Small Ruminant Research 192 (2020) 106248

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Small Ruminant Research

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

Residual intake and body weight gain on the performance, ingestive

behavior, and characteristics of longissimus muscle of Dorper × Santa
Inês lambs
Eduardo Michelon do Nascimento a, Heloise Maggioni a, Carine Inês Schröter Bach a,
Willian Gonçalves do Nascimento a, Sergio Rodrigo Fernandes b, *, Américo Fróes Garcez Neto a
a
Federal University of Paraná, Palotina Campus, Department of Animal Science, Palotina, Paraná State, Brazil
b
State University of Londrina, Department of Animal Science, Londrina, Paraná State, Brazil

A R T I C L E I N F O A B S T R A C T

Keywords: The aim of this study was to evaluate the relationship between residual intake and body weight gain (RIG) and
Backfat thickness the performance, ingestive behavior, and characteristics of longissimus muscle of crossbred ½ Dorper × ½ Santa
Dry matter intake Inês lambs. Nineteen non-castrated male lambs, with a mean age of 4 months and a body weight (BW) of
Feed efficiency
23.04 ± 3.16 kg, were individually fed ad libitum in a feedlot for 90 days. The ration was composed of 640 g/kg
Feed conversion ratio
Cynodon hay and 360 g/kg concentrate feed; it contained 158.6 g/kg crude protein and 2.12 Mcal/kg metabo­
Rumination
Sheep lizable energy on a dry matter (DM) basis. At the end of the trial, lambs were classified as efficient (n = 7),
intermediate (n = 6), or inefficient (n = 6) for RIG based on ± 50 % of the standard deviation for this trait. The
lowest and highest values for dry matter intake relative to BW (DMIBW) were verified in the efficient and inef­
ficient groups, respectively (3.13 vs. 3.43 % BW/day). The feed conversion ratio (FCR) decreased by 15.94 %
(8.47 to 7.12 kg DM/kg gain) from the inefficient to the efficient group. The time spent feeding, in rumination,
and in idleness, as well as the intake rate (IR), were similar among RIG groups, but the rumination rate (RR) was
lower in the efficient group (1.61 g DM/min) compared with the intermediate and inefficient groups (1.82 g DM/
min on average). Average particle size (APS) and chemical composition of the orts were not influenced by RIG
and were similar among the efficiency groups. Backfat thickness, as measured by ultrasound (BFTUS) at the start
and end of the trial, and backfat thickness and loin eye area, as measured directly from the longissimus muscle
(BFTLE and LEA, respectively) after slaughter, were similar among RIG groups. Improving the efficiency for RIG
leads to better feed utilization, without affecting the characteristics of longissimus muscle in lambs.

1. Introduction requirements (Paula et al., 2013; Nascimento et al., 2016). Therefore,

The use of biologically efficient animals may increase the profit­
ability of sheep production, because feeding costs represent about 70–75
% of the total cost of production (Liu et al., 2000). Some feed efficiency
traits used in the selection of beef cattle have been applied to sheep, such
as feed conversion ratio (FCR) and gross feed efficiency (GFE), in which
the former trait is the ratio between dry matter intake (DMI) and average
daily gain (ADG), and the latter trait is the inverse ratio between these
traits. Although FCR and GFE have been widely used as measures of feed
efficiency, selection based on these traits results in animals with large
body sizes, a factor that increases the maintenance and growth
other measures have been proposed to fix these problems, such as re­
sidual feed intake (RFI) and residual body weight gain (RG).
Both RFI and RG were proposed by Koch et al. (1963); RFI is
calculated as the difference between the observed DMI and estimated
dry matter intake (DMIEST) from the mean metabolic weight (MMW) and
ADG. This trait allows for the identification of animals with low DMI for
the same level of production (weight gain). On the other hand, RG is
calculated as the difference between the observed ADG and the esti­
mated average daily gain (ADGEST) from the MMW and DMI. Thus, this
trait allows for the identification of animals with high ADG at the same
level of feed intake or the average DMI verified in the herd.

