Animal Feed Science and Technology 233 (2017) 13–21

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Animal Feed Science and Technology

journal homepage: www.elsevier.com/locate/anifeedsci

Diet energy and feed intake in chickens

Henry L. Classen ∗
Department of Animal and Poultry Science, University of Saskatchewan, Saskatoon, SK, Canada S7N 5A8

a r t i c l e i n f o a b s t r a c t

Article history: Energy homeostasis is of fundamental importance to animal well-being as well as in the
Received 28 October 2015 feeding of species like chickens. Optimizing the balance between energy intake and expen-
Received in revised form 1 February 2016 ditures is required for efficient, highly productive flocks, and understanding its relationship
Accepted 7 March 2016
to production characteristics including energy and feed intake is fundamental to the formu-
lation of chicken diets and levels of other dietary nutrients. A historical perspective on this
Keywords: relationship is that chickens alter feed intake to maintain energy intake when diets con-
Broilers
tain variable dietary energy. In turn levels of other nutrients, and in particular amino acids,
Laying hens
should be adjusted in accordance. However, control of feed intake is complex and even if
Energy homeostasis
Genotype this perspective were correct, multiple factors can compromise the appropriate adjustment
Feed form in feed intake. The validity of broilers altering feed intake in response to dietary energy has
Energy to protein ratio been questioned for some time and more recent evidence suggests that modern strains of
laying hens also are unable to accurately or reliably alter feed intake in response to dietary
energy levels. Based on the premise that chickens do not appropriately respond to dietary
energy, diets must therefore be formulated to ensure adequate dietary energy to meet the
bird’s requirements for maintenance and production, and thereby maximize the protein
accretion potential of the diet.
© 2016 Elsevier B.V. All rights reserved.

1. Introduction

Energy homeostasis is a fundamental mechanism in mammalian and avian species, which provides for animal health and
adaptation to its living environment. By its nature and importance, maintenance of energy homeostasis is highly complex and
is the summation of a wide range of external and internal stimuli that in turn affect intake of nutrients. Stimulated in part by
the increase in human obesity, research on food intake has been extensive in mammalian species and it has provided a good
basic understanding of food intake control (Morton et al., 2014). Research on feed intake control in avian species is far less
extensive (Richards and Proszkowiec-Weglarz, 2007; Song et al., 2013). In addition, despite common signalling molecules,
it is unwise to extrapolate mammalian findings to avian species such as the domestic chicken (Gallus gallus) because of
different effects for important regulatory factors such as ghrelin, which is orexigenic in mammals and anorexigenic in birds
(Kaiya et al., 2013). The rationale for this brief introduction to energy homeostasis is to provide the reader with a reminder
of its complex nature and equally complex impact on feed intake.

Abbreviations: AMEn , apparent metabolizable energy (nitrogen corrected); gut, gastrointestinal tract; CCK, cholecystokinin; PYY, Peptide YY; GLP-1,
glucagon-like peptide-1; GLP-2, glucagon-like peptide-2.
∗ Corresponding author. Fax: +1 3069664560.
E-mail address: hank.classen@usask.ca

http://dx.doi.org/10.1016/j.anifeedsci.2016.03.004
0377-8401/© 2016 Elsevier B.V. All rights reserved.
14 H.L. Classen / Animal Feed Science and Technology 233 (2017) 13–21

