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*Faculdade de Ciências Agrárias e Veterinárias, Universidade Estadual Paulista, Jaboticabal, São Paulo, Brazil;
and †Department of Poultry Science, North Carolina State University, Raleigh, North Carolina 27695
ABSTRACT Two experiments were conducted to de- live BW, the effects of environmental temperature, and
velop and evaluate a model to estimate ME requirements fractional fat (Gf) and protein (Gp) deposition. The second
and determine Gompertz growth parameters for broilers. experiment was carried out to estimate the growth param-
The first experiment was conducted to determine mainte- eters of Ross broilers and to collect data to evaluate the
nance energy requirements and the efficiencies of energy ME requirement model proposed. Live BW, empty
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utilization for fat and protein deposition. Maintenance feather-free carcass, weight of the feathers, and carcass
ME (MEm) requirements were estimated to be 157.8, 112.1, chemical compositions were analyzed until 16 wk of age.
Parameters of Gompertz curves for each component were
and 127.2 kcal of ME/kg0.75 per day for broilers at 13, 23,
estimated. Males had higher growth potential and higher
and 32°C, respectively. Environmental temperature (T)
capacity to deposit nutrients than females, except for fat
had a quadratic effect on maintenance requirements deposition. Data of BW and body composition collected
(MEm = 307.87 - 15.63T + 0.3105T2; r2= 0.93). Energy re- in this experiment were fitted into the energy model pro-
quirements for fat and protein deposition were estimated posed herein and the equations described by Emmans
to be 13.52 and 12.59 kcal of ME/g, respectively. Based (1989) and Chwalibog (1991). The daily ME requirements
on these coefficients, a model was developed to calculate estimated by the model determined in this study were
daily ME requirements: ME = BW0.75 (307.87 − 15.63T + closer to the ME intake observed in this trial compared
0.3105 T2) + 13.52 Gf + 12.59 Gp. This model considers with other models.
(Key words: factorial method, growth parameter, metabolizable energy requirement, modeling energy utilization)
2005 Poultry Science 84:1363–1369
1363
1364 SAKOMURA ET AL.
(Pasternak and Shalev, 1994; Hancock et al., 1995; Hruby The combined pieces were ground in a grinder3 2 times.
et al., 1996; Gous et al., 1999). These samples were dried in a drying oven (55°C) for
In order to determine ME requirements, the factorial 96 h and reground for homogeneity. Carcass and feather
method seems to satisfy the concept of physiological parti- samples were analyzed for nitrogen, fat, and dry matter.
tion of requirements into maintenance and growth (Hur- Chemical analyses followed the methods of AOAC (1990).
witz et al., 1983). The factorial method can be expressed Gross energy was determined by using an adiabatic
by this model: NI = Nm + Nl + Nf, where NI is the nutrient bomb calorimeter.4
intake, and Nm, Nl, and Nf are the requirements of nutrients The energy retention (ER) was calculated by the differ-
for maintenance, lean tissue, and fat deposition, respec- ence between final body energy content and initial energy
tively. The Nm depends on body and carcass composition. content. Heat production was calculated by the difference
The Nl is limited by the genetic potential, and NI and Nf between ME intake (MEi) and energy retained (ER).
are affected by environmental and genetic factors (Em- The MEm was estimated by the regression equation of
mans, 1986). ER as a function of MEi. The intercept on the X-axis of this
Estimation of coefficients not only helps to develop mod- equation represents the MEm for each temperature, and
els to estimate energy requirements but also facilitates the the slope represents the efficiency of energy utilization
understanding of energy metabolism. The present study above maintenance (kg). The linear regression of the loga-
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had the following objectives: To develop a model to esti- rithmic transformation of heat production values as a func-
mate daily energy requirements for broiler chickens; to tion of MEi was used to estimate heat production in the
provide information of growth parameters for the live BW, fasting state (i.e., the net energy requirement for mainte-
carcass, feather, and chemical composition of male and nance). The effect of temperature on MEm was determined
female Ross broilers; and to evaluate the ME model pro- by regression of MEm as a function of temperature.