* Corresponding author.
E-mail addresses: edu.vetufpr@gmail.com (E.M. do Nascimento), helo_maggioni@hotmail.com (H. Maggioni), carineinesbach@yahoo.com.br (C.I.S. Bach),
williangon1974@gmail.com (W.G. do Nascimento), srfernandes83@gmail.com (S.R. Fernandes), americo.garcez@ufpr.br (A.F. Garcez Neto).

https://doi.org/10.1016/j.smallrumres.2020.106248
Received 19 June 2019; Received in revised form 28 August 2020; Accepted 22 September 2020
Available online 29 September 2020
0921-4488/© 2020 Elsevier B.V. This article is made available under the Elsevier license (http://www.elsevier.com/open-access/userlicense/1.0/).
E.M. do Nascimento et al. Small Ruminant Research 192 (2020) 106248

To simultaneously explore the advantages of RFI and RG, Berry and Table 1
Crowley (2012) proposed a combination of both traits in a single index; Ingredients and chemical composition of the diet provided to the
they defined it as residual intake and body weight gain (RIG). Addi­ crossbred ½ Dorper × ½ Santa Inês lambs during the trial.
tionally, this efficiency trait has been evaluated in recent studies with Componenta Content
growing cattle (Favero, 2014; Fernandes, 2014; Grion et al., 2014) and Ingredient (g/kg DM)
finishing cattle (Nascimento et al., 2016; Takeda et al., 2018). However, Cynodon hay 640.3
Lima et al. (2017); Carneiro et al. (2019) and Montelli et al. (2019) Ground corn 158.6
published the first studies that evaluated RIG in sheep; the authors Dried whey permeate 99.6
Soybean meal 68.5
verified differences in performance and feed efficiency traits between
Urea 17.2
lambs classified as efficient and inefficient in terms of RIG, but there Ammonium sulfate 2.2
were no changes in the carcass traits between these groups. Regarding Mineral premixb 9.0
the few studies available in the literature, the relationship between RIG Limestone 3.8
and performance and physiological traits, as well as the ingestive Rumimpex® c 0.2
Elitox® d 0.7
behavior of sheep, is not well understood. Thus, the aim of this study was Chemical composition
to evaluate the influence of RIG on the performance, ingestive behavior, Dry matter (g/kg as fed) 881.1
and characteristics of longissimus muscle of crossbred ½ Dorper × ½ Crude protein (g/kg DM) 158.6
Santa Inês lambs in the growing and finishing phases. Ether extract (g/kg DM) 18.4
Neutral detergent fiber (g/kg DM)e 509.8
Acid detergent fiber (g/kg DM) 254.3
2. Material and methods Lignin (g/kg DM) 48.1
Non-fiber carbohydrates (g/kg DM) 241.8
All procedures involved in this research were performed in accor­ Ash (g/kg DM) 71.6
dance with the National Council of Animal Experimentation Control Total digestible nutrients (g/kg DM) 652.3
Digestible energy (Mcal/kg DM) 2.59
(Conselho Nacional de Controle de Experimentação Animal − CONCEA) and Metabolizable energy (Mcal/kg DM) 2.12
were approved by the Animal Care and Use Committee (Comitê de Ética
a
no Uso de Animais – CEUA) of the Federal University of Paraná (Uni­ DM: dry matter.
b
Guaranteed analysis: 110 g/kg calcium, 55 g/kg phosphorus, 16 g/
versidade Federal do Paraná – UFPR), Palotina Campus, under protocol
kg magnesium, 93 g/kg sodium, 200 mg/kg cobalt; 1400 mg/kg iron;
number 013/2017-CEUA/Palotina.
550 mg/kg fluorine, 150 mg/kg iodine, 1500 mg/kg manganese,
The trial was carried out at the Center of Studies in Small Ruminants
20 mg/kg selenium, 5000 mg/kg zinc.
(Centro de Estudos em Pequenos Ruminantes – CEPER) located on the c
Monensin sodium.
Palotina Campus/UFPR, Palotina, western region of Paraná state, Brazil. d
Micotoxins eliminator.
The climate of this region is defined as subtropical, and it is character­ e
Corrected for ash and protein.
ized as Cfa according to the Köppen classification. The trial period was
from April to July, and the mean ambient temperature and relative length to improve its utilization by the lambs and reduce waste in the
humidity during the trial were 18.1 ◦ C and 74.5 %, respectively. feeder. The ration was supplied as total mixed ration (TMR) and split
into two daily meals (0700 h and 1600 h). Lambs were fed ad libitum
2.1. Animals and management during the adaptation period and the trial, maintaining the orts at 10 %