A wide variety of both nutritional and non-nutritional factors affect feed intake in chickens (Ferket and Gernat, 2006;
Applegate, 2012), but this review will emphasize the relationship of a key dietary input (energy) and feed intake. Dietary
energy, derived from carbohydrates, lipids and protein, is the largest proportion of poultry diets and contributes importantly
to diet cost. It also interacts with other nutrients such as amino acids to promote high and efficient production in chickens.
The focus of this paper is not to review the nature of feed intake regulation in chickens, but rather to examine the nature of
diet energy effects on feed intake as well as to explore factors that may affect or camouflage the effects of energy per se.
The response of growing chickens to dietary energy has been the subject of much research and early research has been
thoroughly reviewed (e.g. Fisher and Wilson, 1974; Pesti and Smith, 1984; Leclercq, 1986). These reviews provide a historical
perspective and undoubtedly have shaped nutritional practice. A commonly accepted role of diet energy is shown in the
following quote written in the beginning of the review by Fisher and Wilson (1974):
“The primary response to dietary energy concentration is seen in feed consumption and in productive efficiency rather
than in production level”
This statement implies that within a reasonable range of dietary energy, feed intake responds to maintain a constant
energy intake, which in turn affects production efficiency, but does not affect production so long as other nutrients are
present at adequate levels. Fisher and Wilson (1974) outlined this response by examining much of the existing literature
(160 comparisons) based on publications where ingredients were known and of a practical nature, data were available on
growth and feed consumption, and the number of energy levels permitted the use of regression analysis. In experiments
meeting these requirements, feed intake was shown to go down with increasing energy, but statistical analysis revealed
diet energy only accounted for 28% of the variation in feed intake. This finding can be interpreted as a further indication
of the multitude of factors that impact feed intake. Interestingly dietary energy was more successful in predicting growth
rate (52% of variation) and feed efficiency (83% of variation). This summary further demonstrated that bird gender, age and
breed influenced the response of growing chickens to dietary energy, and also highlighted other factors that may impact
the comparison of energy levels such as feed density and the nature of feed ingredients. These factors, in fact, confound
interpretation of the specific effect of dietary energy. Of importance, the nature of these and other confounding factors
were not a criterion for inclusion or exclusion in their summary. These factors were similarly not included as a criterion by
others (e.g. Pesti and Smith, 1984). Failure to critically evaluate experiments raises questions about the value of research
summaries, and the lack of recognition or definition of confounding factors, further reduces their predictive ability. In the
following sections, the author will attempt to address the factors that may affect or confound results in experiments designed
to determine the effect of dietary energy on broiler performance and more specifically feed intake.

2. Factors affecting response of broilers and laying hens to dietary energy

2.1. Chicken genotype

Chickens have been extensively selected for a wide range of criteria over many decades. Selection of broiler genotypes has
resulted in positive phenotypic changes in growth, feed efficiency, meat yield, carcass fat, bird health as well as other criteria.
Many of these changes were scientifically demonstrated by Havenstein et al. (2003). Laying hens have similarly undergone
extensive selection for egg production, egg size and quality, body weight, health and welfare indicators. The nature of these
changes has been summarized by Anderson et al. (2013). It is recognized that both broiler chickens and laying hens will
continue to evolve in response to selection pressures, whose specific nature will also change in accordance with market
demands. Recognized physiological changes have occurred as a result of selection and these are the cornerstones of obvious
phenotypic traits, but there are also changes that either are not as obvious or have not been clearly identified, such as
mechanisms of feed intake control. The less clear understanding of the physiological mechanisms on feed intake in avian in
comparison to mammalian species is partially responsible for this lack of clarity in changes (Richards et al., 2010).
A positive correlation between feed intake and growth is expected and scientifically confirmed (Siegel and Wisman, 1966;
Pym and Nichols, 1979). It is also clear that feed intake has a heritable basis and will change in association with selection for
correlated traits or as a result of direct selection (Pym, 2005). That there may be differences in feed intake control mechanisms
in broiler strains has been recognized for some time (Burkhart et al., 1983; Lacy et al., 1986), as have differences in how
rapidly growing breeds respond to dietary energy content with some breeds demonstrating no or little effect (Pym and
Nicholls, 1979; Pym, 2005). Therefore, breed differences are an important confounding factor in the study of feed intake
responses to dietary energy. Furthermore, the continuing selection and sometimes rapid changes within commercial broiler
stock show that understanding breed responses to dietary energy is a moving target requiring periodic re-evaluation.
It is often stated that laying hens have retained the ability to adjust feed intake to dietary energy, but genotype varia-
tion also appears to affect this ability in laying hens as well. Table 1 summarizes the results of a number of experiments
where dietary energy is a portion of the experimental design. In the table, relative changes in dietary energy are related to
proportional changes in feed intake. The author has taken some liberty to summarize across other experimental treatments
to demonstrate relationships. This informal approach to examining the feed intake response to dietary energy is also not
meant to be comprehensive, but rather to demonstrate that certain strains of hens (e.g. Hy-Line W-36) do not appear to
respond, and while others respond but the nature of the response is neither uniform nor predictable. If a response occurs, in
most cases hens overconsume energy and dietary energy increases. Unlike in broiler chickens, where increased growth rate
 H.L. Classen / Animal Feed Science and Technology 233 (2017) 13–21 15

