Modeling Energy Utilization and Growth Parameter
Description for Broiler Chickens1
N. K. Sakomura,*,2 F. A. Longo,* E. O. Oviedo-Rondon,† C. Boa-Viagem,* and A. Ferraudo*
*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-
velop and evaluate a model to estimate ME requirements
and determine Gompertz growth parameters for broilers.
The first experiment was conducted to determine mainte-
nance energy requirements and the efficiencies of energy
utilization for fat and protein deposition. Maintenance
ME (MEm) requirements were estimated to be 157.8, 112.1,
and 127.2 kcal of ME/kg0.75 per day for broilers at 13, 23,
and 32°C, respectively. Environmental temperature (T)
had a quadratic effect on maintenance requirements
(MEm = 307.87 - 15.63T + 0.3105T2; r2= 0.93). Energy re-
quirements for fat and protein deposition were estimated
to be 13.52 and 12.59 kcal of ME/g, respectively. Based
on these coefficients, a model was developed to calculate
daily ME requirements: ME = BW0.75 (307.87 − 15.63T +
0.3105 T2) + 13.52 Gf + 12.59 Gp. This model considers
(Key words: factorial method, growth parameter, metabolizable energy requirement, modeling energy utilization)
INTRODUCTION
Genetic selection of broilers has caused changes in per-
formance and rates of fat and protein deposition in the
carcass. These changes in growth patterns cause modifica-
tions to the nutrient requirements to express the maximum
genetic potential. Better knowledge of energy requirements
and actual efficiencies of energy utilization for protein and
fat deposition are very important to formulate diets that
promote reduction in body fat deposition.
Successful nutrient requirement determination requires
an adequate description of bird growth and body composi-
tion by using growth curves (Hruby et al., 1996). According
to Emmans (1981), one rational way to calculate nutrient
requirements and predict the feed intake of growing broil-
ers needs to start with the understanding of the genetic
2005 Poultry Science Association, Inc.
Received for publication August 19, 2004.
Accepted for publication May 5, 2005.
1
Funding for this research was provided by the Fundação de Amparo
a Pesquisa do Estado de São Paulo, São Paulo, Brasil.
2
To whom correspondence should be addressed: sakomura@fcav.
unesp.br.
1363
live BW, the effects of environmental temperature, and
fractional fat (Gf) and protein (Gp) deposition. The second
experiment was carried out to estimate the growth param-
eters of Ross broilers and to collect data to evaluate the
ME requirement model proposed. Live BW, empty
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feather-free carcass, weight of the feathers, and carcass
chemical compositions were analyzed until 16 wk of age.
Parameters of Gompertz curves for each component were
estimated. Males had higher growth potential and higher
capacity to deposit nutrients than females, except for fat
deposition. Data of BW and body composition collected
in this experiment were fitted into the energy model pro-
posed herein and the equations described by Emmans
(1989) and Chwalibog (1991). The daily ME requirements
estimated by the model determined in this study were
closer to the ME intake observed in this trial compared
with other models.
2005 Poultry Science 84:1363–1369
potential. Leeson and Summers (1980) confirmed that the
nutrient requirements and the growth rate of animals are
very closely related. Gous et al. (1999) commented that
each type of animal has a growth curve that needs to
be determined in ideal environmental conditions. Genetic
lines and genders can differ in several aspects, such as
weight at maturity, composition, and proportions of chemi-
cal components, which affect characteristics of the growth
curve. For adequate description of growth and adequate
estimation of energy and amino acids needs, it is necessary
to distinguish between feather growth and growth of the
empty body, due to significant differences in the amino
acid composition of these two tissues (Emmans, 1989).
In several studies about genetic potential, researchers
have utilized the mathematical function described by
Gompertz (1825). This function has been recently used to
describe several characteristics of growth patterns in birds
Abbreviation Key: kg = efficiency of energy utilization for retention;
kp = efficiency of energy utilization for protein retention; kf = efficiency
of energy utilization for fat retention; km= efficiency of energy utilization
for maintenance; MEi = metabolizable energy intake; MEm = metaboliz-
able energy requirement for maintenance; NEm = net energy require-
ment for maintenance; RE = retained energy; T = temperature.
1364 SAKOMURA ET AL.
(Pasternak and Shalev, 1994; Hancock et al., 1995; Hruby
et al., 1996; Gous et al., 1999).
In order to determine ME requirements, the factorial
method seems to satisfy the concept of physiological parti-
tion of requirements into maintenance and growth (Hur-
witz et al., 1983). The factorial method can be expressed
by this model: NI = Nm + Nl + Nf, where NI is the nutrient
intake, and Nm, Nl, and Nf are the requirements of nutrients
for maintenance, lean tissue, and fat deposition, respec-
tively. The Nm depends on body and carcass composition.
The Nl is limited by the genetic potential, and NI and Nf
are affected by environmental and genetic factors (Em-
mans, 1986).
