Review: Energy partitioning in broiler chickens

G. Lopez and S. Leeson1

Department of Animal & Poultry Science, University of Guelph, Guelph, Ontario, Canada N1G 2W1.
Received 12 July 2007, accepted 20 February 2008.
Lopez, G. and Leeson, S. 2008. Review: Energy partitioning in broiler chickens. Can. J. Anim. Sci. 88: 205
212. In
commercial nutrition and in research studies, metabolizable energy (ME) is the standard measure of energy used in
describing energy requirements and diets for poultry. The provision of dietary energy will influence the intake of all other
nutrients. Broilers exhibit an outstanding ability to control their energy intake by adjusting their feed intake as diet energy
concentration changes. There is still considerable debate on the accuracy, precision and usefulness of different procedures
used for determining ME values of diets and ingredients. ME intake is generally partitioned into energy retained (ER) in
body tissues (mainly as fat and protein) and as heat production (HP): ME HPER. There are few reported estimates of
HP and its components, fasting heat production (FHP), heat production due to physical activity and the thermic effect of
feeding (TEF). Requirements for maintenance (MEm), including major components of FHP and physical activity, are
established at around 155 kcal kg BW0.60. We recently reported that maintenance requirements for young broilers based on
kg BW0.75 were 8% lower than the values estimated using kg BW0.60, and that BW raised to the exponent 0.60, was a more
precise estimator. Gross energy retained in the body as fat (TERF) and protein (TERP), together contribute most of the
total energy retained (TER) in the body. Efficiency of ME utilization above maintenance varies from 70 to 84% for lipid
deposition in adult birds and between 37 and 85% in growing birds.
Key words: Energy, broiler, metabolic rate, energy retention
Lopez, G. et Leeson, S. 2008. Repartition de lenergie chez le poulet de chair. Can. J. Anim. Sci. 88: 205
212. Lenergie
metabolisable (EM) est letalon habituellement employe en nutrition, tant dans le commerce quen recherche, pour decrire
les besoins denergie de la volaille et lenergie contenue dans les aliments. Lenergie des aliments exerce une influence sur
lingestion des autres elements nutritifs. Les poulets de chair montrent une capacite etonnante en modifiant la quantite
daliment ingeree en fonction de la concentration denergie dans la ration. Lexactitude, la precision et lutilite des diverses
methodes servant a` determiner la quantite de EM dans la ration et ses composantes font toujours lobjet dune vive
polemique. En general, on repartit la EM ingeree en plusieurs elements, a` savoir lenergie retenue (ER) dans les tissus de
lorganisme (principalement sous forme de matie`re grasse et de proteines) et celle liberee sous forme de chaleur (CH),
EM CHER. On a relativement peu estime la CH et ses composantes, soit la chaleur produite durant le jeune (CHJ),
celle resultant de lactivite physique et leffet thermique du a` lalimentation (ETA). On fixe les besoins dentretien (EMe),
qui incluent la CHJ et lactivite physique, autour de 155 kcal par kg de poids corporel (PC) multiplie par lexposant 0,60.
Les auteurs ont recemment signale que les besoins dentretien des jeunes poulets calcules en fonction du PC0,75 sont de 8%
inferieurs aux valeurs estimees par kg de PC0,60, et que le PC multiplie par lexposant 0,60 en permet une estimation plus
precise. Lenergie brute conservee dans le corps sous forme de matie`re grasse et de proteines explique la majeure partie de
lenergie totale gardee par lorganisme. Au-dessus des niveaux dentretien, lefficacite dutilisation de la EM varie entre 70
et 84% pour le depot des lipides chez les oiseaux adultes et entre 37 et 85% chez ceux en croissance.
Mots cles: Energie, poulet de chair, vitesse de metabolisme, retention de lenergie

The energy contribution from poultry diets is usually

described in terms of metabolizable energy (ME) and/or
net energy (NE). In commercial nutrition and in most
research studies, ME is the standard measure of energy
used in describing both energy requirements and diets
for poultry. ME can be accurately determined from the
difference between the gross energy of the feed and the
gross energy of the excreta derived from such feed
[National Research Council (NRC) 1994]. ME has been
commonly accepted and extensively used to compare
energy values of feedstuffs, and diets for poultry, and

energy requirements are commonly expressed in these

units. However, ME is rarely measured under commercial conditions.

