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METABOLISM AND NUTRITION Changes in chemical composition in male turkeys during growth V. Rivera-Torres,*† J. Noblet,‡§ and J. van Milgen‡§1 *Techna, BP10, F-44220 Couëron, France; †AgroParisTech, UFR Nutrition animale, qualité des produits et bien-être, Département Sciences de la Vie et de la Santé, F-75005 Paris, France; ‡INRA, UMR1079 Systèmes d’Elevage, Nutrition Animale et Humaine (SENAH), F-35590 Saint Gilles, France; and §Agrocampus Ouest, UMR1079 SENAH, F-35000 Rennes, France ABSTRACT In growing animals, requirements for many nutrients (and energy) are determined by the retention of these nutrients. During growth, this retention changes in an absolute way and also between nutrients and energy, resulting in changing nutrient requirements. The objective of this study was to describe the changes in chemical composition in male growing turkeys. The serial slaughter technique was used to determine the composition of amino acids, lipid, ash, and water in feather-free body (FFB) and feathers in male turkeys offered feed ad libitum from 1 to 15 wk of age. Allometric relations were used to describe changes in body composition. The feather content in the body decreased from 6% at 1 wk of age to less than 3% at 15 wk of age. The water and protein content in FFB decreased with increasing FFB mass, with allometric scalars (b) of, respectively, 0.967 and 0.970, whereas the lipid content increased with increasing FFB mass (b = 1.388). The water, protein, and ash content in fat-free FFB was constant and represented, respectively, 71.6, 24.2, and 4.2% of the fat-free FFB mass. The amino acid content of FFB protein was relatively constant and only the Cys content decreased between 1 and 15 wk of age, whereas the Ile content increased. Feathers were mostly composed of protein, and the protein content did not change during growth. During growth, the Lys, Met, Trp, His, Tyr, Asp, and Glu contents in feather protein decreased, whereas the Cys, Val, and Ser contents increased. The contribution of feathers to whole-body amino acid retention ranged from 5% for His to 33% for Cys. On average, the weight gain of FFB contained 21.3% protein and 12.7% lipid, corresponding to an energy content of 10.1 kJ/g. The weight gain of feathers contained 87.4% protein, corresponding to an energy content of 20.4 kJ/g. The results of the present study can be used in a factorial approach to determine nutrient requirements in growing turkeys. Key words: turkey, growth, body composition, amino acid, allometry 2011 Poultry Science 90:68–74 doi:10.3382/ps.2010-00633 INTRODUCTION mined at different ages, and measurements have to be performed in different conditions to determine the effects of environment and nutritional strategies (Fatufe et al., 2004; Lumpkins et al., 2007; Samadi and Liebert, 2007). The expression of amino acid requirement as a percentage in the diet or relative to dietary energy also requires information on energy utilization. Especially in turkey nutrition, these measurements are costly and time-consuming because of the sexual dimorphism associated with a long production cycle. The composition of BW gain contributes largely to the nutrient requirements of growing animals. A description of the dynamics of protein and amino acid retention can therefore be helpful in determining amino acid requirements. Although this approach has been used in broilers through allometric relations (Gous et al., 1999) and modeling (King, 2001), little information is available in turkeys (Firman and Boling, 1998). Hurwitz et al. (1983) used the factorial method to determine the amino acid requirements of turkeys, but Poultry meat is a major source of animal protein produced in the world. Although the production efficiency is high relative to other animal production