posed herein. Multiple linear regression (MLR) was used to fractionate
MEi in energy retained as fat (ERf) and as protein (ERp):
MATERIALS AND METHODS MEi = MEm + 1/kf ERf + 1/kp ERp. This procedure also
determined the MEm requirement as being the intercept of
Two experiments were carried out. The first experiment equations, and kf and kp represent the efficiencies of energy
was conducted to develop a model to estimate the ME utilization for fat and protein deposition. MLR was also
requirement. The second experiment was conducted to used to partition the BW gain (BWG) into protein (PR) and
study growth parameters and body composition of broilers fat (FR) retention; BWG (g) = a + bPr + cFr, where b and
and to apply these data to evaluate the ME requirement c are coefficients that indicate rates of PR and FR. A model
model. was elaborated to estimate the daily ME requirements
based on the coefficients determined for maintenance and
Development of ME Requirement Model energy requirements for protein and fat deposition.
13°C
Ad libitum 395.7 ± 10.2 151.6 ± 7.4 244.1 ± 5.9
30% restriction 328.1 ± 2.1 102.6 ± 6.4 225.6 ± 5.0
50% restriction 273.8 ± 1.8 72.2 ± 8.5 201.6 ± 7.3
70% restriction 175.9 ± 2.6 12.7 ± 2.3 163.2 ± 4.8
23°C
Ad libitum 362.3 ± 13.8 150.4 ± 11.4 211.9 ± 20.5
30% restriction 295.2 ± 2.7 101.9 ± 0.9 193.3 ± 3.6
50% restriction 242.2 ± 0.4 81.3 ± 8.6 160.9 ± 8.8
70% restriction 151.6 ± 0.2 22.0 ± 8.4 129.6 ± 8.4
32°C
Ad libitum 305.8 ± 16.4 120.2 ± 10.4 185.7 ± 7.0
30% restriction 230.5 ± 1.9 63.1 ± 7.1 167.5 ± 8.6
50% restriction 179.4 ± 0.8 36.2 ± 3.5 143.1 ± 3.1
70% restriction 117.8 ± 0.3 −5.5 ± 2.6 123.3 ± 2.4
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fitted according to the Gompertz (1825) function: Wt = RESULTS
Wm.exp[− exp{− B( t − t* )}], where: Wt = weight (g) of the
animal or body component at time t, expressed as a func- Development of the Energy
tion of Wm; Wm = weight (g) at maturity of animal; B = Requirement Model
rate to maturity (per day); and t* = time (days) taken to
reach the maximum rate to maturity. Based on data of MEi, ER, and heat production observed
According to the equations obtained, the rates of growth (Table 1), the MEm, net energy requirement for mainte-
(g/d) were determined by derivation of the Gompertz nance, and efficiencies of energy utilization above mainte-
equations as a function of time t for BW, feather, and fat nance (kg) and for maintenance (km) were determined by
and protein deposition. Allometric relationships between linear regressions (Table 2). The MEm requirements were
body protein and the other chemical body components 157.8, 112.1, and 127.2 kcal of ME/kg of W0.75/d for chick-
were defined to verify whether the body components could ens reared at 13, 23, and 32°C, respectively. Temperature
be estimated as a function of protein weight. The allometric had a quadratic effect on MEm with a minimum MEm near
relationships were calculated by linear regression analysis. 26°C: MEm= W0.75 (307.87 + 15.63 T + 0.31 T2),. However,
Linear regression equations and the Gompertz equations a slight effect of temperature on km and kg was observed
were fitted by using the Statistica software (1996). in the present experiment.
The MLR equations presented in Table 3 describe the
Evaluation of Model Described partition of BWG into Pr and Fr. According to these equa-
tions, body protein deposition (3.66, 2.53, and 2.62 g) and
Data of BWG and body composition collected in this fat deposition (1.52, 2.32, and 1.52 g) at 13, 23, and 32°C
experiment were fitted into the energy model described were different, probably due to variation in feed intake
herein and into the equations described by Emmans (1989) according to temperature differences.
and Chwalibog (1991). The daily ME requirements esti- The MEm requirements obtained from multiple linear
mated were compared with the MEi observed in the first regression of the MEi as a function of ERf and ERp (Table
trial. 4) were: 132, 114, and 107 kcal of ME/kg of W0.75/d for
TABLE 2. Regression equations of energy retention (ER) and logarithm of heat production (ln HP)
as a function of energy intake (MEi). Values of energy requirement for maintenance
(MEm and NEm) and efficiencies of ME utilization at 13, 23, and 32°C are given
Requirements
(kcal/kg of W0.75 per d) Efficiencies
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less efficiently utilized for protein deposition at low tem- ficient had greater differences: 1.33 g of fat per gram of
perature. protein deposited for males and 1.51 g for females.