Abbreviations: BW, body weight; E:P, energy:

protein; ER, energy retained; ERF, energy retained
in the body as fat; ERP, energy retained in the body
as protein; FHP, fasting heat production; HP, heat
production; ME, metabolizable energy; MEm, ME
intake for maintenance; MEI, ME intake; TEF,
thermic effect of feeding; TER, total energy
retained; TERF, total energy retained in the body
as fat; TERP, total energy retained in the body as
protein

To whom correspondence should be addressed (e-mail:

sleeson@uoguelph.ca).
205

206 CANADIAN JOURNAL OF ANIMAL SCIENCE

The concentration of energy in the diet will greatly

influence the intake of all other nutrients and so the
utilization of ME and these other nutrients. In this
regard, broilers exhibit the ability to control energy
intake by adjusting their feed intake as diet energy
concentration changes (Leeson et al. 1996a, b).
ME is an expensive component of poultry diets and
represents a large portion of the total cost of broiler
production. Many attempts have been made to increase
the accuracy, precision and speed of methods of
determining ME values, and especially for dietary
ingredients (Sibbald 1976, 1979; Farrell 1978; Dale and
Fuller 1984; McNab and Blair 1988). This situation has
been especially critical in the broiler industry, since
broilers consume large amounts of feed compared with
other poultry species, and there are difficulties inherent
in working with young birds in various bioassays
(Leeson and Summers 2005).
There is still considerable debate on the accuracy,
precision and usefulness of different procedures described in the literature for determining ME values of
diets and ingredients. Some of the factors that influence
ME values have been the variability of ME values across
laboratories and the different methods used for measurement (Bourdillon at al. 1990a), and especially the
lack of agreement on the suitability of the adult rooster
to determine ME values for growing broilers. Differences in ME values have been reported when comparing
young broilers and adult roosters (Kussaibati et al.
1982; Mollah et al. 1983; Hartel 1986; Bourdillon et al.
1990b; Carre et al. 1995; Farrell et al. 1997) and adult
laying hens (Petersen et al. 1976; Farrell et al. 1997).
Another factor influencing ME values has been the
practice of adjusting ME values for nitrogen (N)
retention (MEn). This is a common practice in poultry
nutrition, and MEn values of different ingredients are
commonly reported in the literature (NRC 1994). It is
generally accepted that correction to zero N retention is
essential when comparing ME values across species that
have inherently different rates of growth or egg production, and hence different levels of nitrogen retention.
Likewise, N correction seems essential for comparison
of the ME values determined with juvenile vs. mature
birds, since the former will be gaining body protein,
while the latter are usually at zero nitrogen retention.
Some 50 yr since the first application of correction
for nitrogen balance by Hill and Anderson (1958), the
correction factor is still debated, especially for growing
broilers, where dietary protein is penalized assuming
that none of it is deposited as protein tissue (De Groote
1974; Sibbald 1982; McNab 2000; Lopez and Leeson
2007, 2008).
Energy systems, including that for ME, are used to
predict values of feeds relative to energy requirements
for maintenance and production. This scenario is based
on the concept of partitioning energy requirements of
growing animals between requirements for maintenance
and production (Birkett and De Lange 2001a; Lopez