Estimation of coefficients not only helps to develop mod-
els to estimate energy requirements but also facilitates the
understanding of energy metabolism. The present study
had the following objectives: To develop a model to esti-
mate daily energy requirements for broiler chickens; to
provide information of growth parameters for the live BW,
carcass, feather, and chemical composition of male and
female Ross broilers; and to evaluate the ME model pro-
posed herein.
MATERIALS AND METHODS
Two experiments were carried out. The first experiment
was conducted to develop a model to estimate the ME
requirement. The second experiment was conducted to
study growth parameters and body composition of broilers
and to apply these data to evaluate the ME requirement
model.
Development of ME Requirement Model
The experiment was conducted in 3 environmentally
controlled rooms in order to evaluate the effect of tempera-
ture on maintenance energy requirement (MEm) and effi-
ciencies of energy utilization for protein and fat deposition
in chickens by the comparative slaughter technique (Blax-
ter, 1989). Four hundred eighty 14-d-old Ross males were
randomly placed in floor pens (0.1 m2/bird) and assigned
to 1 of 3 rooms with temperatures of 13, 23, and 32 ±
2°C. Four feeding treatments in each room consisted of ad
libitum and 30, 50, and 70% restriction of the ad libitum
intake. Each treatment was replicated 4 times with 10 birds
per replicate. Diets were formulated according to Ross
nutritional recommendations (Agroceres Ross, 2000).
The control group of 12 birds was slaughtered at the
beginning of the experiment (21 d), and at the end (49 d
of age) all birds were processed. Broilers were killed by
CO2 asphyxiation and were weighed before and after
plucking to determine the weight of feathers. Samples of
feathers were taken and ground for subsequent analysis.
The carcasses were frozen at −4°C. The frozen birds from
each pen were combined and chopped into small pieces.
3
Model 114, CAF Rio Claro, SP, Brazil.
4
Reichel & Partner GmbH, Rheinzabern Germany.
The combined pieces were ground in a grinder3 2 times.
These samples were dried in a drying oven (55°C) for
96 h and reground for homogeneity. Carcass and feather
samples were analyzed for nitrogen, fat, and dry matter.
Chemical analyses followed the methods of AOAC (1990).
Gross energy was determined by using an adiabatic
bomb calorimeter.4
The energy retention (ER) was calculated by the differ-
ence between final body energy content and initial energy
content. Heat production was calculated by the difference
between ME intake (MEi) and energy retained (ER).
The MEm was estimated by the regression equation of
ER as a function of MEi. The intercept on the X-axis of this
equation represents the MEm for each temperature, and
the slope represents the efficiency of energy utilization
above maintenance (kg). The linear regression of the loga-
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rithmic transformation of heat production values as a func-
tion of MEi was used to estimate heat production in the
fasting state (i.e., the net energy requirement for mainte-
nance). The effect of temperature on MEm was determined
by regression of MEm as a function of temperature.
Multiple linear regression (MLR) was used to fractionate
MEi in energy retained as fat (ERf) and as protein (ERp):
MEi = MEm + 1/kf ERf + 1/kp ERp. This procedure also
determined the MEm requirement as being the intercept of
equations, and kf and kp represent the efficiencies of energy
utilization for fat and protein deposition. MLR was also
used to partition the BW gain (BWG) into protein (PR) and
fat (FR) retention; BWG (g) = a + bPr + cFr, where b and
c are coefficients that indicate rates of PR and FR. A model
was elaborated to estimate the daily ME requirements
based on the coefficients determined for maintenance and
energy requirements for protein and fat deposition.
Estimation of Parameters
to Describe Growth
A total of 480 Ross broilers were raised from 1 to 112 d
of age in an open-sided house. Two hundred forty chicks
of each sex were distributed in 4 replicates of 60 males and
60 females per pen. The maximum and minimum in-house
temperatures recorded during the 16-wk experimental pe-
riod ranged from 30 to 33.7°C and from 16.6 to 22.6°C,
respectively. Broilers were fed 4 diets formulated according
to Ross guide recommendations (Agroceres Ross, 2000).
The 4 corn-soybean meal diets contained 3,050, 3,180, 3,230,
and 3,180 kcal of ME/kg; 21.5, 19.0, 17.5, and 17.5% CP;
1.22, 1.06, 0.94, and 0.94% lysine; and 0.95, 0.85, 0.72, and
0.72% TSAA, respectively. These diets were fed in the fol-
lowing growing periods from 1 to 21 d, 22 to 42 d, 43 to
63 d, and 64 to 112 d of age.
Every week, a sample of at least 3 birds was selected
from each pen based on average BW of the pen. The birds
were slaughtered and processed as described in the first
experiment. Carcass and feather samples were analyzed
for nitrogen, fat, ash, and dry matter as described in the
previous experiment. The live BW, feather-free BW (with-
out viscera), feather weight, and weight of each chemical
body component were obtained weekly, and data were
 ENERGY MODEL FOR BROILER CHICKENS 1365
TABLE 1. Average metabolizable energy intake (MEi), body energy retention (ER),
and heat production (HP) at different room temperatures and feeding levels

