Livestock Science 231 (2020) 103856

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Livestock Science
journal homepage: www.elsevier.com/locate/livsci

Energy and protein requirements of woolless sheep under tropical T

conditions
Sandra de Sousa Barcelosa, Kaliandra Souza Alvesa, Rafael Mezzomoa, Janaina Barros Luza,
Darley Oliveira Cutrimb, Daiany Íris Gomesa, Luis Rennan Sampaio Oliveiraa,
⁎
Karolina Batista Nascimentoc, Mateus Pies Gionbellic,
a
Department of Animal Science, Amazonia Rural Federal University, Parauapebas, PA 68515-000, Brazil
b
Federal Institute of Tocantins, Pedro Afonso, TO 77710-000, Brazil
c
Department of Animal Science, Federal University of Lavras, Lavras, MG 37200-000, Brazil

A R T I C LE I N FO A B S T R A C T

Keywords: The objective of this study was to determine the energy and protein requirements for maintenance and weight
Comparative slaughter gain of growing woolless sheep and, in addition, to estimate the shrunk body weight (SBW) and empty body
Empty body weight weight (EBW) of the animals according to their body weight (BW) in the fed state. Data from 150 animals from
Growing lambs three comparative slaughter experiments with confined castrated sheep averaging 5 ± 0.96 months of age and
Metabolizable energy
similar BW (20.74 ± 2.99 kg) were used. Three models were evaluated to estimate EBW from SBW: linear,
Metabolizable protein
Net energy
linear with intercept and non-linear. The following requirements were estimated: net energy for maintenance
(NEm), metabolizable for maintenance (MEm), net energy for gain (NEg), efficiency of use of metabolizable
energy for gain, metabolizable protein requirement for maintenance (MPm), net protein requirement for gain
(NPg), and efficiency of use of metabolizable protein for gain (k). Fasting losses corresponded to 6.18% of BW
and the better model to establish the relationship between SBW and EBW was nonlinear. The NEm and MEm were
70.44 and 109.34 kcal/kg EBW0.75/day, respectively. For NEg, the adjusted model was based on the ratio of the
energy retained as a function of the EBW and the desired empty body weight gain (EBG). Considering two lambs
weighing 15 kg and 30 kg with a daily gain of 100 g, the estimated NEg is 181 and 360 kcal/kg EBW0.75/day,
respectively. The estimated kg was 0.407. For MPm, the value of 3.19 g/kg EBW0.75/day was estimated. The
estimated NPg for animals of 15 and 30 kg, with the same gain rate was 13.13 and 13.61 g/day, respectively,
which corresponds to the protein concentration in gain of 170 and 161 g/kg EBG, and efficiency of use of the
metabolizable protein for gain ranging from 0.66 to 0.41. The nutritional requirements of woolless sheep did not
follow the standards assumed by the international feeding systems, except for the energy requirements for
maintenance and efficiency of use of the metabolizable energy for maintenance, which did not differ. The losses
represented by the gastrointestinal tract (GIT) after fasting and slaughtering periods are higher in woolless sheep
compared to those presented by the international feeding systems.

1. Introduction requirements for woolless sheep, the volume of information generated

Although most of the sheep raised in tropical conditions worldwide
are from woolless breeds, little information is available on their nutri-
tional requirements (Galvani et al., 2010; Chay-Canul et al., 2011,
2014). In Brazil, for example, most of the sheep herds are of the Santa
Ines breed and their crosses, due to their great adaptability to tropical
conditions (Souza et al., 2013), resistance, high productivity, and pro-
lificity (Pereira et al., 2017).
Although there are previous efforts to evaluate nutritional
has not yet been sufficient to make it possible to elaborate a system of
requirements for these animals under tropical conditions, once research
with this focus is costly and laborious. Comparative slaughtering, for
example, although allows to determinate the nutritional requirements
under conditions similar to those that the animals are raised and al-
though not require special equipment, is a costly methodology, once it
makes carcasses unfeasible and requires the slaughter of these animals.
In this context, the formulation of the diets for sheep under tropical
conditions is still carried out using international feeding systems, which

⁎
Corresponding author.
E-mail address: mateus.pg@ufla.br (M.P. Gionbelli).

https://doi.org/10.1016/j.livsci.2019.103856
Received 3 May 2019; Received in revised form 25 September 2019; Accepted 30 October 2019
Available online 31 October 2019
1871-1413/ © 2019 Elsevier B.V. All rights reserved.
S.d.S. Barcelos, et al. Livestock Science 231 (2020) 103856