and Leeson 2007). Currently, factorial models are

commonly used to represent the relationship between
ME intake (MEI) and energy retained (ER), including
requirements for maintenance (MEm) (Kielanowski
1965; Birkett and De Lange 2001a; Lopez and Leeson
2007). In this model, the utilization of ME intake above
maintenance depends on the partition of energy retained
as protein (ERP) and as fat (ERF) and their respective
efficiencies for deposition. There is current interest in
broiler research, in quantifying and partitioning daily
ER in the body, as ERF and ERP, and the utilization of
ME intake (Van Milgen et al. 2001; Lopez and Leeson
2005).
As broilers grow, accretion of body fat and body
protein is impacted by nutrition, bird strain, sex,
environmental conditions, body weight (BW) and associated degree of maturity, and by the interaction
between some or all these parameters. Diet composition
can have a major impact on carcass composition
through the deposition of nutrients as fat or protein.
To quantify these body components during growth and/
or over time, several mathematical models have been
developed (Wilson 1977; Wiseman and Lewis 1998).
Genetic improvements in growth rate have been progressive and generally ahead of the established nutrient
requirements, leaving most recommendations quickly
redundant (NRC 1994). This selection for body weight
gain has resulted in broilers that reach market weight
earlier (Nicholson 1998; Remignon and Le Bihan-Duval
2003) at an immature body weight and often without
achieving maximum genetic potential for fat and protein
deposition in terms of absolute quantities deposited
each day. Understanding the energy requirements of the
birds and how modern broilers quantitatively deposit
fat and protein during a commercial life cycle would be
useful to the industry so as to establish useful economic
models that take into account the biology of growth of
broiler chickens.
IMPACT OF BODY COMPOSITION ON ENERGY
REQUIREMENTS
After hatching it is expected that as body weight
increases over time, the quantities and proportions of
body fat and protein increase at different rates (Emmans
1995) with fat deposits potentially increasing faster at
older ages (Leenstra 1986). The relationship between
protein and fat in the body is influenced by nutrition
(Deschepper and de Groote 1995; Wiseman and Lewis
1998), genotype (Edwards and Denman 1975; Leenstra
1988, 1989; Havenstein et al. 1994 a, b), sex (Leeson and
Summers 1980; Cahaner and Leenstra 1992; Leenstra
and Cahaner 1992), environmental conditions (Kubena
et al. 1972; Cahaner and Leenstra 1992; Leenstra and
Cahaner 1992), and body weight (BW) and associated
degree of maturity (Leenstra 1986, 1989; Havenstein
et al. 1994a; Decuypere et al. 2003).
Carcass fat is generally considered to be an unfavorable trait in the broiler industry (Remignon and Le

LOPEZ AND LEESON * ENERGY PARTITIONING IN BROILER CHICKENS

Bihan-Duval 2003), leading to studies in genetic selection (Whitehead and Griffin 1984; Leenstra 1988,
Whitehead 1990; Pym et al. 2004) and nutrition (Zubair
and Leeson 1994; Leeson and Zubair 1997) aimed at
reducing or limiting carcass fat. In most commercial
feeding programs, the desire for fast growth with ad
libitum feeding invariably entails moderate levels of
energy and high crude protein for the starter diets, and
high energy and lower crude protein for the later diets
(Leeson and Summers 2005). It is well documented
that such changes to energy:protein (E:P) in the diet are
associated with increased weight gain as fat (Bartov
et al. 1974; MacLeod 1990, 1991; Wiseman and Lewis
1998; Morris 2004).
Leeson and Summers (1980) investigated body components in male and female broilers to 70 d in response
to increasing energy:protein (E:P) in the diet. These
authors reported that as BW increases, both fat and
protein deposition increase, with body fat content as a
percent of the body weight increasing most dramatically
in both males and females, while body protein content
remained fairly constant. Such increases in carcass fat
deposition are confounded by the usual increase in E:P
in the diet as the bird ages (Wiseman and Lewis 1998)
and the desire to achieve genetic potential for growth
(Havenstein et al. 1994a, b). Protein accretion, on the
other hand, is usually predetermined by the genetic
potential of the bird, assuming the diet supplies adequate amounts and balance of amino acids. Leaner birds
usually result from using a single experimental diet
where protein/AA and energy are constant (Leeson et al.
1996a, b). Selection for reduced abdominal fat (Cahaner
1988; Leclercq 1988) and improved feed efficiency
(Leenstra 1988; Buyse et al. 1998) has produced leaner
broilers, although, again, this will be affected by diet
energy level used in a given study.
Mathematical growth models have been used to study
the influence of change over time in diet energy (Wiseman and Lewis 1998), or dietary protein (Eits et al.
2005) on body components such as fat (Wiseman and
Lewis 1998). These models are used as a tool to predict
BW with desired carcass characteristics. The Gompertz
equation has also been adopted in broiler studies to
appropriately describe growth over time (Wilson 1977;
Emmans 1995; Hurwitz and Talpaz 1997; Darmani et al.
2002) and/or growth of body components in broilers
(Tzeng and Becker 1981; Peter et al. 1997; Wiseman and
Lewis 1998; Gous et al. 1999). This equation describes a
sigmoidal pattern with slow initial growth followed by
acceleration up to a certain age (the inflection point)
followed by subsequent decrease in the rate as body
weight approaches its maximum near to sexual maturity
(Hurwitz and Talpaz 1997). Tzeng and Becker (1981)
fitted the nonlinear Gompertz equation to abdominal
fat, intending to predict total carcass fat over time
(Becker et al. 1979). Usually the methods for evaluating
the growth of body components in broiler studies using
the Gompertz model have been carried out for extended