Feeding level MEi ER HP

and temperature (kcal/kg0.75 per d) (kcal/kg0.75 per d) (kcal/kg0.75 per d)

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

fitted according to the Gompertz (1825) function: Wt =
Wm.exp[− exp{− B( t − t* )}], where: Wt = weight (g) of the
animal or body component at time t, expressed as a func-
tion of Wm; Wm = weight (g) at maturity of animal; B =
rate to maturity (per day); and t* = time (days) taken to
reach the maximum rate to maturity.
According to the equations obtained, the rates of growth
(g/d) were determined by derivation of the Gompertz
equations as a function of time t for BW, feather, and fat
and protein deposition. Allometric relationships between
body protein and the other chemical body components
were defined to verify whether the body components could
be estimated as a function of protein weight. The allometric
relationships were calculated by linear regression analysis.
Linear regression equations and the Gompertz equations
were fitted by using the Statistica software (1996).
Evaluation of Model Described
Data of BWG and body composition collected in this
experiment were fitted into the energy model described
herein and into the equations described by Emmans (1989)
and Chwalibog (1991). The daily ME requirements esti-
mated were compared with the MEi observed in the first
trial.
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RESULTS
Development of the Energy
Requirement Model
Based on data of MEi, ER, and heat production observed
(Table 1), the MEm, net energy requirement for mainte-
nance, and efficiencies of energy utilization above mainte-
nance (kg) and for maintenance (km) were determined by
linear regressions (Table 2). The MEm requirements were
157.8, 112.1, and 127.2 kcal of ME/kg of W0.75/d for chick-
ens reared at 13, 23, and 32°C, respectively. Temperature
had a quadratic effect on MEm with a minimum MEm near
26°C: MEm= W0.75 (307.87 + 15.63 T + 0.31 T2),. However,
a slight effect of temperature on km and kg was observed
in the present experiment.
The MLR equations presented in Table 3 describe the
partition of BWG into Pr and Fr. According to these equa-
tions, body protein deposition (3.66, 2.53, and 2.62 g) and
fat deposition (1.52, 2.32, and 1.52 g) at 13, 23, and 32°C
were different, probably due to variation in feed intake
according to temperature differences.
The MEm requirements obtained from multiple linear
regression of the MEi as a function of ERf and ERp (Table
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