were elaborated from experiments and animals in other conditions of Table 1

temperature and climate than tropical (INRA, 1988; AFRC, 1993; Descriptive statistics of the database used.
CSIRO, 2007; NRC, 2007). Studies on the Santa Ines breed Variable Item
(Galvani et al., 2014; Pereira et al., 2017) show controversial results Minimum Mean Median Maximum SD CV (%)
regarding the energy and protein recommendations contained in these
1
iSBW 14.60 20.74 20.65 27.65 2.99 14.40
systems, demonstrating the importance of specific research for local
iEBW2 10.82 15.87 15.89 21.42 2.38 15.03
conditions and breeds. Additionally, previous research also suggests fBW3 16.54 31.24 32.16 42.50 4.95 15.84
that there are differences in nutritional requirements between wool and fSBW4 15.10 28.37 29.80 40.20 5.28 18.60
woolless sheep (Chay-Canul et al., 2016). fEBW5 10.04 22.71 23.82 33.39 4.94 21.77
The objective of this study was to determine the energy and protein SBG (kg/d)6 −0.03 0.12 0.10 0.31 0.07 57.69
EBG (kg/d)7 −0.02 0.11 0.10 0.24 0.06 55.79
requirements for maintenance and weight gain for growing castrated
SBW/BW 0.88 0.94 0.94 0.98 0.02 1.88
male woolless lambs, using a database from tropical condition experi- EBW/SBW 0.66 0.80 0.80 0.91 0.05 6.10
ments, once there is no previous research which uses only castrated RE (kcal/kg −13.98 45.90 38.78 101.72 25.43 55.40
male woolless sheep in tropical conditions. In addition, this research EBW0.75/d)8
HP (kcal/kg 52.82 176.24 159.36 323.42 56.90 32.28
aimed to generate data to make possible the comparison about nutrient
EBW0.75/d)9
requirements between wool and woolless sheep in warm and tropical RP (g/d)10 −16.79 17.74 15.23 42.93 10.37 58.47
climates around the world and also to estimate the shrunk body weight MEI (kcal/kg 85.58 221.77 197.73 399.00 74.18 33.45
and empty body weight of the animals according to their fed body EBW0.75/d)11
weight. MPI (g/d)12 21.67 69.52 68.62 127.43 24.31 34.97

1
iSBW = final shrunk body weight;.
2. Materials and methods 2
iEBW = initial empty body weight;.
3
fBW = final body weight;.
2.1. Data acquisition 4
fSBW = final shrunk body weight;.
5
fEBW = final empty body weight;.
6
Data from three comparative slaughter experiments (n = 150) of SBG = shrunk body gain;.
7
castrated male woolless sheep was used. Experiments were carried out EBG = empty body gain;.
8
at the Universidade Federal Rural da Amazônia (UFRA - Federal Rural RE = retained energy;.
9
University of Amazonia) in Brazil. All experimental procedures were HP = heat production;.
10
approved by the Ethics Committee on Animal Use of UFRA (protocols RP = retained protein;.
11
MEI = metabolizable energy intake;.
03/2014, 04/2013 and 01/2012 for the Exp. 1, 2 and 3, respectively). 12
MPI = metabolizable protein intake.
The following information on each study were used: feeding treat-
ment, days in feeding, average intake of metabolizable energy intake
daily gain considering the requirements for late maturity sheep ac-
(MEI) and crude protein intake (CPI), body weight (BW), shrunk body
cording to NRC (2007). The experiment lasted 20 days for adaptation
weight (SBW), initial and final total body content of ether extract (EE)
and 63 for the feeding treatment. Additional description of methodo-
and crude protein (CP) of each animal. A descriptive statistics of the
logical procedures was presented by Cutrim et al. (2016).
main variables used is shown at Table 1. Main details of each experi-
ment were:
Experiment 1. Fifty-four castrated male lambs, of Santa Ines breed, 2.2. Estimation of weight relations
averaging 6 mo of age and 19.08 ( ± 2.76) kg were used. Nine lambs
were selected at random to compose the baseline group. The remaining At the end of experiments, the lambs were weighed at 3:00 pm (to
45 animals were distributed at random into three feeding treatments assess the BW) and again after a 16-hour solid fasting to assess the SBW.
and fed individually. All treatments were composed of 40% of roughage The animals were slaughtered the following morning. A similar pro-
and 60% of concentrate. The average energy content of the diets was tocol was followed in experiment 2. In experiment 3, when the animals
3.07 Mcal/kg in the dry matter (DM) basis in order to achieve 200 g/ reached 30 kg BW, those in the group with the highest intake (4.96%
day of average daily gain considering the requirements for late maturity BW) were slaughtered, and for every two animals slaughtered from this
sheep according to NRC (2007). The experiment lasted 14 days for group, one animal from the group with the lowest intake (3.4% BW)
adaptation and 63 for the feeding treatment. Additional description of was slaughtered. The remaining animals were subjected to a 16 h solid
methodological procedures was presented by Luz et al. (2017). fast and subsequently weighed again (to assess BW and SBW).
Experiment 2. Fifty-four castrated male lambs, of Santa Ines breed, The fits of three models were tested to estimate SBW as a function of
averaging 6 mo of age and 23.17 ( ± 2.39) kg were used. Nine lambs nonfasting (fed) BW and EBW as a function of SBW. The following
were selected at random to compose the baseline group. The remaining models were tested:
45 animals were distributed at random into three feeding treatments
and fed individually. All treatments were composed of 50% of roughage
Linear with intercept Y = aX + b
and 50% of concentrate. The average energy content of the diets was Linear without intercept Y = Ax
2.02 Mcal/kg in the DM basis in order to achieve 200 g/day of average Nonlinear Y = aXb
daily gain considering the requirements for late maturity sheep ac-
cording to NRC (2007). The experiment lasted 20 days for adaptation
and 63 for the feeding treatment. Additional description of methodo- The relationships between daily SBG, average daily gain (ADG), and
logical procedures was presented by Santos et al. (2016). empty body gain (EBG), were determined using the first derivative of
Experiment 3. Forty-two castrated male lambs, of Santa Ines breed, the models used to estimate the weight ratios as a function of the daily
averaging 4 mo of age and 19.79 ( ± 1.92) kg were used. Six lambs weight gain rate.
were selected at random to compose the baseline group. The remaining
36 animals were distributed at random into three feeding treatments 2.3. Energy and protein intake
and fed individually. All treatments were composed of 60% of roughage
and 40% of concentrate. The average energy content of the diets was In all experiments, feed intake was assessed daily by weighing and
2.43 Mcal/kg in the DM basis in order to achieve 250 g/day of average sampling the provided feed and leftovers. The sheep from each