systems, improving the efficiency remains a key issue to reduce production costs, the use of natural resources, and the impact on the environment. An excess supply of protein in the diet compared with the amino acid requirement of the bird is the main cause of the inefficient use of dietary nitrogen. In poultry nutrition, amino acid requirements are usually based on empirical definitions of the amino acid levels that optimize a response criterion such as maximization of BW gain, carcass yield, or feed efficiency (e.g., Moore et al., 2004; Barbour et al., 2008; Dozier et al., 2008). The requirements are deter©2011 Poultry Science Association Inc. Received January 6, 2010. Accepted October 4, 2010. 1 Corresponding author: [email protected] 68 BODY COMPOSITION IN MALE TURKEYS recommendations were given for different age classes, rather than being based on the dynamics of BW gain. Emmans (1989) described the change in body composition in male and female turkeys as a function of whole-body mass or protein mass of carcass and feathers. In addition, he described the amino acid requirement for maintenance and growth in feather-free body and feathers. However, a possible change in the amino acid profile of feathers and the feather-free body was not considered. As observed by Ferket et al. (1997), the amino acid profile of the feather-free body does not seem to change during growth, in contrast to that of feathers. The objective of this study was to determine the composition of BW and BW gain in male growing turkeys, with emphasis on the change in the relative contributions of feathers and carcass during growth. MATERIALS AND METHODS Birds and Diets The experiment was carried out at the same time as an experiment in which we measured the energy expenditure by indirect calorimetry (Rivera-Torres et al., 2010), and turkeys from the same batch were used in both experiments. In brief, 2 groups of medium-size male turkeys (B.U.T. 9) were obtained from a hatchery (France Dinde, Saint Hervé, France) on the day of hatching. Turkeys were housed in flat-deck cages and given ad libitum access to feed and water throughout the experiment. Four different diets were used during 4 successive periods of 4 wk each, and the nutrient supply of these diets met or exceeded NRC (1994) recommendations during each period. The anticipated digestible Lys contents were 1.48, 1.32, 1.08, and 1.00% for diets distributed during periods 1 through 4. Measurements and Sampling The change in body composition was determined by periodically slaughtering turkeys from each group. From 1 wk of age onward, 2 turkeys from each group were slaughtered every 2 wk so that at the end of the experiment, 4 replicate measurements were obtained at 8 different ages. Turkeys that were selected for slaughter were kept in a light room for 4 to 6 h before slaughter, with ad libitum access to water but without access to feed. They were then killed by intravenous injection of a mixture of embutramide, mebezonium, and tetracaine chlorydrate (T 61, Intervet SA, Beaucouzé, France) after anesthesia by an intravenous injection of ketamine (Imalgène 1000, Merial, Lyon, France). Each turkey was weighed immediately after this procedure. The body was then soaked in hot water (60°C) for 2 min, plucked, and placed in a room at ambient temperature until the skin was dry. The feather mass was calculated as the difference between BW before and after plucking. The birds were then eviscerated manually. The viscera included both the empty gastrointestinal tract and internal organs (including reproductive 69 organs). The feather-free body (FFB; i.e., carcass plus empty viscera) was stored in plastic bags and frozen at −20°C. During plucking, some feathers of each bird were picked from different parts of the body to create a sample representative of the feathering