Temperature Regression equations1 r2 MEm NEm kg2 km3

13°C ER = −98.69 + 0.63 MEi 0.99 157.83 0.60

ln HP = 4.78 + 0.00187 MEi 0.91 119.3 0.76
23°C ER= −66.1 + 0.59 MEi 0.96 112.09 0.57
ln HP = 4.50 + 0.0024 MEi 0.93 89.99 0.80
32°C ER = −83.74 + 0.66 MEi 0.99 127.16 0.64
ln HP = 4.57 + 0.0022 MEi 0.96 96.25 0.76
1
Slopes were significantly different from zero (P < 0.01).
2
kg = efficiency of ME utilization above maintenance.
3
km = efficiency of ME utilization for maintenance (NEm/MEm).
1366 SAKOMURA ET AL.
TABLE 3. Multiple linear regression equations of BW gain indicating higher weight at maturity (Wm) and higher rates
(BWG; g) as a function of protein (PR) and fat (FR) retention
to maturity (B), and resulting in similar age of maximum
Temperature Regression equations1 R2 growth (t*) between both sexes (Tables 5 and 6). The param-
13°C BWG = 3.38 + 3.66 PR + 1.52 FR 0.98 eters of feather-free BW and feather growth had the same
23°C BWG = 6.19 + 2.53 PR + 2.32 FR 0.93 tendency observed for live BW.
32°C BWG = 6.28 + 2.62 PR + 1.52 FR 0.94 Males exhibited higher potential for nutrient deposition
1
Slopes were significantly different from zero (P < 0.01). Slope coeffi- than females, except for fat deposition. Values (Table 5) of
cients correspond to the rates of PR and FR. rate to maturity (B) for protein, water, and ash deposition
were higher for females, resulting in lower t* values. In
contrast, the Wm and t* values for fat deposition in females
chickens kept at 13, 23, and 32°C, respectively. The effi- were higher and the B value was lower, which was consis-
ciency of energy utilization for fat (0.92) at 13°C was higher tent with the higher fat content of the older female broilers.
than those at 23°C (0.55) and at 32°C (0.70). On the other The allometric relationships (Table 7) determined by lin-
hand, the kp (0.36) at 13°C was lower than those at 23°C ear regression of BW fat (BWf), water (BWw), and ash (BWa)
(0.58) and at 32°C (0.47). This result indicates that dietary on body protein weight (BWp) showed the differences in
energy was more efficiently utilized for fat deposition and body composition between males and females. The fat coef-

less efficiently utilized for protein deposition at low tem-
perature.
The ME requirement for each gram of fat and protein
deposition was calculated by considering the kf and kp
determined at the 3 temperatures and energy value for
fat (9.37 kcal/g) and for protein (5.66 kcal/g; Leeson and
Summers, 2001). The ME requirements for broilers reared
at 13, 23, and 32°C were 10.21, 16.96, and 13.40 kcal of
ME/g for fat and 15.85, 9.74, and 12.17 kcal of ME/g for
protein deposition. The average ME requirement was 13.52
kcal/g for fat and 12.59 kcal/g for protein deposition.
Based on coefficients for maintenance, fat, and protein
gain, the following model was determined: ME = W0.75
(307.87 − 15.63T + 0.3105 T2) + 13.52 Gf + 12.59 Gp, where
ME is the daily requirement per bird per day, W is live
BW, T is environmental temperature, and Gf and Gp are
fat and protein deposition, respectively (g/bird per d).
Description of Body, Feather,
and Body Chemical Growth
The data in Table 5 show the estimated growth parame-
ters of male and female Ross broiler chickens described by
Gompertz function. The potential growth is described in
terms of live BW, feather-free empty weight, protein, water,
lipid, and ash BW. The growth rates of whole BW, feathers,
and protein and fat deposition are presented in Table 6.
The growth parameters for live BW show the superiority
of the males’ growth potential compared with the females,
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ficient had greater differences: 1.33 g of fat per gram of
protein deposited for males and 1.51 g for females.
Evaluation of ME Requirement Model
Growth data and body chemical composition collected
in this experiment were fitted into the model previously
described. Estimations of ME requirement made by the
model determined in experiment 1 were compared with
models described by Emmans (1989) and Chwalibog (1991)
and the ME intake observed in experiment 2 (Table 8). It
was verified that the composition of the BW gain influences
the ME requirements. It is important to consider this differ-
ential growth to estimate accurate values. Our model pre-
dicted ME requirements closer to the ME intake observed
in the trial. The sum square error (SSE) of this model was
lower compared with the models of Emmans (1989) and
Chwalibog (1991). These models underestimated the ME
requirements compared with ME intake observed.
DISCUSSION
The MEm requirements determined herein are in the
ranges of values estimated by Nieto et al. (1995) for male
broiler chickens (111.4 to 143 kcal/kg of W0.75 per d). The
quadratic effect of temperature on MEm observed in this
work is in agreement with the results of Hurwitz et al.
(1980), who observed a reduction in the MEm requirement
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