2
S.d.S. Barcelos, et al.
experiment were subjected to digestibility trials. While under perfor-
mance evaluation, the animals were adapted to the collection bags for
three days, followed by a five-day fecal collection period.
For chemicals analysis, all samples (feed, orts and fecal samples)
were individually dried in a forced dry oven (65 °C) for 72 h and
grinded to 1 mm (Wiley mill; A. H. Thomas, Philadelphia, PA). Samples
were analyzed following AOAC (1990) methods, for DM (method
967.03), ash (method 942.05) and CP (method 981.10) contents, being
nitrogen concentration determined by the Kjeldahl method, with CP
content calculated as N × 6.25. The EE content was assessed using an
XT15 (Ankom®, Macedon, NY, USA) extractor using petroleum ether,
according to the American Oil Chemist's Society method (Am5-04;
AOCS, 2009). Neutral detergent fiber (NDF) and acid detergent fiber
(ADF) contents were analyzed according to Van Soest et al. (1991)
using heat stable α-amylase.
The animals’ digestible energy intake (DEI) was calculated by
multiplying the digestible fraction of each nutrient by its caloric value,
using the following equation NRC (2001):
DEI = 5.6 × dCPI + 9.4 × dEEI + 4.2 × dNDFapI + 4.2 × dNFCI (1)
where DEI = digestible energy intake (Mcal/day), dCPI = digestible
crude protein intake (kg/day), dEEI = digestible ether extract intake
(kg/day), dNDFapI = digestible ash- and protein-corrected neutral de-
tergent fiber intake (kg/day) e dCNFI = digestible non-fibrous carbo-
hydrates intake (kg/day). The DEI was then multiplied by 0.82
(NRC, 2000) to calculate the metabolizable energy intake (MEI). The
total digestible nutrient intake was calculated assuming that 1 kg total
digestible nutrients (TDN) equaled to 4.409 Mcal of digestible energy
(DE).
The microbial crude protein (CPmic) content of daily ration was
calculated by multiplying the TDN intake (kg/day) by the average
microbial efficiency value of 75 g CPmic/kg TDN intake obtained from
previous sheep studies (Bayão et al., 2014; Freire et al., 2014;
Santos et al., 2015; Souza, 2016; Alves Júnior et al., 2017;
Pereira, 2017; Santos Neta, 2017). The true fraction of CPmic was set to
80%, as was digestibility, represented by the coefficient of 0.64. The
rumen-undegradable protein (RUP) intake was estimated as the dif-
ference between the crude protein intake and the rumen degradable
protein (RDP) intake. The RUP digestibility was considered as 80%
(NRC, 2000).
Metabolizable protein intake (MPI) was estimated by adding the
CPmic and RUP intake (RUPI) from Eq. (2).
MPI = (CPmic × 0.64) + (RUPI × 0.80) (2)
where MPI = Metabolizable protein intake (g/day), CPmic = microbial
crude protein and RUPI = rumen-undegradable protein intake.
2.4. Slaughter and body composition
Pre-harvest animal care and handling procedures followed the
Brazilian Sanitary and Industrial Inspection Regulation for Animal
Origin Products (Brazil, 1997). Before slaughter, feed was withheld
from animals for 16 h, although they had ad libitum access to water the
entire time, performing a 16 h solid fasting. Slaughter was performed by
electrical stunning followed by exsanguination. After skinning and
eviscerating the animals, each body component (blood, skin, legs, head,
viscera and carcass) was individually weighed. The gastrointestinal
tract, bladder and gallbladder contents were discarded to calculate the
EBW, which was defined as the SBW minus the digestive tract contents.
Half the carcass, all organs, viscera, legs, head, skin and blood were
weighed, frozen, ground and homogenized, and 250-g samples were
subsequently collected per animal to analyze EE (Am5-04;
AOCS, 2009), CP (method 981.10; AOAC, 1990), ash (method 942.05;
AOAC, 1990) and water content.
Body retained energy (RE) was calculated from the protein and fat
content, using caloric values of 5.6405 and 9.3929 Mcal/kg,
3
Livestock Science 231 (2020) 103856
respectively (ARC, 1980). The relationship between SBW and EBW for
all animals was used to develop nonlinear regression equations within
each study to determine the initial EBW and initial body composition of
the remaining animals. The baseline animals of each experiment were
used to improve the accuracy of the estimation of initial body compo-
sition, especially at lighter weights.
After slaughtering the EBW and final body compositions were de-
termined. The total EBG and the protein, fat, ash and water in the EBG
were calculated based on the difference between final and initial body
compositions.
2.5. Energy requirement calculations
Heat production (HP) was calculated as the difference between MEI
and energy retention (Lofgreen and Garrett, 1968). The net energy re-
quired for maintenance (NEm) was estimated by nonlinear regression of
HP as a function of MEI trough the model described below (Ferrell and
Jenkins, 1998):
HP = a × e b × MEI (3)
0.75
where HP = heat production (kcal/kg EBW /day), a and b = coeffi-
cients, e = the natural logarithm and MEI = is ME intake (kcal/kg
EBW0.75/day).
The metabolizable energy required for maintenance (MEm, kcal/kg
EBW0.75/day) was calculated by iteration, assuming that MEm is the
value by which HP is equal to MEI. NEm divided by the MEm was
considered the ME use efficiency for maintenance (km).
Using the animals fed above maintenance data, a regression equa-
tion (Eq. (4)) was fitted, plotting RE as a function of the daily gain in
EBW and metabolic EBW (kg0.75) to determine the net energy re-
quirement for gain (NEg) as proposed by Chizzotti et al. (2008).
RE = a × EBW 0.75 × EBG b (4)
0.75
where RE = retained energy (kcal/kg EBW /day),
EBW0.75 = metabolic empty body weight, EBG = empty body gain (kg/
day), a = the antilog of the intercept and b = the slope of the linear
regression of the logarithm of RE (Mcal/kg0.75 of EBW/d) on the
logarithm of EBG.
The partial efficiency of ME utilization for growth (kg) was assumed
to be the slope of the linear regression of RE on MEI above the main-
tenance level, assuming that RE is null when MEI above main-
tenance = 0 (model intercept equal to 0). Each animal's MEIg value was
calculated as the difference between total metabolizable energy intake
and metabolizable energy required for maintenance (MEIg = MEI –
MEm).
RE = k g × MEIg (5)
where RE = retained energy (kcal/kg EBW 0.75/day), kg = efficiency of
use of ME to NE for growth and MEIg = MEI for gain (kcal/kg EBW 0.75/
day).
The metabolizable energy required for gain (MEg) was calculated by
dividing NEg by kg. The sum of MEm and MEg was considered the total
metabolizable energy requirement (MEt).
2.6. Protein requirement calculations
The metabolizable protein required for maintenance (MPm) was
calculated by linear regression with the intercept (Eq. (6)) between MPI
and EBG for performance and maintenance.
MPI = a + b × EBG (6)
where MPI= metabolizable protein intake, EBG = empty body gain
(kg/day) and a and b = regression parameters.
The intercept of this equation divided by the mean metabolic EBW
was considered as the MPm.
S.d.S. Barcelos, et al. Livestock Science 231 (2020) 103856