207

periods of time up to 10 to 16 wk (Tzeng and Becker

1981; Peter et al. 1997; Wiseman and Lewis 1998; Gous
et al. 1999). Under these conditions, the asymptotic
values of liveweight or body components are obviously
better estimated. However, as todays broilers reach
commercial body weight earlier (approximately 6 wk)
without achieving maximum genetic potential for fat
and protein deposition, the asymptotic value of liveweight or body components will never be realized, and
this may lead to potentially unreliable predictions of
body weight, fat or protein deposition using such model
predictions (Lopez and Leeson 2007).
PARTITIONING OF METABOLIZABLE ENERGY
ME intake is generally partitioned into energy retained
(ER) in body tissues (mainly as fat and protein) and as
heat production (HP): ME HPER (Close 1990;
Lawrence and Fowler 2002). In thermoneutral conditions, HP represents heat associated with the utilization
of ME intake for maintenance (MEm) and productive
processes, which, in juvenile broilers, represents 52
64%
of intake (Fuller et al. 1983; Van Milgen et al. 2001;
Noblet et al. 2003). Therefore, ER represents the
difference between ME and HP and so ME
HP ER.
When evaluating ER, it is necessary to measure energy
retained as both fat (ERF) and protein (ERP), along
with their efficiency of utilization of ME, usually termed
kf for fat and kp for protein deposition. Estimates
of ERF and ERP in broilers have been determined
using such methods as indirect calorimetry (Farrell
1974; Fuller et al. 1983; Van Milgen et al. 2001; Noblet
et al. 2003) and comparative slaughter (Fuller et al.
1983; MacLeod 1991). However, values for kf and kp
have only been estimated using statistical models from
experiments involving varying degrees of quantitative
feed restriction. (Boekholt et al. 1994).
There have been limitations in nutritional studies
using growing broilers to adequately define ME utilization and, consequently, their energy requirements. There
are few reported estimates of HP and its components,
fasting heat production (FHP), heat production due to
physical activity and the thermic effect of feeding (TEF)
(Van Milgen et al. 2001; Noblet et al. 2003). Feeding
broilers from 21 to 35 d of age, Van Milgen et al. (2001)
subdivided HP into its three major components using
indirect calorimetry, showing that FHP and physical
activity together represents 36
37% of ME intake. In
their experiment, Van Milgen et al. (2001) described
physical activity as a major component of maintenance
(8
10%). Requirements for MEm, including major
components of FHP and physical activity, were established at 152
157 kcal kg BW0.60. These values were
similar (155 kcal kg BW0.60) to those reported by Lopez
and Leeson (2005) using comparative slaughter, and the
same estimate of metabolic body modifier (kg BW0.60).
There are few estimates of the efficiency of ME
utilization for fat and protein deposition. In most
published studies, estimates for kf and kp are established