Temperature Regression equations1 MEm kf kp MEf2 MEp3

(kcal/kg0.75 per d) (kcal/g)

13°C MEi = 132.02 + 1.09. ERf + 2.80 ERp 132 0.92 0.36 10.21 15.85
23°C MEi = 114.13 + 1.81. ERf + 1.72 ERp 114 0.55 0.58 16.96 9.74
32°C MEi = 107.09 + 1.43. ERf + 2.15 ERp 107 0.70 0.47 13.40 12.17
Average 13.52 12.59
1
Slopes were significantly different from zero (P < 0.01), R2 = 0.96, 0.87, and 0.97, respectively, for regression
equations at 13, 23, and 32°C.
2
ME requirements calculated based on energy of fat (9.37 kcal/g) and kf.
3
ME requirements calculated based on energy of protein (5.66 kcal/g) and kp.
 ENERGY MODEL FOR BROILER CHICKENS 1367
TABLE 5. Estimates of growth parameters for live weight, feather-free BW, protein, fat, water,
and ash weights of male and female Ross broilers by using the Gompertz equation

Parameters
1
Variables Sex Wm (g) B (per d)2 t* (d)3 r2

Live weight Males 6,050 0.0389 39 0.99

Females 4,801 0.0362 38 0.99
Feather-free BW Males 5,939 0.0356 42 0.99
Females 4,457 0.0349 39 0.98
Feather weight Males 213 0.0653 31 0.90
Females 180 0.0588 30 0.89
Body protein (BWp) Males 616 0.0483 31 0.87
Females 368 0.1092 25 0.78
Body fat (BWf) Males 818 0.0455 42 0.93
Females 1,234 0.0297 56 0.93
Body water (BWw) Males 3,773 0.0327 43 0.98
Females 2,313 0.0379 34 0.97
Body ash (BWa) Males 201 0.0283 50 0.96
Females 127 0.0320 42 0.95