a Table 2
MPm =
EBW 0.75 (7) Summary of equations used to energy and protein requirements and adjust BW
of woolless sheep.
where MPm= requirement of metabolizable protein for maintenance
(g/kg EBW0.75/day), EBW0.75 = mean metabolic EBW, a = intercept of Variable to be Unit Relation Equation code
estimated
Eq. (6). Those calculations followed the methodology proposed by
Wilkerson et al. (1993). SBW1 Kg SBW = 0.938 × BW Eq. (11)
The net protein required for gain was calculated by fitting a retained SBG2 Kg SBG = 0.938 × ADG Eq. (12)
protein (RP) model as a function of RE and daily gain in EBW, using EBW3 Kg EBW = 0.057 × SBG 1,.135 Eq. (13)
EBG4 Kg EBG = SBG × 0.576× SBW 0.135
Eq. (14)
only performance animals per Eq. (8) (Chizzotti et al., 2008).
NEm kcal/ HP = 70.4 × e0.00402×MEI Eq. (15)
RP = a × EBG + b × RE (8) EBW0.75
RE5 kcal/d RE = 462 × EBW 0.75 × EBG 1.046 Eq. (16)
where RP= retained protein (g/day), a and b= parameters of the kg6 – RE = 0.407× MEIg Eq. (17)
MPm7 g/EBW0.75 MPI = 30.1 + 352 × EBG Eq. (18)
multiple regression between EBG (kg/day) = empty body gain and RP8 g/d PR = 182 × EBG – 0.005 × RE Eq. (19)
RE = retained energy (kcal/day), respectively. k9 – PR = 0.428 × MPIg Eq. (20)
The metabolizable protein use efficiency for gain (k) was considered K – k = 0.868 – 0.0204× EBW Eq. (21)
the slope of the regression curve between PR and the available meta-
1
bolizable protein intake for gain (MPIg). Each animal's MPIg was cal- SBW = shrunk body weight;.
2
SBG = shrunk body gain;.
culated as the difference between the total metabolizable protein intake 3
EBW = empty body weight;.
and metabolizable protein required for maintenance (MPIg = MPI – 4
EBG = empty body gain;.
MPm). The following model was fitted: 5
RE = retained energy;.
6
RP = k × MPIg kg = efficiency of energy utilization for growth;.
(9) 7
MPm = requirement of metabolizable protein for maintenance;.
8
RP = retained protein;.
where RP= retained protein (g/day), k = efficiency of use of meta- 9
k = efficiency of use of metabolizable protein for gain.
bolizable protein for gain, MPIg = metabolizable protein intake for gain
(g/day). The model presented at Eq. (9) was used to achieve a fixed k
3.1. Body weight relationships
(Valadares Filho et al., 2016). The k was considered a variable and
calculated by dividing RP by MPIg (k = RP/MPIg). This result was re-
The fit of a linear model without an intercept (AIC = −62.3) de-
lated to EBW using the following model:
scribed the relationship between BW and SBW better than the linear
k = a − −b × EBW (10) models with intercepts (AIC = −60.3) or nonlinear models
(AIC = −60.3). Based on the AIC evidence ratio, the linear model had a
The metabolizable protein requirements for gain (MPg) were cal- 73.1% probability of showing the best fit, versus 26.9% probability for
culated by dividing the NPg by k. Therefore, the total metabolizable the other models. The following final model was fitted to estimate the
protein requirements (MPt) were obtained using the following equa- SBW as a function of BW of growing castrated male woolless sheep:
tion: MP = MPm + MPg.
SBW = 0.938(± 0.002) × BW (Rxy = 0.993) (11)