The ME requirement for each gram of fat and protein
deposition was calculated by considering the kf and kp Evaluation of ME Requirement Model
determined at the 3 temperatures and energy value for
fat (9.37 kcal/g) and for protein (5.66 kcal/g; Leeson and Growth data and body chemical composition collected
Summers, 2001). The ME requirements for broilers reared in this experiment were fitted into the model previously
at 13, 23, and 32°C were 10.21, 16.96, and 13.40 kcal of described. Estimations of ME requirement made by the
ME/g for fat and 15.85, 9.74, and 12.17 kcal of ME/g for model determined in experiment 1 were compared with
protein deposition. The average ME requirement was 13.52 models described by Emmans (1989) and Chwalibog (1991)
kcal/g for fat and 12.59 kcal/g for protein deposition. and the ME intake observed in experiment 2 (Table 8). It
Based on coefficients for maintenance, fat, and protein was verified that the composition of the BW gain influences
gain, the following model was determined: ME = W0.75 the ME requirements. It is important to consider this differ-
(307.87 − 15.63T + 0.3105 T2) + 13.52 Gf + 12.59 Gp, where ential growth to estimate accurate values. Our model pre-
ME is the daily requirement per bird per day, W is live dicted ME requirements closer to the ME intake observed
BW, T is environmental temperature, and Gf and Gp are in the trial. The sum square error (SSE) of this model was
fat and protein deposition, respectively (g/bird per d). lower compared with the models of Emmans (1989) and
Chwalibog (1991). These models underestimated the ME
Description of Body, Feather, requirements compared with ME intake observed.
and Body Chemical Growth
DISCUSSION
The data in Table 5 show the estimated growth parame-
ters of male and female Ross broiler chickens described by The MEm requirements determined herein are in the
Gompertz function. The potential growth is described in ranges of values estimated by Nieto et al. (1995) for male
terms of live BW, feather-free empty weight, protein, water, broiler chickens (111.4 to 143 kcal/kg of W0.75 per d). The
lipid, and ash BW. The growth rates of whole BW, feathers, quadratic effect of temperature on MEm observed in this
and protein and fat deposition are presented in Table 6. work is in agreement with the results of Hurwitz et al.
The growth parameters for live BW show the superiority (1980), who observed a reduction in the MEm requirement
of the males’ growth potential compared with the females, of broilers reared at 27°C and increasing needs up to 34°C.
TABLE 4. Multiple linear regression equations of MEi as a function of energy retention as fat (ERf) and
as protein (ERp). Estimates of MEm, efficiencies of energy retention as fat (kf) and protein (kp), and ME
requirements for fat (MEf) and protein (MEp) deposition in broilers at different temperatures
Parameters
1
Variables Sex Wm (g) B (per d)2 t* (d)3 r2
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1
Wm (g) = weight at maturity.
2
B (per d) = rate of maturing.
3
t* (d) = time at which growth rate is maximized.
These data confirm the physiological explanation that birds According to Kessler and Brugalli (1999), broilers reared
reared above or under their thermoneutral temperatures under ideal temperature conditions have a maximum utili-
need to change their metabolism to dissipate heat or in- zation of energy intake for lean tissue deposition. At low
crease heat production. temperatures, birds increase feed intake and body heat
The efficiencies of energy utilization for maintenance production, and the excess of nutrients ingested promotes
and above maintenance determined in the present experi- lipogenesis. At high temperatures, birds decrease energy
ment are similar to those estimated by Sakomura et al. intake to minimize the heat increment and promote an
(2003) for broiler breeder pullets (75, 76, and 72% for km intermediate deposition for lean tissue and fat. Reported
and 69, 69, and 62% for kg at 15, 22, and 30°C, respectively). values of kf and kp have high variability due to several
Balnave (1974) described variability in the efficiency for factors, such as genetics, sex, age, diet, and methods used
maintenance ranging between 66 and 78%. Efficiency for for their determination. Macleod (1990) estimated kf and
ME utilization above maintenance has been cited between kp for female broilers to be between 102 and 103% and
52 and 78% (Nieto et al., 1995). According to De Groote between and 47 and 57%, respectively. In contrast, Nieto
(1974) and Nieto et al. (1995), this variability in efficiencies et al. (1995) determined the value of kf for male broilers to
could be related with composition of the diets. be between 64 and 127% and kp to be between 40 and 58%.