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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
reared above or under their thermoneutral temperatures
need to change their metabolism to dissipate heat or in-
crease heat production.
The efficiencies of energy utilization for maintenance
and above maintenance determined in the present experi-
ment are similar to those estimated by Sakomura et al.
(2003) for broiler breeder pullets (75, 76, and 72% for km
and 69, 69, and 62% for kg at 15, 22, and 30°C, respectively).
Balnave (1974) described variability in the efficiency for
maintenance ranging between 66 and 78%. Efficiency for
ME utilization above maintenance has been cited between
52 and 78% (Nieto et al., 1995). According to De Groote
(1974) and Nieto et al. (1995), this variability in efficiencies
could be related with composition of the diets.
The feeding level affects body composition by influenc-
ing the lipogenesis activity (Leeson and Summers, 1980;
Kessler and Brugalli, 1999). Boekholt et al. (1994) found
that Hybro broiler chickens had daily body protein and
fat retention coefficients of 3.8 g and 1.1 g, respectively. In
the present study, average deposition coefficients were 2.94
g of protein and 1.79 g of fat retention per day (Table 3).
According to Kessler and Brugalli (1999), broilers reared
under ideal temperature conditions have a maximum utili-
zation of energy intake for lean tissue deposition. At low
temperatures, birds increase feed intake and body heat
production, and the excess of nutrients ingested promotes
lipogenesis. At high temperatures, birds decrease energy
intake to minimize the heat increment and promote an
intermediate deposition for lean tissue and fat. Reported
values of kf and kp have high variability due to several
factors, such as genetics, sex, age, diet, and methods used
for their determination. Macleod (1990) estimated kf and
kp for female broilers to be between 102 and 103% and
between and 47 and 57%, respectively. In contrast, Nieto
et al. (1995) determined the value of kf for male broilers to
be between 64 and 127% and kp to be between 40 and 58%.
The comparisons between ME requirements for growth
should be made with caution, because of variations in
the body composition, growth rate, and protein and fat
deposition rates according to the genetics, age, and envi-
ronment where birds are raised.
The ME requirements for protein and fat deposition ob-
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

BW growth Protein deposition Fat deposition Feather growth

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
fat and 11.95 kcal/g of protein deposited. In the same way,
Boekholt et al. (1994) estimated that broilers need 10.90
and 8.57 kcal of ME/g for fat and protein deposition, re-
spectively.
The growth potential has been described by the Gomp-
occurred at 39 d for males and at 38 d for female broilers.
The differences between males and females are noticeable
after 21 d. Males gained more weight as they aged, and
the difference between males and females became more
significant.

ertz (1825) function in several studies (Emmans, 1981, 1989;
Knı́zetová et al., 1991; Hancock et al., 1995; Gous et al.,
1999). The growth parameters estimated in the present
study are in the same range of live weight (5,800 to 6,100
g for males and 4,300 to 5,200 for females) presented by
Knı́zetová et al. (1991), Hancock et al. (1995), and Gous et
al. (1999). Values of rate to maturity (B) for live BW are
similar to those estimated by Gous et al. (1999) of 0.037
and 0.038 and superior to those estimated by Hancock et
al. (1995) and Knı́zetová et al. (1991), which changed from
0.033 to 0.037. The Wm values of empty feather-free weight
obtained in this study were higher for males (5,939 g) and
lower for females (4,457 g) than those presented by Gous
et al. (1999), which were 5,500 to 5,700 g and 4,500 to 4,800
g for males and females, respectively.
One discrepancy was observed between the estimated
values in the present work and the previous reports. The
t* values obtained in this experiment show the precocity
of modern chickens. This parameter corresponds to the
inflexion point of the growth curve. Gous et al. (1999) and
Hancock et al. (1995) estimated maximum growth rate (t*)
at ages from 40 to 43 d and 43 to 45 d for males and females,
respectively. In the present experiment, the inflexion point
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As a bird develops, the proportion of feather protein in
total protein gain changes, which must be taken into ac-
count in calculation of amino acid requirements (Emmans,
1989). According to Gous et al. (1999), a description of
mature feathering and growth rates is necessary to predict
potential feather growth. In this study, these parameters
were higher than those described by Gous et al. (1999),
who found average values of Wm for feather growth of
300 and 230 g, values for B of 0.040 and 0.042, and values
for t* of 43 and 40 d for males and females, respectively.
The feathering of birds seems to be related to the genetic
line and mainly to sex. Females are selected for earlier
feathering for sex differentiation (Hancock et al., 1995).
Gous et al. (1999) indicated that the feathering characteris-
tics could be modified over time as a consequence of a
selection process.
Our results show that the growth rates for feathers were
initially higher for females, but the males had higher rates
after 21 d until 70 d of age. Kessler et al. (2000) observed
that females exhibited higher growth rates from 21 to 28
d of age and lower growth rates after 35 d. The males had
higher rates between 28 and 35 d and remained constant
until the age at processing.