2.7. Parameter estimation
The estimation of parameters followed the mixed models metho-
dology considering the study as the random effect for all analyses. The
MIXED procedure of SAS version 9.2 (SAS Inst., Inc., Cary, NC) was
used to estimate the parameters for linear models, and the NLMIXED
procedure was used to estimate the parameters for nonlinear models.
All models structure were described above (Eqs. (1)–(10)). All other
effects except study were used as fixed effects. The Gauss-Newton
method was used for convergence. For all analyses, fixed effects were
considered different from zero if the P-value was less than 0.05. The
random effect of study was maintained in all analyses.
2.8. Model comparisons
The fit of different model structures to the same database, when
necessary, was compared using the Akaike information criterion (AIC)
(Akaike, 1974). The probability of one model being a better fit than
another was calculated using the AIC evidence ratio (Motulsky and
Christopoulos, 2003), based on the absolute difference in AIC score
between the models, given by: AIC = 2 L + 2 K, wherein L is the natural
logarithm of the model's likelihood function with the parameters, and
K is the number of parameters. The model with the lowest AIC score
was considered the best fitted model.
3. Results and discussion
where SBW = fasting body weight and BW = body weight.
The linear model resulting from the relationship between FBW and
BW in this study showed that FBW accounted for 93.82% of the non-
fasting body weight, representing a fixed relationship of 6.18% fasting
losses. Similar results were found in studies applying nutritional models
to small ruminants, which defined FBW as 96% of the total sheep BW
(Cannas et al., 2004; Tedeschi et al., 2010).
To estimate the SBG as a function of ADG, the factor from Eq. (11)
was used because the weight gain rate is the first derivative of the
weight gain model:
(12)
SBG = 0.938 × ADG
where SBG = shrunk body gain (g/day) and ADG = average daily gain
(g/day).
Similar to SBW, the SBG accounted for 94% of the ADG. This
measure is calculated as the difference between two FBW weighings,
and the derivative of a constant is zero. Therefore, the same factor was
used.
The relationship between SBW and EBW was better described by a
nonlinear model (AIC = 12.5) than by linear models with (AIC = 13.7)
or without (AIC = 26.2) intercepts. The AIC evidence ratio showed
best-fit probabilities of 63.3% vs. 35.7% for the nonlinear model vs.
linear model with an intercept and of 100% vs. 0.0% for the nonlinear
model vs. linear model without an intercept. The following final model
was fitted to estimate the EBW as a function of SBW for growing ca-
strated male woolless sheep:
EBW = 0.507(± 0.052) × SBW 1.135(±0.028) (Rxy = 0.976) (13)

The summary of models generated in this study is shown at Table 2. where EBW = empty body weight and SBW = fasting body weight.