of the body. Feather samples were then pooled by age and ground with a cutting mill (Grindomis GM200, Retsch, Newtown, PA). A subsample was further ground using an ultracentrifugal mill (ZM100, Retsch), whereas another subsample was ground with a ball mill (Vangoumil 300, Merck, Darmstadt, Germany) for amino acid analyses. The individual frozen FFB samples were cut into smaller pieces. For turkeys weighing less than 1 kg, this was done manually, whereas for turkeys weighing more than 1 kg, a cutter-slicer was used (RF15, Hobart, Marne la Vallée, France). The samples were then ground twice (4346SF, Hobart) using, respectively, a 5-mm and a 2-mm die. After manual homogenization, a 600-g sample was taken for heavier turkeys, whereas all material was used for turkeys weighing less than 1 kg. The FFB samples were weighed, freeze-dried, and weighed again. The samples were then minced and stored in a cold room. The freeze-dried FFB samples were later ground with an ultracentrifugal mill (ZM100, Retsch) for chemical analyses. Subsamples of these were taken, pooled by age, and ground with a ball mill (Vangoumil 300, Merck) for amino acid analyses. Chemical Analyses Chemical analyses were carried out in samples of individual turkeys for FFB and in samples pooled by age for feathers. Samples were analyzed for DM (ISO 64961983), ash (ISO-5984), N [NF V18-120, 1997 (with a LECO3000 instrument, Leco, St. Joseph, MI)], and fat (NF V18-117, 1997) using methods of the International Organization for Standardization (ISO methods, http://www.iso.org) and the AFNOR group (NF methods, http://www.afnor.org/en). The fat content was not determined in feathers. The DM content of FFB corresponded to the product of the DM contents determined during freeze-drying and oven-drying. Energy content was measured with an adiabatic bomb calorimeter (ISO 9831-1998; IKA C5000, IKA, Staufen, Germany). Amino acid analyses were performed by the Ajinomoto Eurolysine S.A.S. laboratory in Amiens (France) with a JLC-500/V AminoTac Amino Acid Analyzer (JEOL, Croissy-sur-Seine, France) after hydrolysis with 6 N HCl at 110°C for 23 h under reflux. The Cys and Met contents were obtained after performic acid oxidation before hydrolysis. The Trp content was determined after hydrolysis at 120°C for 16 h with barium hydroxide and separation by reverse-phase HPLC and fluorometric detection. Statistical Analyses All statistical analyses were performed using the GLM and NLIN procedures of SAS (SAS Inst. Inc., 70 Rivera-Torres et al. Cary, NC). A Gompertz growth function was used to describe the change in BW (BWt, in kg) as a function of age (t, in d). The function was log-transformed to account for heteroscedasticity of the errors: log (BWt) = log(BWm × exp{−exp[−B × (t − t*)]}), [1] where BWm is the BW at maturity (kg), B is the rate of maturing (d−1), and t* is the age at maximal growth rate (d). A high value of B implies an early-maturing animal. Body composition traits were analyzed by analyses of variance with age as a fixed effect. Traits were also analyzed using allometric equations to describe the development of a tissue relative to a reference compartment (e.g., empty BW). When described as a mass, the equation is Y = a × Xb, [2a] where Y is the mass of a tissue or a nutrient (g) and X is the mass of the reference compartment (g). The parameters a and b are the shape and scale parameters of the allometric equation, respectively. Because most of the traits were measured as contents (e.g., amino acid content in FFB protein), a transformed equation was used: Y/X = a × X(b−1). [2b] When the scale parameter b differs from unity, the content increases (b >1) or decreases (b <1) with increasing mass. The allometric analysis differs from the ANOVA in that mass (and not age) is seen as a driving force for potential changes in