The feeding level affects body composition by influenc- The comparisons between ME requirements for growth
ing the lipogenesis activity (Leeson and Summers, 1980; should be made with caution, because of variations in
Kessler and Brugalli, 1999). Boekholt et al. (1994) found the body composition, growth rate, and protein and fat
that Hybro broiler chickens had daily body protein and deposition rates according to the genetics, age, and envi-
fat retention coefficients of 3.8 g and 1.1 g, respectively. In ronment where birds are raised.
the present study, average deposition coefficients were 2.94 The ME requirements for protein and fat deposition ob-
g of protein and 1.79 g of fat retention per day (Table 3). tained in this work are similar to values determined with
TABLE 6. Growth rates1 of live BW, feathers, protein, and fat deposition according
to age and sex of Ross broilers
Age (d) Males Females Males Females Males Females Males Females
(g/d)
7 24.7 24.9 4.1 2.3 1.3 2.1 0.56 0.88
14 43.5 38.2 7.1 4.9 3.6 3.8 2.03 2.12
21 62.8 50.4 9.6 13.5 7.1 6.0 3.90 3.30
28 77.7 59.1 10.8 14.1 10.6 8.4 4.99 3.86
35 85.6 63.2 10.7 9.6 13.1 10.6 4.95 3.72
42 86.3 63.0 9.6 5.3 13.8 12.3 4.17 3.16
49 81.4 59.4 8.1 2.6 13.2 13.3 3.17 2.47
56 72.9 53.6 6.5 1.3 11.7 13.6 2.25 1.83
63 62.8 46.8 5.1 0.60 9.8 13.4 1.54 1.30
70 52.6 39.9 3.9 0.28 7.9 12.6 1.02 0.90
1
Growth rates were calculated from derivates of Gompertz equations.
1368 SAKOMURA ET AL.
TABLE 7. Allometric relationships of body fat (BWf), water (BWw),
and ash (BWa) with body protein (BWp)
Chemical
fraction1 Males r2 Females r2
BWf Log BWf = 1.022 + 1.334 log BWp 0.97 Log BWf = −1.288 + 1.506 log BWp 0.93
BWw Log BWw = 0.585 + 0.988 log BWp 0.97 Log BWw = 0.613 + 0.973 log BWp 0.96
BWa Log BWa = −1.002 + 1.080 log BWp 0.98 Log BWa = −1.011 + 1.085 log BWp 0.95
1
Chemical composition in grams.
turkeys by Emmans (1989), which were 13.38 kcal/g for occurred at 39 d for males and at 38 d for female broilers.
fat and 11.95 kcal/g of protein deposited. In the same way, The differences between males and females are noticeable
Boekholt et al. (1994) estimated that broilers need 10.90 after 21 d. Males gained more weight as they aged, and
and 8.57 kcal of ME/g for fat and protein deposition, re- the difference between males and females became more
spectively. significant.
The growth potential has been described by the Gomp-
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As a bird develops, the proportion of feather protein in
ertz (1825) function in several studies (Emmans, 1981, 1989; total protein gain changes, which must be taken into ac-
Knı́zetová et al., 1991; Hancock et al., 1995; Gous et al., count in calculation of amino acid requirements (Emmans,
1999). The growth parameters estimated in the present 1989). According to Gous et al. (1999), a description of
study are in the same range of live weight (5,800 to 6,100 mature feathering and growth rates is necessary to predict
g for males and 4,300 to 5,200 for females) presented by potential feather growth. In this study, these parameters
Knı́zetová et al. (1991), Hancock et al. (1995), and Gous et were higher than those described by Gous et al. (1999),
al. (1999). Values of rate to maturity (B) for live BW are who found average values of Wm for feather growth of
similar to those estimated by Gous et al. (1999) of 0.037
300 and 230 g, values for B of 0.040 and 0.042, and values
and 0.038 and superior to those estimated by Hancock et
for t* of 43 and 40 d for males and females, respectively.