4
S.d.S. Barcelos, et al.
The nonlinear model observed in this study considered that the
weight ratios and weight gain rates varied according to the animal's
weight. Even after solid-fasting for more than 16 h, the animals still had
ingest in their gastrointestinal tracts (GIT). This study suggests a non-
linear relationship between SBW and EBW, with the decreased GIT
content proportional to the increased body weight. A nonlinear re-
lationship between SBW and EBW was also found for other ruminant
species (Gionbelli et al., 2015). Therefore, for growing animals such as
those in this study, feed accounted for a higher EBW loss in 20 kg an-
imals (25%) than in those that reached the mature weight (17% for
40 kg animals). Santa Ines sheep have adult weight between 32 and
50 kg (Malhado et al., 2009; Regadas Filho et al., 2013). This value does
not represent the maximum animal weight, the definition of an op-
timum mature weight depends on the species, breed and environmental
conditions (Malhado et al., 2009). In addition, the GIT sizes of the
animals in this study may have been larger than those of animals used
in international studies.
To estimate the average daily EBW gain rate, the result of the first derivative of
Eq. (13) as a function of SBG was used, generating the following equation:
EBG = SBG × 0.576 × SBW 0.135
where EBG = empty body weight gain (g/d), SBG = fasting average
daily weight gain (g/d) and SBW = fasting body weight.
Converting ADG into EBG without considering SBG seems in-
adequate because losses from fasting are disregarded, thus over-
estimating the net requirements by 6% in this study. Thus, EBG was
estimated using SBG and the nonlinear equation above (Eq. (14)). This
result showed that the ratio between EBG and SBG ranged from 0.82 to
0.90, for animals of 15 and 30 kg, with average daily weight gains
ranging from 100 to 300 g/day.
3.2. Energy requirements for maintenance
Fig. 1 shows the nonlinear relationship between HP and MEI. The
following general equation was used to calculate NEm:
HP = 70.4(± 2.1) × e 0.00402 (±0.00010) × MEI
The value estimated for NEm was 70.44 kcal/kg EBW0.75/day.
Considering a 95% confidence interval for this parameter [66.14;
74.74], values reported by studies using animals with similar char-
acteristics fell within the interval observed in this study (Regadas Filho
et al., 2013; Pereira et al., 2017). The conversion of NEm into the me-
tabolic SBW basis (SBW0.75) yielded 59.37 [56.11; 63.40] kcal/kg
BW0.75/day, which parameter confidence interval is within the re-
commended means of the main international systems for sheep
[NRC (2007): 62 kcal/kg SBW0.75/day; AFRC (1993): 62.2 kcal/kg
SBW0.75/day]. The value at which heat production equaled the MEI was
109.34 kcal/kg EBW0.75/day, which corresponded to the MEm, and thus
Livestock Science 231 (2020) 103856
the km was 0.644. The National Research Council (NRC, 2007) uses a
fixed value for energy use efficiency for maintenance, which is identical
to that observed in this study. Thus, a common value of 0.644 could be
established to km for animals at different climate conditions.
Besides influenced by factors related with the animals physiological
condition (NRC, 2007), the maintenance requirement in ruminants can
be also influenced by body composition, it means, by the relative
quantities of protein and fat being synthetized (Garrett, 1980). Protein
deposition is energetically less efficient than fat synthesis, because
muscle has a more intense activity (Garrett, 1980). Once Santa Ines
sheep were selected at semiarid region (characterized by longer dry
period with constant food shortage), a rapid fat tissue deposition is
expect as a survival strategy under these conditions (Regadas Filho
et al., 2011, 2013), it means, primarily in fat visceral tissues increasing
the cost of energy for maintenance (CSIRO, 2007). Besides that, higher
values of ME requirements were obtained by Salah et al. (2014) of
tropical ruminant livestock than the published values for either tem-
perate genotypes.
However, as demonstrated in this research, it seems that's there are
no effects of these parameters on Santa Ines sheep energy requirement
(14)
under tropical condition compared with temperate animals. These re-
sults corroborate to Regadas Filho et al. (2013) that differences between
responses when using international tables and those observed in
woolless sheep may not result from maintenance requirements.
3.3. Energy retention and metabolizable energy use efficiency for gain
All energy retained in the EBW of animals as fat and protein is
understood as the energy requirement for weight gain (Garrett et al.,
1959). Accordingly, to calculate the NEg, the fitted model was based on
the relationship between energy retention as a function of EBW and the
target EBG:
RE = 462(± 53) × EBW 0.75 × EBG1.046(±0.059) (16)
This model considers that variations in weight gain affect the re-
quirements for weight gain. This type of fit improves the model com-
(15)
pared with those that only consider a fixed gain composition, regardless
of weight gain rate.
Small adjustments to the model may be valuable for result accuracy,
as studies indicate that the method used is one of the sources of var-
iation in NEg values (Oliveira et al., 2017). The results show that the
concentration of energy in gain depends approximately 96% on body
weight and 4% on the weight gain rate. Therefore, two lambs weighing
15 kg and 30 kg who gain 100 g/day will have NEg values of 181 and
360 kcal/kg EBW0.75/day, respectively, as presented in Table 3.
The NEg values found are similar to Galvani et al. (2014) generated
using data from crossbred Santa Ines lambs of 15 kg BW. However, the
values found by Galvani et al. (2014) for lambs with BWs ranging from
Fig. 1. Nonlinear relationship between heat production (HP) and
metabolizable energy intake (MEI) of woolless sheep in feedlot. The
symbols represent data from experiment 1 (●), experiment 2 (□) and
experiment 3 (▲).
5
S.d.S. Barcelos, et al. Livestock Science 231 (2020) 103856

Table 3
Energy and protein requirements for woolless sheep in feedlot.
BW SBW1 EBW2 ADG NEm3 NEg4 TNE5 MEm6 MEg7 TMEt8 MPm9 MPg10 TMPt11