Table 1
Effect of dietary metabolisable energy on feed intake of pullets and laying hens.

Diet energy Feed intake

1
Reference Breed MJ/kg % g/b/d %1
2,4
dePersio et al. (2015) Hy-Line W-36 11.6 0 104 0
11.8 2.4 103 −1.0
12.1 4.8 104 −0.1
12.4 7.2 103 −1.0

Rama Rao et al. (2014) Babcock BV300 10.0 0 105 0

Energy (3) × protein (3)5 10.7 6.3 103 −2.4
11.3 12.5 98 −7.1

Jiang et al. (2013)2,3 Hy-Line Brown 11.5 0 104 0

Energy (3) × calcium (3)5 12.7 10.2 94 −9.2
13.4 16.2 93 −10.5

Rama Rao et al. (2013)4 Babcock BV300 9.6 0 114 0

10.0 4.4 111 −2.0
10.5 8.7 110 −3.2
10.9 13.1 104 −7.7
11.3 17.4 102 −10.4
11.7 21.8 101 −11.3

Pérez-Bonilla et al. (2012)2,4 Hy-Line Brown 11.1 0 115 0

11.5 3.8 114 −0.8
11.9 7.5 111 −3.5
12.4 11.3 110 −4.3

Frikha et al. (2009)2,4,6 Hy-Line Brown 11.2 0 56 0

Energy (3) × feed form (2)5 Pullets 11.8 5.3 55 −3.2
12.4 10.7 52 −7.2

Yuan et al. (2009)2,3 Bovans White 11.6 0 105 0

Energy (4) × breed (2)5 Dekalb White 11.8 1.6 105 −0.2
12.0 3.2 104 −1.1
12.2 4.8 102 −3.1

Gunawardana et al. (2008)2,7 Hy-Line W-36 11.7 0 97 0

Energy (3) × protein (3)5 12.0 2.8 98 0.7
12.3 5.7 95 −1.8
12.7 8.6 95 −2.1

Valkonen et al. (2008)2,4,8 Lohmann LSL

Energy (2) × housing (2)5 Phase 1 10.1 0 125 0
11.0 8.9 116 −6.7
Phase 2 10.0 0 133 0
11.0 10.0 125 −6.4
Phase 3 9.8 0 138 0
10.8 10.2 125 −8.9

Wu et al. (2007)2,4 Hy-Line W-36 11.7 0 92 0

Energy (4) × protein (3)5 11.8 0.9 93 0.6
11.9 1.7 92 −0.4
12.0 2.6 93 0.3

Jalal et al. (2007)2,3 Hy-Line W-36 11.8 0 108 0

Energy (2) × enzyme (2)5 Babcock B-300 12.1 3.2 109 0.1
Babcock BV300
Shaver White

Jalal et al. (2006)3 Hy-Line W-36 11.7 0 89 0

Cage space (4) × energy (3) × breed (4)5 11.9 1.8 88 −0.5
12.1 3.6 87 −1.7

Junqueira et al. (2006)2 ISA Brown 11.9 0 111 0

Energy (3) × protein (3)5 12.4 3.5 108 −2.7
12.8 7.0 110 −0.9

Scheideler et al. (2005)2,3 Hy-Line W-36 11.7 0 100 0

Energy (2) × enzyme (2)5 Babcock B-300 12.1 3.3 98 −2.1

Wu et al. (2005)2,3 Bovans White 11.4 0 108 0

Energy (4) × breed (2)5 Dekalb White 11.7 2.9 105 −2.0
12.1 5.8 103 −4.1
12.4 8.7 101 −6.0
16 H.L. Classen / Animal Feed Science and Technology 233 (2017) 13–21