208 CANADIAN JOURNAL OF ANIMAL SCIENCE

through experiments involving the feeding of graded

levels of energy or with a wide range of induced energy
intakes so as to obtain different rates of protein and
lipid deposition (Boekholt et al. 1994). This approach
has been questioned in studies with growing pigs, the
argument being that in most animals fat and protein
deposition are confounded (Noblet et al. 1999). Currently, the model that is most commonly used to predict
the effect of diet on growth (ERF and ERP) and
efficiency is based on the equation of Kielanowski
(1965). In this model, ME intake (MEI) is defined as:
MEI MEm(1=kf ERF)(1=kpERP)
where MEmME for maintenance as a function of
body weight (BWb), and kf and kp efficiencies of
utilization of ME for fat and protein deposition,
respectively (Boekholt et al. 1994; Birkett and de Lange
2001a; Lawrence and Fowler 2002).
MAINTENANCE ENERGY REQUIREMENTS
Maintenance energy requirement is described as the
state of the animal where its body composition remains
constant and when there is no production or work/
activity (Larbier and Leclercq 1994; Wenk et al. 2001).
There are few estimates of maintenance energy requirements in poultry (MacLeod et al. 1988; Sakomura et al.
2003) and, in particular, growing broilers (Van Milgen
et al. 2001; Lopez and Leeson 2005).
MEm represents a large portion of the MEI in
broilers, being in the order of 42
44% (Lopez and
Leeson 2005), and is influenced by the method used to
express MEm. The MEm of broilers is traditionally
reported as a function of BW raised to the 0.75 power
(Fuller et al. 1983; MacLeod 1990; Boekholt et al. 1994;
Buyse et al. 1998). Recent information suggests that
MEm in broilers is, in fact, more adequately described
by BW exponents other than 0.75 (MacLeod 1991, 1997;
Van Milgen et al. 2001; Noblet et al. 2003; Lopez and
Leeson 2005). This difference directly influences the
partitioning of energy between maintenance, fat and
protein deposition and, by inference, efficiency of energy
use. Underestimation of MEm for younger or smaller
birds implies less energy for production and therefore,
by calculation, greater apparent efficiency for growth
(higher kp and kf). The converse effect, but to a less
degree is expected for overestimation of MEm for older
heavier birds (Lopez and Leeson 2005). Few studies
have been conducted in order to comprehensively
understand the efficiencies for protein and fat deposition
in poultry, and most such data are based on BW0.75
(Boekholt et al. 1994), or data extrapolated from studies
with pigs (Birkett and de Lange 2001b; Noblet et al.
1999). Lopez and Leeson (2005) reported that maintenance requirements for young broilers based on kg
BW0.75 were 8% lower than the values estimated using
BW0.60, and that BW raised to the exponent 0.60, was a