al. (1995) and Knı́zetová et al. (1991), which changed from
The feathering of birds seems to be related to the genetic
0.033 to 0.037. The Wm values of empty feather-free weight
line and mainly to sex. Females are selected for earlier
obtained in this study were higher for males (5,939 g) and
lower for females (4,457 g) than those presented by Gous feathering for sex differentiation (Hancock et al., 1995).
et al. (1999), which were 5,500 to 5,700 g and 4,500 to 4,800 Gous et al. (1999) indicated that the feathering characteris-
g for males and females, respectively. tics could be modified over time as a consequence of a
One discrepancy was observed between the estimated selection process.
values in the present work and the previous reports. The Our results show that the growth rates for feathers were
t* values obtained in this experiment show the precocity initially higher for females, but the males had higher rates
of modern chickens. This parameter corresponds to the after 21 d until 70 d of age. Kessler et al. (2000) observed
inflexion point of the growth curve. Gous et al. (1999) and that females exhibited higher growth rates from 21 to 28
Hancock et al. (1995) estimated maximum growth rate (t*) d of age and lower growth rates after 35 d. The males had
at ages from 40 to 43 d and 43 to 45 d for males and females, higher rates between 28 and 35 d and remained constant
respectively. In the present experiment, the inflexion point until the age at processing.
ME model
Age MEi1 present Emmans Chwalibog
(d) (kcal/d) study2 P − O3 (1994)4 P − O3 (1991)5 P − O3
Downloaded from http://ps.oxfordjournals.org/ at Cleveland Health Sciences Library on May 11, 2014
indicates that males develop more meat and better carcass 1995. The evaluation of the growth parameters of six strains
composition (Kessler et al., 2000). The reason for the sig- of commercial broiler chickens. Br. Poult. Sci. 36:247–264.
nificant increments of fat deposition in females after 56 d Hruby, M., M. L. Hamre, and C. N. Coon. 1996. Non-linear and
is the bird’s need for extra fat deposition during develop- linear functions in body protein growth. J. Appl. Poult. Res.
5:109–115.
ment of the reproductive system (Gous et al., 1999). Hurwitz, S., I. Plavinik, I. Bengal, H. Talpaz, and I. Bartov. 1983.
Knowledge of an adequate description of body nutrient The amino acid requirements of growing turkeys. 1—Model
deposition and ME models to determine accurate require- construction and parameter estimation. Poult. Sci. 62:2208–
ments for broiler chickens can help to establish specific 2217.
feeding programs and define optimum age for processing, Hurwitz, S., M. Weiselberg, and U. T. Eisner. 1980. The energy
to provide better carcass quality, and reduce production requirements and performance of growing chickens and tur-
keys as affected by environmental temperature. Poult. Sci.
costs. The model proposed in this study includes the effect 59:2121–2128.
of environmental temperature, which affects body compo- Kessler, A. M., and I. Brugalli. 1999. Recentes avanços do efeito
sition and efficiencies of energy utilization for protein and da nutrição no crescimento especı́fico de componentes da car-
fat deposition. This factor improves the accuracy for pre- caça de frangos de corte. Pages 1–19 in Proceedings of the
dicting daily ME requirements. Simpósio Internacional Sobre Tecnologia de Processamento e
Qualidade de Carne em Aves, Concórdia, SC, Brazil.
Kessler, A. M., P. N. Snizek, and I. Brugalli. 2000. Manipulação
ACKNOWLEDGMENTS da quantidade de gordura na carcaça de frangos. Pages 107–
133 in Proceedings of the Conferência Apinco de Ciência e
The authors thank the Fundação de Amparo a Pesquisa
Tecnologia Avı́colas. FACTA, Campinas, SP, Brazil.
do Estado de São Paulo (FAPESP) for the financial support. Knı́zetová, H., J. Hyanek, B. Knize, and J. Roubicek. 1991. Analysis
of growth curves of fowl. I. Chickens. Br. Poult. Sci.
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