Table 1 (Continued)

Diet energy Feed intake

Reference Breed MJ/kg %1 g/b/d %1

2,4
Bohnsack et al. (2002) Hy-Line W-36 11.7 0 93 0
Fat sources (2) × fat level (4)5 12.19 3.7 93 0
12.59 7.3 92 −0.7
12.99 10.5 90 −3.2

Leeson et al. (2001)2,3,10 Shaver White

Diet dilution4 10.3 0 115 0
10.9 5.9 109 −5.2
11.5 11.8 103 −10.4
12.1 17.6 103 −10.4
Reformulation2,4 10.3 0 127 0
10.9 5.9 117 −7.9
11.5 11.8 105 −17.3
12.1 17.6 97 −23.6

Harms et al. (2000)2,4 Hy-Line W-36 10.6 0 119 0

Energy (3) × breed (4) Hy-Line W-98 11.7 11.1 109 −7.8
Significant interaction5 Hy-Line Brown 12.9 22.2 108 −9.2
Dekalb White

Grobas et al. (1999)3 ISA Brown 11.2 0 122 0

11.8 4.9 116 −4.9

Keshavarz (1998)11 Babcock B300

Energy (2) × protein (2)5 Experiment 1 11.8 0 4109 0
12.7 7.8 4042 −1.6
Energy (2) × protein (4) 5
Experiment 2 11.8 0 6199 0
12.7 7.8 5975 −3.6

Keshavarz and Nakajima (1995) Babcock B300 11.8 0 101 0

Fat constant 12.7 7.8 101 −0.3

Peguri and Coon (1991)3,12 DeKalb XL 11.1 0 104 0

Energy (4) × temperature (6)5 11.5 4.2 103 −0.8
12.0 8.3 101 −2.5
12.5 12.5 98 −5.7
1
Percent change in relationship to the lowest diet energy level.
2
Diet energy primarily increased by adding fat.
3
Protein or essential amino acid levels were estimated to be equal across energy treatments.
4
Protein or essential amino acid levels increased proportionally with dietary energy.
5
Factorial arrangement treatments are indicated in italics. Values in tables summarize the feed intake response to dietary energy across main effects.
6
AME means of three diets − AMEn was 11.5, 12.1 and 12.7 MJ/kg from 1 to 45 days, 11.1, 11.7 and 12.3 MJ/kg from 46 to 85 days and 11.0, 11.6 and
12.1 MJ/kg from 86 to 120 days of age for the low, medium and high energy diets, respectively.
7
Energy levels consisted of 0, 79, 158 and 238 additional kcal of ME/kg based on diets containing 11.5, 11.6 and 11.8 MJ/kg. Table values are average ME
and feed intake values for the diets.
8
Trial initiated at 21 weeks of age and Phases 1, 2 and 3 lasted 20, 16 and 16 weeks, respectively.
9
Mean values ((12.10 + 12.05)/2 = 12.08; (12.53 + 12.45)/2 = 12.49; (12.92 + 12.80)/2 = 12.86).
10
In the first experiment, energy levels were adjusted by diet dilution, while in the second experiment diets were re-formulated to achieve diet energy
levels.
11
Feed intake: Experiment 1, g/bird 8–18 weeks of age; experiment 2, g/bird 0–18 weeks of age.
12
Average feed intake values from hens housed at 16.1, 18.9, 22.2, 25.0, 27.8 and 31.1 ◦ C.

matches the need to eat more feed, the reduced or lack of response in laying hens is associated with selection for reduced
body weight, improved feed efficiency and increased egg mass production.
It is clear that chicken genotype affects feed intake responses to dietary energy and that the response is not stationary
as the nature of strains changes rapidly with constant selection pressure applied to pure lines incorporated into the final
commercial hen, as well as the potential for changes in the pure lines.