composition. Several authors have reported important changes in body composition during the first week after hatching (Leeson and Summers, 1980; Ferket et al., 1997). We also observed that body composition in the first week differed from that later on (see below). Therefore, the allometric analysis was carried out using data from 3 wk of age onward. The results of the Gompertz function and the allometric relations were used to determine the average composition of BW gain between 3 and 15 wk of age. In addition, the allometric relations were used to determine the average contribution of feathers to wholebody amino acid retention. The ash, water, and protein masses in FFB were described as a function of fat-free FFB mass, as indicated in equation 2a. RESULTS Daily gain was in line with the performance potential provided by the breeder and averaged 140 g/d between 1 and 15 wk of age. With the Gompertz function (equation 1), the estimated mature BW (BWm) was 18.1 ± 0.9 kg, whereas the rate of maturing (B) was 0.0274 ± 0.0009 /d. The age at maximal growth rate (t*) was estimated at 62 ± 2 d. The viscera and feather contents generally decreased with increasing age (Table 1), resulting in an increase in carcass content from 75.2 to 90.2% of BW between 1 and 15 wk. These tendencies are also reflected in the scale parameter (b) of equation 2b, which was greater than unity for the carcass and lower than unity for the viscera and feathers (Table 2). The allometric scalar for viscera was lower than that of feathers, indicating that the viscera content decreased more rapidly than the feather content. In general, the protein and ash contents in FFB declined with increasing age, whereas the lipid content, and consequently the gross energy content, increased substantially from 3 wk of age onward (Table 1). The composition of FFB at 1 wk of age appeared to be different from the general tendency observed at the other ages. The water content in particular was low (60.3%) at 1 wk of age, resulting in relatively high values for protein, ash, and lipid contents. This justifies the differentiation in body composition between 1 wk of age and later ages in the allometric analysis (Table 2). The changes in chemical composition of FFB were reflected in the allometric scalars (for FFB), which were lower than unity for the protein and water contents and were greater than unity for the lipid and gross energy contents. In addition, the scalars for water, ash, and protein were very similar (0.966 to 0.970), meaning that their ratios in FFB gain were relatively constant and that the composition of fat-free FFB would be constant during the growing period. That was further illustrated in the allometric relationships (Equation 2b) for water, ash, and protein content relative to the fat-free FFB mass, for which the allometric scalars did not differ significantly from unity. The fat-free FFB contained on average 71.6, 4.2, and 24.2% water, ash, and protein, respectively. Feathers were mostly composed of protein (Table 1) and the protein content did not change with feather mass (Table 2), resulting in an average protein content of 88.6% in feathers. Similarly, the gross energy content in the feathers did not vary during growth (P = 0.19) and corresponded to 20.64 kJ/g of feathers. The ash content decreased with increasing feather mass, whereas the water content increased (Table 2). The amino acid profile differed between FFB and feather protein (Tables 3 and 4). The Lys, Met, Trp, and His contents in FFB protein were at least 2-fold greater than those in feather protein, whereas the Cys content was 6-fold greater in feathers than in FFB protein. The variation in the amino acid contents of protein from 3 to 15 wk of age (from 0.8 to 14 kg of BW) appeared to be greater in feathers than in FFB, as indicated by the allometric