TABLE 8. Evaluation of prediction models of daily ME requirements of male

Ross broilers by using data from experiment 2

ME model
Age MEi1 present Emmans Chwalibog
(d) (kcal/d) study2 P − O3 (1994)4 P − O3 (1991)5 P − O3

7 to 14 148 154 6 121 −27 127 −21

15 to 21 255 272 17 217 −38 222 −33
22 to 28 401 405 4 331 −70 342 −59
29 to 35 516 536 20 445 −71 448 −68
36 to 42 586 567 −19 465 −121 501 −85
43 to 49 660 713 53 602 −58 619 −41
50 to 56 721 619 −102 513 −208 520 −201
ESS6 14,356 73,910 59,336
1
MEi = metabolizable energy intake of Ross male broilers observed in the experiment 1.
ME = (307.87 − 15.63 T + 0.311 T2) BW0.75 + 13.52 Gf + 12.59 Gp; where T = temperature, BW0.75 = metabolic
2

BW, Gf = fractional fat deposition, and Gp = fractional protein deposition.

3
Difference between ME predicted by the models and observed MEi.
4
ME = 0.275 BP + 11.95 Gp + 13.38 Gf (BP = protein BW, g; Emmans, 1989).
5
ME = 111.1 BW0.75 + 8.03 Gp + 13.38 Gf (Chwalibog, 1991).
6
ESS = predicted (P) minus observed (O) error sum of squares.
 ENERGY MODEL FOR BROILER CHICKENS
The body chemical growth parameters in this study
showed the same trend of response observed by Gous et
al. (1999) and Hancock et al. (1995). These results show
the superiority of protein deposition in males during the
whole period of growth. As for females, fat deposition
increases as they age. In this experiment, rates of protein
deposition were different between sexes. Males had higher
rates of protein deposition from 21 to 42 d and females
from 21 to 35 d. After this age, protein deposition rates
decreased faster for females than for males, indicating
higher potential of males for protein deposition. Males
increased fat deposition until 28 d with a maximum plateau
from 28 to 49 d and then decreased after this age. However,
females increased their fat rates until 42 d and kept high
rates until 70 d. The higher and longer plateau for protein
deposition observed for males in comparison to the females
indicates that males develop more meat and better carcass
composition (Kessler et al., 2000). The reason for the sig-
nificant increments of fat deposition in females after 56 d
is the bird’s need for extra fat deposition during develop-
ment of the reproductive system (Gous et al., 1999).
Knowledge of an adequate description of body nutrient
deposition and ME models to determine accurate require-
ments for broiler chickens can help to establish specific
feeding programs and define optimum age for processing,
to provide better carcass quality, and reduce production
costs. The model proposed in this study includes the effect
of environmental temperature, which affects body compo-
sition and efficiencies of energy utilization for protein and
fat deposition. This factor improves the accuracy for pre-
dicting daily ME requirements.
ACKNOWLEDGMENTS
The authors thank the Fundação de Amparo a Pesquisa
do Estado de São Paulo (FAPESP) for the financial support.
REFERENCES
Agroceres Ross. 2000. Manual de Manejo de Frango de Corte.
Agroceres Ross, Rio Claro, SP, Brazil.
AOAC. 1990. Official Methods of Analysis. Vol. II 15th ed. Associ-
ation of Official Analytical Chemists, Washington, DC.
Balnave, D. 1974. Biological factors affecting energy expenditure.
Pages 25–46 in Energy Requirements of Poultry. T. R. Morris
and B. M. Freeman, ed. British Poultry Science, Edinburgh, UK.
Boekholt, H. A., P. H. Van Der Grinten, V. V. A. M. Schreus, M.
J. N. Los, and C. P. Leffering. 1994. Effect of dietary energy
restriction on retention of protein, fat and energy in broiler
chickens. Br. Poult. Sci. 35:603–614.