2.2. Diet composition and digestible nutrient content

When comparing the response of chickens to dietary energy, one of two approaches can be taken: 1) formulation of
one diet and then diluting it with materials with little nutritional value or effect, or 2) formulation of diets using practical
feed ingredients. Both approaches can be criticized for potentially influencing the response of birds to dietary energy. In
the case of diet dilution, the assumption that diluting materials have little or no nutritional influence other than reducing
diet energy content can be questioned. Use of practical ingredients has the advantage of using feedstuffs that might be used
in commercial application, but has the disadvantage that ingredients may influence research results in a way unrelated to
energy. As a consequence, there is considerable potential for bird response to be related to the nature of ingredients that are
 H.L. Classen / Animal Feed Science and Technology 233 (2017) 13–21 17

either not found in all diets or are incorporated at different levels. Most research on the effect of dietary energy has taken
the practical feed ingredient approach. Some of the ingredient factors that may influence broiler response are nutrient or
anti-nutrient levels, nutrient digestibility and post-digestion metabolic effects.
Formulation of diets relies on the use of dietary specifications derived from a variety of sources, which are usually based
on averages of results from digestibility research. In research designed to estimate the impact of dietary energy on broilers,
this has the potential to reduce accuracy since specific samples of ingredients can vary considerably from these averages.
In research on nutritional requirements, it is well accepted that the level of the nutrients in diets should be measured to
confirm experimental assumptions. Yet this has not been the case for the vast majority of studies on dietary energy. Further
reducing accuracy in these studies is the failure to recognize known age differences in energy retention. It appears logical
that ingredients or diets used in this research should be assessed for energy retention in an age appropriate manner to reduce
the potential for inaccurate nutrient profiles. Method of measuring energy retention can also impact research results and
this is briefly discussed later in the review.
A key aspect of diet formulation is the level, balance, and digestibility of amino acids. Diets from early research on dietary
energy were formulated based on total amino acid content and a limited number of amino acids (sulphur amino acids,
lysine). Lower density, higher fibre ingredients frequently have lower amino acid digestibility and this has the potential to
impact growth and other response criteria. Failure to formulate for amino acids in a comprehensive fashion (beyond sulphur
amino acids and lysine) may result in the diet being limited by another amino acid, thereby affecting performance potential.
Finally, amino acid balance and the ideal protein concept (Emmert and Baker, 1997) can also affect bird response to dietary
energy. Amino acids in excess of that required for protein synthesis and other aspects of body metabolism are catabolised,
and this process incurs an energy cost (Sklan and Plavnik, 2002) and is metabolically stressful. It is only in most recent years
that some or all of these issues related to amino acids have been taken into account in dietary energy research.
The nature and amount of fibre in diets can also affect broiler response and changing fibre is a key method of reducing
diet energy. That soluble fibre can be an anti-nutritional factor has long been recognized (Choct et al., 1992) and therefore
alleviation of its effects via exogenous dietary enzymes is required for proper dietary energy comparisons. Initial research
on dietary energy in my lab utilized barley as a key ingredient in low energy diets. This research occurred before the
commercialization of exogenous enzymes so the soluble ␤-glucan content in barley affected not only the digestible nutrient
content of the barley, but also the digestibility of remainder of the diet. Insoluble fibre, often considered to have little
nutritional impact other than nutrient dilution, can also affect feed intake and digestive function (Svihus and Hetland, 2001;
Svihus, 2011).
Fat is the most common ingredient used to increase dietary energy and possibly has the most potential to be confounding
in dietary energy studies. Fat is recognized as providing more nutritional benefit than can be attributed to its metabolizable
energy, the measure of energy most used in previous research (Pesti and Smith, 1984). This “extra-caloric effect” can be
partially attributed to the lower heat increment of fat. To derive this beneficial effect, fat must be digested and absorbed
and this is not only affected by fat source but also by bird age; younger birds do not utilize fats as well as older birds and
the degree of difference is related to fatty acid chain length and saturation (Tancharoenrat et al., 2013). However, fat effects
on digestive function also increase utilization of other dietary nutrients and this effect is complicated by level and nature of
fat in the diet (Mateos and Sell, 1980; Mateos et al., 1982). Further, fat can affect diet palatability, thereby affecting research
results and in particular feed intake. Use of net energy (or equivalent) should enhance the accuracy of energy delivery in
diets varying in fat content, but other beneficial effects of fat are more difficult to estimate. With few exceptions research
on broiler and laying hen response to dietary energy has failed to account for the important influence of dietary fat.
In addition to fibre and fat, other ingredient factors appear to affect feed intake independent of diet energy. Scott (2005)
summarized a number of studies measuring the AMEn of cereal grains and found a lack of relationship between feed intake
and AMEn . This finding supports the hypothesis that broiler chickens are unable to adjust intake to match energy require-
ments. In contrast, strong positive correlations were found between feed intake and body weight in birds fed wheat and
barley diets. Scott (2005) proposes that differences in feed intake among grain samples relate to differences in time to hydrate
the diet. Based on this principle, one can visualize similar responses with other types of ingredients, possibly facilitated by
the presence of nutrient receptors in the gut (Geraedts et al., 2011; Rasoamanana et al., 2012; Duca and Lam, 2014). Receptors
are found on the apical face of the gut and are localized in enteroendocrine cells. Specific receptors for calcium, protein and
AA, fatty acids, and monosaccharides detect luminal concentrations and affect secretion of gastro-intestinal peptides such
as ghrelin, serotonin, cholecystokinin (CCK), PYY, glucagon-like peptide-1 (GLP-1), and GLP-2. These regulatory peptides
affect a wide range of physiological responses including gut motility, epithelial repair, satiation, and nutrient homeostasis.
Effects occur via paracrine and endocrine mechanisms that also involve direct action on vagal afferent nerves that transmit
neurological signals to the brain.