scalars (Tables 3 and 4). The amino acid composition of FFB was relatively constant during growth and only the Cys and Ile contents were affected by FFB protein mass. In feathers, the Cys, Val, and Ser contents increased with increasing protein 71 BODY COMPOSITION IN MALE TURKEYS Table 1. Body composition of male turkeys from 1 to 15 wk of age1 Significance2 Age (wk) Item Empty BW (g/bird) Empty body (% of BW) Carcass3 Viscera Feathers Feather-free body (%) Water Ash Protein (N × 6.25) Lipid Gross energy (kJ/g) Feathers4 (%) Water Ash Protein (N × 6.25) Gross energy (kJ/g) 1 3 5 7 9 11 13 15 186 75.2 18.5 6.3 60.3 4.5 26.3 8.4 9.34 10.8 1.9 86.7 20.56 862 77.1 18.9 4.0 66.7 4.1 23.2 5.5 7.61 11.6 1.4 88.5 20.11 2,310 84.3 11.3 4.4 67.7 3.5 20.9 7.5 7.89 11.4 1.6 88.6 21.17 4,112 86.7 10.7 2.6 63.9 4.1 23.7 6.1 8.64 9.5 1.1 90.5 21.49 6,463 91.7 5.3 3.0 64.7 3.7 20.4 8.7 8.86 10.5 1.4 89.5 21.08 8,947 92.0 4.7 3.3 64.4 3.9 22.0 6.8 8.72 11.9 1.1 88.2 20.77 11,840 89.5 7.6 2.9 61.1 3.6 21.1 12.5 10.16 11.4 0.9 88.6 19.86 13,944 90.2 7.2 2.6 60.9 3.5 20.8 14.2 10.44 13.5 1.2 86.5 19.95 RSD P-value 754 4.3 4.1 1.2 1.8 0.3 1.1 2.7 0.69 NA NA NA NA <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 <0.01 NA NA NA NA 1Values correspond to the average of 4 replicate measurements per age (for feathers: 1 pooled measurement per age). = residual SD; NA = not applicable. P= probability of an age effect. 3Feather-free eviscerated body. 4No statistical analysis was carried out because there was no replicate measurement. 2RSD mass, whereas the Lys, Met, Trp, His, Tyr, Asp, and Glu contents decreased. Threonine, Leu, Phe, and Arg were the only amino acids that remained constant in both FFB and feather protein. The allometric relations were used to determine the average composition of BW gain (Table 2) and the average amino acid composition of whole-body protein retention (Figure 1). As expected, BW gain was mainly due to carcass gain, and viscera and feathers represented only a minor part of BW gain (4.7 and 2.6%, respectively). The FFB gain mostly consisted of water, and contained 21% protein, 13% lipid, and 4% ash. Feather gain mostly consisted of protein (87%). Lysine, Leu, and Arg were the most abundant essential amino acids in the deposited protein, each representing more than 6%. Most amino acids were deposited in FFB, but the contribution of FFB and feathers varied largely between amino acids. Feathers contributed only marginally (6%) to whole-body Lys and Met retention, but they represented 33% of whole-body Cys retention. Table 2. Allometric relations describing the change in body composition during growth in male turkeys from 3 to 15 wk of age Parameter estimate2 Item1 Empty body Carcass Viscera Feather-free body Feathers Feather-free body Water Ash Protein Lipid Gross energy Feathers Water Ash Protein Gross energy 1The a (SD) 0.539 (0.052) 3.687 (2.334) 0.915 (0.018) 0.120 (0.061) 0.848 (0.047) 0.050 (0.008) 0.281 (0.032) 0.003 (0.003) 3.36 (0.56) 0.089 (0.010) 0.020 (0.003) 0.913 (0.012) 21.6 (0.71) b (SD) RSD3 Composition of gain4 1.057* (0.011) 0.558* (0.082) 1.007* (0.002) 0.846* (0.062) 0.967* (0.007) 0.966 (0.020) 0.970* (0.013) 1.388* (0.102) 1.114* (0.02) 1.049* (0.021) 0.907* (0.032) 0.994 (0.003) 0.991 (0.006) 0.045 0.043 0.011 0.011 0.021 0.004 0.014 0.032 0.776 0.011 0.002 0.011 0.608 92.9% 4.5% 97.4% 2.6% 62.4% 3.6% 21.3% 12.7% 10.1 kJ/g 11.5% 1.1% 87.4% 20.4 kJ/g values for each subitem (Y) are expressed relative to that of the main item (X). the allometric relation Y/X = a·X(b−1), where a and b are the shape and the scale parameters, respectively, and both Y and X are expressed in grams (gross energy is expressed in kilojoules). Body composition was determined at 7 different ages with 4 replicate measurements per age (1 measurement per age for feathers). An asterisk (*) indicates the value is significantly different from 1 (P < 0.05). 