15 14.07 10.20 0.100 402.12 181.12 583.24 624.20 445.01 1069.20 18.21 19.90 38.11
0.150 402.12 276.75 678.86 624.20 679.97 1304.16 18.21 29.81 48.02
0.200 402.12 373.87 775.99 624.20 918.59 1542.79 18.21 39.71 57.92
0.250 402.12 472.11 874.23 624.20 1159.98 1784.18 18.21 49.60 67.81
0.300 402.12 571.27 973.38 624.20 1403.60 2027.80 18.21 59.49 77.70
20 18.76 14.14 0.100 513.71 240.98 754.68 797.42 592.08 1389.50 23.26 23.10 46.37
0.150 513.71 368.21 881.92 797.42 904.69 1702.11 23.26 34.60 57.86
0.200 513.71 497.43 1011.14 797.42 1222.18 2019.60 23.26 46.07 69.34
0.250 513.71 628.14 1141.85 797.42 1543.35 2340.77 23.26 57.54 80.80
0.300 513.71 760.06 1273.77 797.42 1867.48 2664.90 23.26 68.99 92.26
25 23.46 18.22 0.100 621.18 300.72 921.90 964.25 738.87 1703.12 28.13 27.27 55.41
0.150 621.18 459.50 1080.68 964.25 1128.99 2093.24 28.13 40.83 68.96
0.200 621.18 620.76 1241.94 964.25 1525.20 2489.45 28.13 54.35 82.48
0.250 621.18 783.88 1405.06 964.25 1926.00 2890.24 28.13 67.86 95.99
0.300 621.18 948.51 1569.69 964.25 2330.49 3294.74 28.13 81.36 109.49
30 28.15 22.41 0.100 725.49 360.38 1085.86 1126.15 885.45 2011.60 32.85 33.14 65.99
0.150 725.49 550.65 1276.14 1126.15 1352.96 2479.11 32.85 49.58 82.43
0.200 725.49 743.90 1469.39 1126.15 1827.77 2953.92 32.85 65.99 98.84
0.250 725.49 939.39 1664.87 1126.15 2308.07 3434.23 32.85 82.37 115.22
0.300 725.49 1136.67 1862.16 1126.15 2792.81 3918.96 32.85 98.73 131.58

1
SBW = shrunk body weight, kg (SBW=0.938×BW);.
2
EBW = empty body weight, kg (EBW=0.507×SBW1.135);.
3
NEm = net energy requirement for maintenance, kcal/kg EBW0.75/d (HP = 70.4 × e0.00402×MEI);.
4
NEg = net energy requirement for gain, kcal/d (RE = 462×EBW0.75×EBG1.046), where EBG = empty body gain (EBG=0.938×SBG×0.576×SBW0.135);.
5
TNE = total net energy, kcal/d (TNE = NEm+ NEg);.
6
MEm = metabolizable energy for maintenance, kcal/d (MEm = 109.3 kcal/kg EBW0.75/d);.
7
MEg = metabolizable energy for gain, kcal/d (MEg = NEg/0.41);.
8
TME = total metabolizable energy, kcal/d (TME = MEm+ MEg);.
9
MPm = metabolizable protein for maintenance, g/d (MPm = 3.19 g/kg EBW0.75/d);.
10
MPm = metabolizable protein for gain, g/d (MPg = NPg/k), where NPg = net protein for gain (RP = 182×EBW-0.005×RE), and k = efficiency of use of
metabolizable protein for gain (k = 0.868–0.0204×EBW);.
11
TMP = total metabolizable protein, g/d (TMP= MPm+ MPg).

25 to 30 kg were significantly lower than those of the present study (up
to 25% lower), even when considering differences in the fat deposition
rates of lambs in the same breed groups.
The kg value estimated using Eq (17) was 0.407 (Fig. 2). The kg
determination is important to convert the NEg into MEg, once it has
direct application on diets formulation for ruminants (Marcondes et al.,
2016). However, the results shown in this study are higher than those
reported in previous studies using the same approach (Oliveira et al.,
2017; Pereira et al., 2017).
RE = 0.407(± 0.259) × MEIg (17)
Using the same approach, a study with 1/2 Santa Ines × 1/2 Dorper
crossbred sheep fed low-quality roughage diets found a kg of 0.41
(Galvani et al., 2014). This is explained by the high visceral fat reten-
tion in those animals because the efficiency of energy retention in the
body depends on the proportions of energy retained as protein and fat,
and fat deposition efficiency is higher than protein deposition effi-
ciency. Because fat is the main energy reserve form for the animals and
has more calories than protein (ARC, 1980), the higher the fat gain
percentage, the higher its energy concentration, thus confirming this
study's findings Fig. 3.
Total nutrient requirements were calculated using the factorial
method adopted in most systems. Per the model used, MEt increases
with BW and ADG and may be lower or higher than recommended
NRC (2007), depending on body weight.
3.4. Protein requirements for maintenance
The relationship between metabolizable protein intake (MPI) and
empty body gain (EBG) is shown in Fig. 4. The following model was

Fig. 2. Relationship between retained energy (RE) and metabo-

lizable energy intake for gain (MEIg) in the three experiments. The
symbols represent data from experiment 1 (●), experiment 2 (□)
and experiment 3 (▲).

