and biological functions that rely on arginine. Also, excess lysine decreased the arginine:glycine amidinotransferase activity (Jones et al., 1967), which catalyzes the first step of creatine synthesis, depressing muscle creatine concentration (Chamruspollert et al., 2002). The antagonism between arginine and lysine is more pronounced in birds that are not able to synthesize arginine endogenously due to the lack of carbamoyl phosphate synthetase I and a low renal activity of argininosuccinate synthetase, argininosuccinate lyase, and ornithine transcarbamylase (Ball et al., 2007). Therefore, arginine is an essential amino acid, and its requirement must be supplied by diet to support broiler growth.
Optimal dietary arginine levels in modern broiler chickens - Image 8
The effect of heat and cold stress on arginine requirements
The environmental conditions in arginine studies are important to be considered to avoid any interference in the estimated SID arginine levels. Blood flow is redirected to peripheral organs involved in heat dissipation, when broilers are exposed to a warm environment. Therefore, dietary arginine requirement may increase in high temperature as this amino acid is a substrate for nitric oxide synthesis, which plays an important role in vasodilation and blood flow (Perez Laspiur et al., 2006). Likewise, high environment temperature demonstrated to decrease arginine absorption in the intestinal epithelium increasing their dietary needs (Brake et al., 1998). Besides alleviating the adverse effect of heat stress, arginine supplementation has shown to recover broiler performance of birds submitted to cold stress (Saki et al., 2013; Kodambashi Emami et al., 2017). In birds subjected to low-temperature stress, the sympathetic nervous activity is stimulated to produce heat (thermogenesis) by increasing the bird cellular metabolic rate and cell oxygen intake (Estler and Ammon, 1969). The relative underdeveloped cardio-respiratory system of modern broilers fails to fulfill the required oxygen demand, resulting in hypoxemia (Aftab and Khan, 2005). Thus, the bird increases heart rate and blood flow to compensate for the oxygenation deficit that may result in pulmonary hypertension and ascites (Olkowski et al., 1999). A previous study reported that arginine supplementation in broilers exposed to cool environmental temperature attenuated the effects of pulmonary hypertension (Wideman et al., 1995) by increasing the endogenous pulmonary vasodilator nitric oxide. Therefore, either cold or heat stress increase arginine needs being an important amino acid to attenuate the negative effects on performance of broilers submitted to suboptimal temperatures. In addition, the reactive oxygen species production is higher in birds raised in high- or low-temperature environments, increasing the oxide nitric production for its well-documented potent cellular antioxidant function. Therefore, arginine levels could be insufficient to support broiler growth or meet their metabolic requirements during stress periods.

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Related topics:
Iván Camilo Ospina-Rojas
CJ Bio
Ramalho Rodrigueiro
CJ Bio
Lyssa Otani
CJ Bio
Alvaro Dubois
23 de octubre de 2020
Dear Ospina-Rojas. Thanks for the reply but my doubt is really about the regression generated with this LRP-QR (97.53= 9.069+1.4213x-0.0056*x2) which is presented in fig. 3. With just one value per paper, regressing it against Arg:Lys ratio sounds a bit odd. Not saying it’s necessarily wrong, but considering that both weight gain and Arg:Lys were standardized based on the point of maximum response, in theory you are setting your requirement as 100% or very close to it. The fact that the quadractic model gave you a much larger requirement is by itself a testimony of the problem of setting requirements based on quadractic curves. LRP model predicted 104% which is much closer to the expected value of 100%. Having this average between the two models diminish but still maintain the problem. Thank you.
Juarez Donzele
Universidade Federal de Viçosa - UFV
17 de octubre de 2020

The different considerations that have been made on this matter seem relevant to us. Now regarding my placement, which would be more coherent to use only the works in which diets with sub-optimal level of digestible lysine (LD) were used, I do it because I understand that this is the most accurate methodology for determining the ratio of any amino acid (Aas ) with LD. When I use a diet with a sub-optimal LD level, in addition to the advantage of avoiding a possible excess of LD consumption, which would result in underestimating the relationship, there is also a second advantage that would be related to the fact that, with the use of a sub-optimal level of LD, the accuracy to determine the deficiency of a second Aas, in this case arginine, would be increased. Just check the quadratic regression to see that it is more sensitive to determine the relationship with the LD in the ascendant than at the inflection point of the curve, which represents the animal's requirement. I tried to focus on aspects related to the methodology used in the choice of works.