more precise estimator. Close (1990) indicated that

differences in MEm are mainly affected by changes in
body composition. Since, at a given body weight, fat
tissue contributes little to heat production compared
with that of muscle, it is suggested that maintenance
energy requirements are lower in fat animals than in
lean animals (Close 1990). MacLeod et al. (1988) found
significantly higher fasting heat production and N
retention in broiler lines selected for leanness than those
selected for fatness, suggesting an increased maintenance energy requirement in lean birds. Questions still
remain whether fat animals have lower MEm or if
animals with a lower MEm become fatter (Van Milgen
et al. 1998).
RETAINED ENERGY AS FAT AND PROTEIN AND
EFFICIENCY OF ENERGY RETENTION
As broilers grow, accretion of fat and protein is the
result of the interaction among bird strain, sex, environmental conditions, nutrition, body weight (BW) and
degree of maturity. It is expected that mature animals
retain energy mainly as fat, while growing animals retain
energy as both fat and protein. Gross energy retained in
the body as fat (TERF) and protein (TERP), together
contribute most of the total energy retained (TER) in
the body. TER is calculated from accumulation of fat
and protein and by using corresponding energy values
of 9.5 kcal g1 fat and 5.7 kcal g1 of protein
(Znaniecka 1967; Hakansson and Svensson 1984). Since
fat and protein accretion likely differ in their efficiencies
of transfer of energy from feed to tissue (Buttery and
Boorman 1976; Pullar and Webster 1977), changes in
the proportion of both fat and protein during growth
influence the total energy in the body and especially the
efficiency of such gain.
Various studies have indicated that the efficiency of
protein deposition is lower than that for fat deposition
(Petersen 1970; De Groote 1974; Boekholt et al. 1994).
De Groote (1974) reported that the efficiency of ME
utilization above maintenance for lipid deposition in
adult birds varies from 70 to 84% and between 37 and
85% in growing birds. Petersen (1970), using White
Plymouth Rock birds, estimated efficiencies of 0.51 and
0.78 for protein and fat, respectively, indicating a need
for 11.2 kcal of ME g1 protein and 12.2 kcal of ME
g1 of fat deposited. More recent information in
growing broilers suggests higher energetic efficiencies
for protein (0.66) and fat (0.86) deposition, (Boekholt et
al. 1994), indicating lower needs for protein (8.63 kcal
ME g1) and fat deposition (10.9 kcal of ME g1).
Similar efficiencies (0.65 and 0.83) are reported in
comparable studies with growing pigs (Noblet et al.
1999). Moreover, due to the close association between
body water and body protein in lean meat, the ME
requirements per gram of lean tissue gain are much
lower than those per unit of fat tissue gain. Both
increases in ERP/ERF and reductions in energy needs
per gram of protein deposition contribute to increase in

LOPEZ AND LEESON * ENERGY PARTITIONING IN BROILER CHICKENS

feed efficiency in modern broilers (Lopez et al. 2007).

Broilers may also have been inadvertently selected for
greater rate of protein synthesis and/or reduced protein
degradation (Urdaneta and Leeson 2004).
Variation in kp and kf (Petersen 1970; De Groote
1974; Boekholt et al. 1994) is due to such factors as the
nature of the diet, animal effects and the function for
which feed is used by the animal (Birkett and de Lange
2001a). For example, studies in growing pigs show that
ME from dietary fat is used very efficiently for fat
deposition (0.90), while lower values are reported for the
formation of fat from protein (0.66). This is due to the
different energetic efficiencies in the chemical transformations involved in the synthesis of fat and protein from
absorbed nutrients (Van Milgen et al. 2001).
Metabolizable energy intake is well documented to
influence body composition (Hakansson and Svensson
1984; Boekholt et al. 1994; Wiseman and Lewis 1998)
and therefore body TER. Boekholt et al. (1994) fed
broilers from 60 to 100% of normal daily energy intake
and reported that daily retention of fat and protein was
linearly related to energy retention suggesting that in
growing broilers each additional unit of gain generated
by energy intake over 43 kcal kg W0.75 d was composed
of constant amounts of protein and fat, but different
proportions of energy as protein (15%) and fat (85%).
These data suggest that at an ER of 43 kcal kg W0.75 d,
ERF is zero and only protein is retained, perhaps at the
expense of fat mobilization (Boekholt et al. 1994).
CONCLUSIONS
Today, broilers reach commercial body weight very
early, at an immature body weight and often without
achieving maximum genetic potential for fat and protein
deposition in terms of absolute quantities deposited
each day. It is calculated that within the commercial
growing range of 0
42 d, broilers deposit body fat and
protein that together represent 35 to 40% of their
daily ME intake (Lopez and Leeson 2005). Quantifying
and partitioning TER as TERF and TERP as major
components of the requirement of ME in growing
broilers can be used in the industry to establish useful
models that will have economic consequences allowing
better management decisions and taking into account
the biological need of growth of modern birds within the
juvenile commercial age range.
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