2.3. Feed form and feed processing

Feed form is well recognized to influence broiler performance with birds fed pelleted diets growing more rapidly and
more efficiently (McKinney and Teeter, 2004; Skinner-Noble et al., 2005; Brickett et al., 2007; Abdollahi et al., 2013a). A
primary reason for the increase in productivity is thought to relate to bird behaviour, more specifically, reduced energy use
due to less time eating and more time resting (McKinney and Teeter, 2004). However, other mechanisms may also play a role.
Heat and moisture processing have been shown to affect nutrient digestibility, with the nature of the response ingredient
18 H.L. Classen / Animal Feed Science and Technology 233 (2017) 13–21

dependent (Abdollahi et al., 2013b). Diet presentation can also impact both the ability of birds to consume feed as well as
its palatability for consumption (Latshaw, 2008). Volumetric density is reduced in mash feeding and this can impact the
ability to consume sufficient nutrients for maximum production, particularly when diets are low in nutrient density. Particle
size in mash diets can further impact diet palatability and this effect is modified by the level of dietary fat. Therefore, the
characteristics of a mash diet can confound broiler response to dietary energy. Regardless of the mechanism, pelleting diets
affects the effective caloric value of feed (McKinney and Teeter, 2004). Pellet quality is also impacted by diet ingredients,
so experiments should be designed to ensure comparable quality across energy treatments. It would seem rational that
research related to the response of birds to diet energy should employ the processing technique that it is being used to
predict.

2.4. Bird age

The response of broilers to dietary energy has been shown to be affected by bird age, with younger birds seemingly less
able to respond appropriately because of physical limits in digestive tract capacity (Brickett et al., 2007). An age effect could
logically be related to the reduced digestive ability of young birds (Sklan, 2001) and the nature the response could again be
affected by diet ingredients and feed form.