Alvaro Dubois
16 de octubre de 2020
Dear Ospina-Rojas. Congratulation for the very good review and discussion on the importance of arginine in general metabolism. About the mathematical analysis of the data, besides the importance of taking into account lysine levels used in the trials, as mentioned in the other comments, I would like to point that in the regression calculated in fig. 2, a large part of the difference in weight gain and arginine intake comes from the difference in age between the different sets of data and not due to difference in arginine levels used in each paper. In this way, there's a large degree of confounding between the two factors (age and arginine intake). For this reason, although you can establish a large correlation between the two factors, you can not properly define a cause:effect relation. The second approach (fig. 3) is a more proper way of analyzing the results but today mixed models analysis is considered the best way to do it. Still on fig. 3. I couldn't quite figure out what is the regression in red ((LRP+QR). Considering that you would have only one value for it in each study, what is exactly the regression meaning? Thank you.
Juarez Donzele
Universidade Federal de Viçosa - UFV
15 de octubre de 2020
Iván Camilo Ospina-Rojas, I believe that Dr Andreas' considerations are pertinent. As for the information used to obtain the digestible arginine (ARGD): digestible lysine (L D) ratio, I think it should be used only from studies where this determination was made using a diet with a sub-optimal level of LD. This is because in fact the birds demand is Aas gram per day not percentage. In this way, using the diet with suboptimal level of LD, the ARGD: LD ratio would be more accurately determined, since it would avoid the possibility of birds consuming LD above the requirement, which would reflect on the determination of the ideal relationship between these two amino acids. Thus, the use of this methodology, a diet with a sub-optimal level of LD, is more appropriate when compared to the determination of the diet in which the LD level was in the requirement of the birds.
Dr. Andreas Lemme
Evonik Animal Nutrition
15 de octubre de 2020

Dear Camilo, it is always a challenge to compare literature data from different institutes produced under different conditions and to extract conclusions and/or recommendations. However, compiling the data is one issue and you determined (modeled) the SID Arg requirement accordingly. Indeed, changing production conditions might influence optimal concentrations.
You also explained how you determined the optimal SID Arg:Lys ratio and I would have two questions in this context:
1) why did you chose the Brazil tables for SID Lys? There are many more sources/recommendations available globally. However, my point is that the recommended ratio is highly dependent on the Lys level of the reference and thus to a large degree independent from the Arg-research - not considering Lys level of the individual studies considered.
2) you analysed the dose-response by three models and finally averaged the results. Why didn't you decide for one? The outcome of the different models would have tremendous impact on practical feed formulation.
Best regards

Juarez Donzele
Universidade Federal de Viçosa - UFV
29 de septiembre de 2020

Iván Camilo Ospina-Rojas, good article, with an enlightening focus. I take advantage of the material presented here, which somehow comes to consolidate some information that we publish, right here at ENGORMIX with the title; Digestible lysine levels obtained by two methods of formulation of diets for 22-to-42-day-old broilers, with a focus on differentiating diets with or without the use of industrial amino acids (Aas). In this study, it was evidenced that diets in which the levels of digestible lysine (LD) (0.90-1.00-1.10 and 1.20%) were obtained by varying corn and soybean meal (full protein) provided better performance of chickens compared to those fed diets in which LD levels, from the baseline level of 0.90%, were achieved using Aas. It should also be considered that in diets supplemented with Aas, because they contain a lower level of CP, they will consequently contain lower levels of amino acids considered non-essential and a reduction in the amount of arginine, which corresponds to that above the ratio arginine x lysine proposed in the protein. ideal for the category of birds being studied. Just as this study showed that the level of arginine can limit the growth of chickens, despite being at the established requirement level, several studies have also shown that some non-essential Aas such as; glutamine, glycine, aspartate can also compromise the performance chickens when using diets with low LD levels. It is concluded, finally, that the practice of reducing CP of the diet with supplementation of Aas must be carried out with great discretion.

George Entz
28 de septiembre de 2020

Thank you for your response. I can't seem to find any commercially available Arginine in western Canada, so far.
If it's not available, Is there a way to reduce the negative effects, of a low arginine:lysine ratio.

Some research is suggesting DEB levels and Methionine source could be factors looking into it.

My thoughts are this, it would be great to use L-Arginine or even feeding a Corn-SBM diet (versus a Wheat-SBM) but cost restriction on both corn and L-arginine makes it so that's it's a hard ROI to use them here.

George Entz
19 de septiembre de 2020

The ideal Arg:Lysine Ratio can not always be obtained in some situations.

For example: When the grower Diet is reduced below 20% Crude protein due to feeding program (RWA + VGF birds vaccinated with coccivac B52 to control coccidiosis) and having to use 16 % crude protein wheat with SBM as the main ingredients with no L-arginine to supplement.

So my question is what can be done to reduce some of the negative effects then of a low Arg level in a diet (dig 0.90%)? This problem would be for days 14-26 of age. After that, the CP can be raised.

Looking forward to your comments.

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