3RSD = residual SD. 4Average composition of BW gain from 3 to 15 wk of age calculated from the allometric relations from 0.8 to 14 kg of BW. 2From 72 Rivera-Torres et al. Table 3. Amino acid composition (%) in feather-free body protein in male growing turkeys and allometric relations describing the change in amino acid composition during growth1 Parameter estimate2 Age (wk) Amino acid 1 3 5 7 9 11 13 15 Lys Thr Met Cys Trp Ile Leu Val His Phe Tyr Arg Gly Ser Ala Asp Glu Pro Total 6.82 3.92 1.93 1.10 1.08 3.88 6.96 4.57 2.42 3.81 3.15 6.68 8.23 4.05 6.24 8.29 13.05 5.19 91.4 6.94 3.83 2.04 1.05 1.09 3.87 6.83 4.46 2.60 3.65 2.99 6.42 7.76 3.80 6.15 8.20 12.87 5.02 89.6 7.23 3.92 2.04 1.02 1.07 3.99 7.01 4.55 2.67 3.73 3.07 6.52 7.69 3.89 6.26 8.43 13.21 5.11 91.4 6.95 3.81 2.02 0.99 0.99 3.96 6.80 4.50 2.57 3.63 2.93 6.52 8.02 3.73 6.23 8.25 12.97 5.31 90.2 7.13 3.83 2.00 0.96 1.04 4.07 6.92 4.65 2.69 3.67 2.96 6.55 7.98 3.64 6.38 8.38 13.26 5.24 91.4 7.10 3.76 2.02 0.98 1.03 4.02 6.80 4.52 2.63 3.56 2.92 6.39 7.63 3.56 6.21 8.23 13.07 4.98 89.4 7.19 3.79 2.01 0.97 1.04 4.07 6.84 4.55 2.62 3.59 2.89 6.43 7.68 3.66 6.21 8.27 13.16 5.09 90.1 7.31 3.86 2.04 0.94 1.04 4.07 6.93 4.51 2.65 3.62 2.95 6.50 7.65 3.69 6.26 8.38 13.42 5.10 90.9 a (SD) 0.059 0.041 0.021 0.017 0.014 0.031 0.069 0.042 0.025 0.041 0.036 0.065 0.084 0.051 0.059 0.080 0.114 0.050 (0.006) (0.004) (0.001) (0.001) (0.002) (0.002) (0.005) (0.003) (0.003) (0.003) (0.004) (0.004) (0.011) (0.007) (0.004) (0.006) (0.008) (0.008) b (SD) 1.012 (0.006) 0.996 (0.005) 0.997 (0.003) 0.968* (0.006) 0.982 (0.010) 1.015* (0.003) 1.000 (0.005) 1.004 (0.005) 1.003 (0.006) 0.992 (0.005) 0.988 (0.006) 1.000 (0.004) 0.995 (0.008) 0.980 (0.009) 1.004 (0.004) 1.002 (0.004) 1.009 (0.004) 1.001 (0.010) 1Amino acid composition was determined in samples pooled by age (4 turkeys per pool). the allometric relation 100·Y/X = a·X(b−1), where a and b are the shape and scale parameters, respectively, 100·Y/X is the amino acid content (%), and X (g) is the feather-free body protein mass. Only data between 3 and 15 wk of age were used in the analysis. An asterisk (*) indicates the value is significantly different from 1 (P < 0.05). 2From DISCUSSION Growth functions, such as the Gompertz function, are frequently used in biology to describe the change in BW as a function of time. Because of the simplicity and robustness of this function, the parameters may be compared among different studies. Emmans (1989) estimated the mature BW in different strains of turkeys from 1942 (Asmundson, 1942) to 1980 (Leeson and Summers, 1980), and he suggested that genetic selection for greater BW resulted in an increase in mature BW and in earlier maturing turkeys. This hypothesis was confirmed later by Anthony et al. (1991), who compared randombred turkeys and turkeys selected for increased BW at 16 wk of age. Although the estimated BW at maturity was close to the estimate by Emmans for 1980 data and to that of Anthony et al. (1991), the rate of maturing (B) was greater than those estimated Table 4. Amino acid composition (%) in feather protein in male growing turkeys and allometric relations describing the change in amino acid composition during growth1 Parameter estimate2 Age (wk) Amino acid Lys Thr Met Cys Trp Ile Leu Val His Phe Tyr Arg Gly Ser Ala Asp Glu Pro Total 1Amino 1 3 5 7 9 11 13 15 2.24 4.37 0.65 6.58 0.74 4.67 7.58 7.32 1.62 5.32 4.72 7.27 8.05 10.49 3.85 7.32 10.90 9.03 102.7 1.77 4.49 0.50 6.95 0.52 4.68 7.99 7.95 0.78 5.18 3.61 6.81 7.93 11.10 4.70 7.03 10.53 9.58 102.1 1.74 4.51 0.43 7.16 0.51 4.81 8.10 8.29 0.75 5.18 3.43 6.84 7.95 11.05 4.62 6.95 10.52 9.83 102.7 1.67 4.54 0.41 7.16 0.49 4.81 8.15 8.24 0.74 5.32 3.41 6.95 7.88 11.32 4.70 7.00 10.50 10.12 103.4 1.53 4.37 