2.5. Energy to protein ratio

The relationship of energy to protein (indispensable amino acids) in experimental diets examining the effect of diet energy
has important implications on broiler response. Experiments have generally followed one of two approaches, providing
amino acids (digestible or total) at or above bird requirement, or balancing amino acids to the level of dietary energy. The
latter approach is primarily based on the assumption that birds have the ability to maintain energy intake by adjusting feed
intake to match the energy in the diet. If this is incorrect, as discussed above, or the response is unpredictable because of
other factors, this approach can produce invalid results because lower energy diets would be amino acid deficient. Broiler
response is affected by the most limiting factor in the diet, and therefore diets containing insufficient levels of indispensable
amino acids or energy will have lower response potential. The partitioning of energy and protein for maintenance and
production purposes will certainly be affected by dietary balance. The protein (amino acid) sparing effects of dietary energy
is a good example of partitioning change as less protein is used to meet the energy requirements of the bird and more is
directed towards growth, lean tissue accretion and egg production. The use of calorie to protein ratio for describing the
relationship between these two important diet fractions is outdated other than to discuss this relationship in the most
general of terms. More relevant is to describe the relationship of energy to the levels of ideal digestible amino acids, specific
effects and limitations of dispensable amino acids, and minimum levels of protein to supply nitrogen for non-essential amino
acid synthesis.

2.6. Environment and disease

Numerous non-nutritive factors can affect bird performance and hence have the potential to affect the feed intake
response of broilers to dietary energy (Applegate, 2012). It is beyond the scope of this report to examine them in detail,
but environmental factors of significance include temperature, environmental contaminants such as ammonia, housing
density, water and feed availability and lighting programs. Disease challenge and immunological stress are also important
in defining bird response. The nature of these effects is complex but among the mechanisms of action, partitioning of dietary
energy as a result of increased heat production and confounding effects on feed intake merit mentioning (Korver et al., 1998;
Liu et al., 2015).

3. Accounting for confounding factors in broiler energy research

With the above factors in mind, research was undertaken to investigate the impact of dietary energy, while attempting
to minimize confounding factors (Cho, 2012). Diets were formulated to contain 11.30, 11.85, 12.41 and 12.97 MJ/kg of AMEn
using practical ingredients (main ingredients − corn, wheat, barley, soybean meal, canola meal and canola oil), and fed
to Ross x Ross 308 mixed sex broilers housed in litter floor pens. The following aspects of the experimental design were
implemented to reduce confounding influences on broiler response.

• Ingredient AMEn and apparent ileal amino acid digestibility were determined at 5–6 and 21 days of age in broiler chickens
and diets were formulated using age appropriate values.
• Diets were formulated on a digestible amino acid basis with ideal protein balance.
• Diets were formulated to maintain a constant ratio between the ether extract content and AMEn of the diets.
• Diets were pelleted and fed in crumble or pellet form. Crumble and pellet quality, as assessed by sieving (particle size)
and pellet quality index procedures, demonstrated minor differences among diets. To reduce the impact of ingredients on
pellet quality, pellet binders were added to the diets.
 H.L. Classen / Animal Feed Science and Technology 233 (2017) 13–21 19

90% 80% 70%

4.0

Feed intake (kg/bird)

3.8

3.6

3.4

3.2
11.30 11.85 12.41 12.97
AMEn (MJ/kg)

Fig. 1. Effect of dietary energy (MJ/kg) on feed intake of broilers fed 90, 80 and 70% of Aviagen (2007) amino acid specifications (Cho, 2012).

70% 80% 90%

2.5
Body weight (kg)

2.3

2.1

1.9

1.7
11.30 11.85 12.41 12.97
AMEn (MJ/kg)

Fig. 2. Effect of dietary energy (MJ/kg) on body weight of broilers fed 90, 80 and 70% of Aviagen (2007) amino acid specifications (Cho, 2012).

• Exogenous enzymes were used to negate the impact of soluble fibre found in wheat and barley.
• Experimental environmental conditions (temperature, ventilation) were optimized or specified (e.g. space allocation).
• Anti-coccidial agents and growth promotants were used to minimize subclinical and clinical disease.