6
S.d.S. Barcelos, et al.
Fig. 3. Energy concentration of EBG (Mcal/kg of EBG) as a function of per-
centage of fat in the EBG.
used to estimate the MPm:
MPI = 30.1(± 2.1) + 352( ± 16) × EBG (18)
When dividing the intercept of Eq. (18) by the animals’ average
metabolic weight (9.45 kg), the MPm obtained was 3.19 g/kg EBW0.75/
day. Converting this value into BW, an estimate of 2.56 g/kg BW0.75/
day was obtained, which is higher than the values recommended by the
systems INRA (1988) and AFRC (1993) that set MPm to 2.51 and 2.18 g/
kg BW0.75/day. However, these systems estimated MPm by the sum of
nitrogen losses in the urine, feces and skin, divided by the metaboliz-
able protein use efficiency, which varies considerably among nutri-
tional systems (NRC, 2007).
The model used in this study considered the effect of MP intake on
animal growth dynamics and is indicated as an alternative to better
estimate the MPm (Watson et al., 2017), because it considers microbial
protein synthesis efficiency. Furthermore, this study considered that the
inefficiency of rumen-degraded N conversion into microbial N was
compensated by urea being recycled from the liver to the rumen and
salivary glands (Rotta et al., 2016).
3.5. Protein retention and metabolizable protein use efficiency for gain
PR = 182(± 16) × EBG − −0.005(± 0.004) × RE (19)
The following general equation was used to estimate NPg:
Tissue proportions are affected by body weight, age, genetic group,
energy intake level and sex. Accordingly Garrett (1980), the genetic
group would more strongly affect body composition than the nutri-
tional level. Most studies indicate decreases in net protein requirements
per kilogram of weight gain with increases in body weight
7
Livestock Science 231 (2020) 103856
(Rodrigues et al., 2016; Pereira et al., 2017), thus confirming this
study's findings. The results showed 13.13 and 13.61 g/day NPg for
animals with 15 and 30 kg live weight, respectively, with the same
weight gain rate (100 g/day), which corresponds to protein con-
centrations in weight gain of 170.06 and 160.54 g/kg EBG, respec-
tively. Therefore, with the increase of lambs body weight, a change in
weight gain composition occurs, being verified a decrease in protein but
an increase in fat deposition in its body. From a practical point of view,
this evidence indicate, that with higher body weight lambs production
tend to become more economically and nutritionally expensive, since
for the same weight gain a greater quantity of food will be required.
A static metabolizable protein use efficiency for a gain (k) of 0.427
was assessed using the model below and is represented in Fig. 5.
PR = 0.428(± 0.012) × MPIg (20)
The k depends on the amino acid profile quality in the digestible
RUP and on the ratio between digestible RUP and true digestible mi-
crobial protein in the small intestine (Rotta et al., 2016). However,
many other factors may affect the metabolizable protein use efficiency
for gain because NPg depends on the composition of gain and weight
gain rate; thus, these factors directly affect k. Therefore, another model
was fitted to estimate a variable k (Fig. 6), considering that it varies
with EBW (Eq. (21)), which was used to estimate the protein require-
ments for gain (Table 3).
k = 0.868(± 0.118) − −0.0204(± 0.0056) × EBW (21)
Although the AFRC (1993) consider the fixed efficiency of MP
conversion into gain of 0.59, the French system (INRA, 1988) has
considered that k decreases with the increase in body weight. This
decreasing efficiency with increasing body weight was confirmed by
Wilkerson et al. (1993). According to the model constructed in this
study, sheep at 15 kg BW (10.20 kg EBW) have an efficiency of 0.66,
whereas sheep at 30 kg BW (22.41 kg EBW) have an efficiency of 0.41.
The total protein requirements obtained here were higher than
those reported by a study using animals of similar age and breed
(Oliveira et al., 2017). This difference likely occurred because these
studies (NRC, 2007; Oliveira et al., 2017) set the k value to 0.70, which
may have underestimated the MPt recommendations and disregarded
that mature animals would less efficiently deposit protein as muscle.
The results of this study demonstrate that the recommended values
increase with body weight due to k, which decreases as body weight
increases.
The lack of studies similar to this study limits comparing data and
simultaneously demonstrates the need for updating studies on nutri-
tional requirements and nutrient efficiency of shorn sheep. The findings
did not support the hypothesis that woolless sheep have energy and
protein requirements lower than those wool lambs. The EBW, EBG and
Fig. 4. Relationship between metabolizable protein intake (MPI)
and empty body gain (EBG) of woolless sheep in feedlot. The
symbols represent data from experiment 1 (●), experiment 2
(□) and experiment 3 (▲).
S.d.S. Barcelos, et al.
the model chosen to estimate these parameters are crucial for the cor-
rect practical application of the requirement results. Based on that, it is
possible to say that further studies are necessary.
4. Conclusion
The models summary generated with this research can be used as a
tool to estimate the energy and protein requirements of woolless sheep
in warm and tropical climates around the world. In conclusion, the
energy requirements for maintenance and the metabolizable energy use
efficiency for maintenance found in this study did not differ to those
reported in international feeding systems. However, the nutritional
requirements of shorn crossbred sheep under tropical conditions do not
follow the pattern assumed by most nutritional systems for most vari-
ables estimated in this study. Thus the use of international feed system
standards to predict the nutritional requirements of woolless sheep
under tropical conditions may possibly affect the productive responses
observed. In addition, the results found in this study did not support the
hypothesis that woolless sheep have energy and protein requirements
lower than wool lambs and more studies similar to this are need to
investigate it.
Declaration of Competing Interest
The authors declare no conflict of interest.
Acknowledgments
We thank the Coordenação de Aperfeiçoamento de Pessoal de Nível
Superior (CAPES, Brazil) for providing financial support to Sandra S.
Barcelos.
8
Livestock Science 231 (2020) 103856
Fig. 5. Relationship between retained protein (RP) and metabolizable
protein intake available for gain (MPIg) of woolless sheep in feedlot. The
symbols represent data from experiment 1 (●), experiment 2 (□) and
experiment 3 (▲).
Fig. 6. Relationship between the efficiency of use of metabolizable
protein for gain (k) and empty body weight (EBW). The symbols re-
present data from experiment 1 (●), experiment 2 (□) and experiment 3
(▲).
Supplementary materials
Supplementary material associated with this article can be found, in
the online version, at doi:10.1016/j.livsci.2019.103856.
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