0.39 7.21 0.42 4.76 7.88 8.20 0.72 NA NA 6.84 7.64 11.05 4.43 6.77 10.00 9.89 92.1 1.52 4.54 0.43 7.27 0.42 5.03 8.21 8.37 0.74 5.33 3.13 7.09 7.79 11.83 4.36 6.70 10.35 10.11 103.2 1.29 4.34 0.37 7.69 0.43 4.86 8.21 8.47 0.65 5.31 3.10 6.97 8.03 11.90 4.59 6.73 10.02 9.87 102.8 1.25 4.43 0.34 7.43 0.38 4.86 8.28 8.48 0.66 5.32 3.02 6.95 8.14 11.79 4.61 6.70 9.98 10.16 102.8 a (SD) b (SD) 0.100 (0.047) 0.051 (0.006) 0.025 (0.011) 0.045 (0.006) 0.025 (0.010) 0.037 (0.004) 0.069 (0.008) 0.058 (0.004) 0.018 (0.005) 0.044 (0.003) 0.095 (0.010) 0.059 (0.005) 0.075 (0.012) 0.072 (0.012) 0.061 (0.011) 0.096 (0.006) 0.143 (0.017) 0.077 (0.009) 0.866* (0.034) 0.990 (0.009) 0.871* (0.032) 1.034* (0.010) 0.878* (0.028) 1.020 (0.008) 1.012 (0.008) 1.025* (0.005) 0.934* (0.020) 1.012 (0.005) 0.924* (0.007) 1.012 (0.006) 1.004 (0.011) 1.033* (0.012) 0.980 (0.013) 0.976* (0.004) 0.977* (0.009) 1.018 (0.008) acid composition was determined in samples pooled by age (4 turkeys per pool). NA = not applicable. the allometric relation 100·Y/X = a·X(b−1), where a and b are the shape and scale parameters, respectively, 100·Y/X is the amino acid content (%), and X (g) is the feather protein mass. Only data between 3 and 15 wk of age were used in the analysis. An asterisk (*) indicates the value is significantly different from 1 (P < 0.05). 2From BODY COMPOSITION IN MALE TURKEYS by Emmans (1989) and Anthony et al. (1991). Genetic selection resulted in early-maturing birds with a rapid growth rate. Although the allometric relation indicated that the viscera content in BW decreased with increasing body mass, viscera content was not a continuously declining fraction of BW because it decreased by 40% during the first 5 wk of age, but increased again after 11 wk of age (Table 1). Similar to our observations, Leeson and Summers (1980) and Hurwitz et al. (1983) observed a sharp decline in viscera content during the first 5 wk of age. At later ages, the viscera content in BW appeared to remain stable (Leeson and Summers, 1980) or even to increase (Hurwitz et al., 1983). Lilja (1983) and Hurwitz et al. (1991) suggested that this may be explained by an early development of internal organs, such as the liver and intestines, and a later development of other organs, such as the testes and spleen. The change in composition of FFB during growth observed in our study is in agreement with data in the literature. The relatively high protein and lipid contents at 1 wk of age in FFB were also observed by Leeson and Summers (1980) and Hurwitz et al. (1983). During the first week posthatch, the lipid content appeared to decrease because of the mobilization of fat reserves from the yolk sac. During this period, the poult is in a transition phase between using endogenous nutrients from the yolk sac to using dietary nutrients (Sklan et al., 1996; Noy and Sklan, 2001). Later on, the lipid content in FFB increased with age, as observed by Hurwitz et al. (1983). The lipid content in FFB at 15 wk of age was close to values observed by Crouch et al. (2002) in female breeder turkeys and those reported by Sell (1993) in male turkeys at a similar age. It is known that, apart from age, the diet can also affect the partitioning of energy retention between protein and lipids in broiler chickens and turkeys (Priyankarage et al., 2008). Defining body composition as a function of the fat-free FFB mass enables the body composition to be described independently of the protein-to-lipid ratio in the body. In our study, the water, ash, and protein contents represented a constant proportion of the fat-free FFB mass, which also means that water and ash are constant relative to protein (2.9 and 0.17 of the protein mass in FFB, respectively). Similarly, Eits et al. (2002) measured an ash-to-protein ratio of 0.15 in the FFB of broilers. However, Emmans (1989) and Eits et al. (2002) observed that the water-to-protein ratio in fatfree FFB decreased in turkeys and broiler chickens during growth. These differences may be due to changes in the relative contribution of carcass and viscera in FFB during growth. As observed by Eits et al. (2002), the constant ash-to-protein ratio in FFB may be due to the combination of an increasing ash-to-protein ratio in viscera and a decreasing water-to-protein ratio in carcass. Similarly, the constant water-to-protein ratio observed in our study may be due to a decreasing contribution of viscera in the feather-free protein mass and to an in- 73 creasing contribution of carcass, which has a low water content in comparison with viscera. Apart from Cys and Ile, the amino acid content in FFB was relatively constant during growth. Our values were close to those reported by Fisher and Scougall (1982) and Hurwitz et al. (1983), suggesting that genetic selection has not significantly affected the amino acid composition in FFB. Apart from the rapid change during the first week, the content of several amino acids in feather protein changed during growth, with an increase in Cys and Val and a decrease in Lys, Met, Trp, His, and Tyr. Similar changes in amino acid composition were observed by Fisher et al. (1981), specifically for His, the aromatic amino acids, and the sulfur-containing amino acids. These changes may be explained by a change in the nature of feathers, as suggested by Fisher and Scougall (1982) and Stilborn et al. (1997). Similar to the lipid and protein contents in FFB, the amino acid content in feather protein also changed very rapidly during the first wk of age. For example, there was a 2-fold decrease in the His content in feather protein from 1 to 3 wk of age. A similar reduction in His content was observed by Stilborn et al. (1997) in broilers. The origin of these changes is not entirely clear but may be related to the fact that, after hatching, the young poult can consume essential nutrients, some of which may be limiting during the in ovo stage. In ovo administration of amino acids in broiler breeder eggs has been shown to change the amino acid content of the embryo (Ohta et al., 2001). The changing contribution of feathers and FFB to whole-body protein during growth implies that the amino acid retention is not constant during the growth of turkeys. Although feather mass represents only a small part of BW (3 to 4%), the contribution of feather protein retention to total protein retention ranged from Figure 1. Average amino acid composition of whole-body protein gain and the contribution of feathers and feather-free body to the composition of whole-body protein gain in male growing turkeys from 3 to 15 wk of age (0.8 to 14 kg of BW). 74 Rivera-Torres et al. 13% at 0.8 kg of BW to 9% at 14 kg of BW. This, combined with the fact that feathers have a high Cys content, resulted in a reduction in the contribution of feathers to whole-body Cys retention (from 36% at 0.8 kg of BW to 31% at 14 kg of BW). The Cys-to-Lys retention ratio concomitantly declined from 31% at 0.8 kg of BW to 26% at 14 kg of BW. It is therefore likely that the Cys requirement (relative to Lys) decreases with increasing BW. The current study does not provide direct information concerning the amino acid requirement of turkeys. The requirement is determined by the amino acid composition of retained protein, the efficiency with which a limiting amino acid can be used for growth, and the maintenance amino acid requirement. The ideal amino acid profile is an aggregate representation of these components. Assuming that the ideal amino acid profile is constant implies that the relative contribution of these components (or of body tissues) does not change during growth. 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