In the first experiment, starter (0–10 days), grower (11–26 days) and finisher (27–35 days) diets were formulated to
contain the digestible amino acid requirements outlined by Aviagen (2007). The four levels of energy were fed starting from
0, 10 or 26 days of age until trial end. Body weight and feed intake were monitored every 5 days for the length of the trial.
With minor exceptions, dietary energy did not affect feed intake regardless of when treatments were initiated. Body weight
gain increased and feed to gain ratio (mortality corrected) decreased with dietary energy with significant effects most often
found during the starter and grower phases. The results demonstrate that this strain of broilers did not increase feed intake
to maintain the same energy intake. Effects of growth rate and feed efficiency were similar to previous research, but not to
the same degree.
Because the levels of digestible amino acids in the first trial were relatively high, it was of interest to examine the
interaction of dietary energy with lower levels of dietary amino acids. The second trial used the same ingredients and energy
treatment levels, but these were fed in combination with three levels of amino acid supplementation that approximated 90,
80 and 70% of Aviagen (2007) recommendations. Significant interactions were found for most response criteria assessed. At
the highest level of dietary amino acids, body weight and carcass and breast yield increased, and feed to gain ratio decreased
with dietary energy, but feed intake was not affected. With the intermediate level of amino acids, dietary energy did not affect
response criteria. With the low levels of dietary amino acids, weight gain, feed intake, and carcass and meat yield decreased
with diet energy content. The interactions between diet energy and amino acids for feed intake and body weight are shown
in Figs. 1 and 2, respectively. These data confirm the lack of energy effect on feed intake in diets containing moderate to high
levels of dietary amino acids. Performance data from the 90% amino acid level suggest that the low levels of energy were
inadequate to provide for bird maintenance and maximum growth. It supports the concept that increased energy spares
amino acids from catabolism and permits their use in protein synthesis. The carcass and breast meat yield responses were
similar to that of body weight, which reinforces this conclusion. No obvious increase in fat content was noted. Therefore, not
all increases in body weight associated with increasing diet energy are due to increased fat deposition. At the 80% level of
amino acids, production characteristics were unaffected suggesting that all energy levels were adequate to match the growth
potential provided by the lower levels of amino acids. The reduction in feed intake, growth and carcass characteristics with
increasing energy in the 70% amino acid diets may have occurred because of the large imbalance between energy and amino
acids in the diet. Increased fat content was noted for birds fed 70% amino acids and high energy, suggesting that energy
homeostasis and feedback from lipid reserves may affect feed intake in more imbalanced diets; of note, feed to gain ratio
was unaffected by diet energy for this amino acid level.
20 H.L. Classen / Animal Feed Science and Technology 233 (2017) 13–21

Results from more recent research in broiler chickens are not uniformly in agreement with our research. Results are
variable with some research supporting the ability of broilers to eat to meet energy requirements (Leeson et al., 1996; Lemme
et al., 2005; Dozier I.I.I. et al., 2006), while other research failing to see an effect of dietary energy on feed intake (Plumstead
et al., 2007; Latshaw, 2008; Delezie et al., 2010; Li et al., 2011; Mbajiorgu et al., 2011). Although more recent papers are
more likely to account for the confounding papers described above, at least some of these factors remain unaccounted for in
most research. An opportunity exists to complete additional research in this area (possibly as an international collaborative
approach) to more fully understand the reasons for variable responses and to provide additional data for modelling energy
levels in broiler and laying hen diets. It is the author’s opinion that research to provide this understanding must be based on
current genotypes and industry practice, and avoid confounding factors that reduce the value of research.

4. Conclusions

Energy is an expensive component of broiler and laying hen diets with energy, and it is increasing in prominence due to
increased competition for starch- and oil-enriched feedstuffs. This reality is likely not to change in the near future and as
the poultry industry responds to world demand for high quality animal protein, the efficient use of ingredients will become
paramount. Research has convincingly found that diet energy does not accurately predict feed intake and in at least some
genotypes does have any effect. Therefore, generating a comprehensive understanding of the amount of energy required for
maintenance and production, and the balance between energy and ideal amino acid content is of increasing importance.

Conflict of interest

None.

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