Blaxter, K. L. 1989. Energy Metabolism in Animals and Man.
Cambridge University Press, Cambridge, UK.
Chwalibog, A. 1991. Energetics of Animal Production. Acta Agric.
Scand. 41:147–160.
De Groote, G. 1974. Utilization of metabolizable energy. Pages
113–133 in Energy Requeriments of Poultry. T. R. Morris and
B. M. Freeman ed. British Poultry Science, Edinburgh, UK.
1369
Emmans, G. C. 1989. The growth of turkeys. Pages 135–166 in
Recent Advances in Turkey Science. C. Nixey and T. C. Grey.
Butterworths, London.
Emmans, G. C. 1986. Growth, body composition and feed intake.
World’s Poult. Sci. J. 43:208–227.
Emmans, G. C. 1981. A model of the growth and feed intake of
ad libitum fed animals, particularly poultry, in computers in
animal production. G. M. Hillyie, C. T. Whittemore, and R.
G. Gunn, ed. British Society of Animal Production, London.
Gompertz, B. 1825. On the nature of the function expressive of
the law of human mortality and on a new method of determin-
ing the value of life contingencies. Trans. R. Philos. Soc.
115:513–585.
Gous, R. M., E. T. Moran Jr., H. R. Stilborn, G. D. Bradford, and
G. C. Emmans. 1999. Evaluation of the parameters needed to
describe the overall growth, the chemical growth, and the
growth of feathers and breast muscles of broilers. Poult. Sci.
78:812–821.
Hancock, C. E., G. D. Bradford, G. C. Emmans, and R. M Gous.
Downloaded from http://ps.oxfordjournals.org/ at Cleveland Health Sciences Library on May 11, 2014
1995. The evaluation of the growth parameters of six strains
of commercial broiler chickens. Br. Poult. Sci. 36:247–264.
Hruby, M., M. L. Hamre, and C. N. Coon. 1996. Non-linear and
linear functions in body protein growth. J. Appl. Poult. Res.
5:109–115.
Hurwitz, S., I. Plavinik, I. Bengal, H. Talpaz, and I. Bartov. 1983.
The amino acid requirements of growing turkeys. 1—Model
construction and parameter estimation. Poult. Sci. 62:2208–
2217.
Hurwitz, S., M. Weiselberg, and U. T. Eisner. 1980. The energy
requirements and performance of growing chickens and tur-
keys as affected by environmental temperature. Poult. Sci.
59:2121–2128.
Kessler, A. M., and I. Brugalli. 1999. Recentes avanços do efeito
da nutrição no crescimento especı́fico de componentes da car-
caça de frangos de corte. Pages 1–19 in Proceedings of the
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
da quantidade de gordura na carcaça de frangos. Pages 107–
133 in Proceedings of the Conferência Apinco de Ciência e
Tecnologia Avı́colas. FACTA, Campinas, SP, Brazil.
Knı́zetová, H., J. Hyanek, B. Knize, and J. Roubicek. 1991. Analysis
of growth curves of fowl. I. Chickens. Br. Poult. Sci.
32:1039–1053.
Leeson, S., and J. D. Summers. 1980. Production and carcass
characteristics of the broiler chicken. Poult. Sci. 59:786–798.
Leeson, S., and J. D. Summers. 2001. Scott’s Nutrition of the
Chicken. 4th ed. University Books, Guelph, Ontario, Canada.
Macleod, M. G. 1990. Energy and nitrogen intake, expenditure
and retention at 20°C in growing fowl given diets with range
of energy and protein contents. Br. J. Nutr. 64:625–637.
Nieto, R., C. Prieto, I. Fernandez-Figares, and J. F. Aguilera. 1995.
Effect of dietary protein quality on energy metabolism in grow-
ing chickens. Br. J. Nutr. 74:163–172.
Pasternak, H., and B. A. Shalev. 1994. The effect of a feature of
regression disturbance on the efficiency of fitting growth
curves. Growth Dev. Aging 58:33–39.
Sakomura, N. K., R. Silva, H. P. Couto, C. Coon, and C. R. Pacheco.
2003. Modeling metabolizable energy utilization in broiler
breeder pullets. Poult. Sci. 82:419–427.
Statistica. 1996. Version Release 5.1. for Windows. Statsoft,
Tulsa, OK.