of the amount of feed provided. Adjustments to the amount of ration
Nineteen non-castrated male crossbred ½ Dorper × ½ Santa Inês supplied were performed daily based on the amount of orts. Samples of
lambs, with an average age of 4 months and body weight (BW) of hay and concentrate feed were collected every week to monitor DM
23.04 ± 3.16 kg (mean ± standard deviation [SD]), were used. The an­ content and to adjust the amount of ration provided to the lambs.
imals were allocated to individual covered pens, with a slatted floor and The chemical analysis of the feedstuffs was performed at the Labo­
an area of 1.5 m2. Each pen had an individual feeder and water drinker. ratory of Animal Nutrition of the Palotina Campus/UFPR. Samples were
The lambs were kept in the pens for 104 days, in which the first 14 days collected on days 8, 35, and 80 of the trial; dehydrated in a forced
were an adaptation period to the experimental management, and the last ventilation oven at 55 ◦ C until they reached a constant weight (after
90 days comprised the trial period. approximately 72 h of drying); and subsequently processed in a Wiley
Anthelmintic treatment (levamisole 10 %, at a dose of 1 mL/20 kg knife mill with a 1 mm mesh sieve. The contents of DM, crude protein
BW; and nitroxinil 34 %, at a dose of 1.5 mL/50 kg BW) was adminis­ (CP), ether extract (EE), lignin, and ash were determined according to
tered subcutaneously to all lambs at the first day of the adaptation the procedures of the Association of Official Analytical Chemists (AOAC,
period. Subsequently, parasitological monitoring was performed every 1990). The content of neutral detergent fiber (NDF) and acid detergent
21 days by assessing the fecal egg count (FEC) using the modified fiber (ADF) were determined using nonwoven bags and following the
McMaster technique (Gordon and Whitlock, 1939) and the degree of procedures described by Van Soest et al. (1991). The contents of ash and
anemia according to the Famacha® method (Molento et al., 2004). The neutral detergent insoluble nitrogen (NDIN; AOAC, 1990) were deter­
anthelmintic treatment was repeated in the lambs that presented a mined in the NDF residue to correct this fraction for ash and protein. The
FEC ≥ 600 and a Famacha score ≥ 3 during the trial. The treatment non-fiber carbohydrates (NFC) content was calculated as proposed by
protocol of infected animals after starting the trial was set up with the Hall et al. (1999), using the equation: NFC =
aim to obtain the same FEC and Famacha scores between treated and 100 − (CP + EE + NDF + Ash). The digestible fractions of CP, EE, NDF,
non-treated lambs. and NFC were estimated from equations adapted for growing and fin­
The diet was formulated to obtain accurate feed efficiency mea­ ishing cattle raised under Brazilian conditions, as proposed by Detmann
surements during a minimum feedlot period of 84 days (Castilhos et al., et al. (2006a, 2006b, 2006c, 2007). These fractions were used for the
2011), and to allow the lambs to reach approximately 35 kg BW at the calculation of total digestible nutrients (TDN), digestible energy (DE),
end of the trial, which is the average BW for slaughter in the western and metabolizable energy (ME), which were determined according to
region of Paraná state (Rozanski et al., 2017). Thus, the ration was Weiss et al. (1992).
formulated to meet the requirements for an ADG of 150 g/day (NRC,
2007). It was composed of 640 g/kg Cynodon hay and 360 g/kg
concentrate feed, on a dry matter (DM) basis (Table 1). Hay was ground
in a stationary shredder into particles that were approximately 3 cm in

2
E.M. do Nascimento et al.
2.2. Performance and feed efficiency traits
The daily DMI was calculated as the difference between the amount
of DM in the provided feed and in the orts. Each day, the orts of each
animal were collected before the morning meal, weighed, and stored in
plastic bags. At the end of each week, composite samples (approximately
250 g) were formed from the orts collected over the previous 7 days.
These samples were dehydrated in a forced ventilation oven at 55 ◦ C
until they reached a constant weight to determine their DM content. The
mean DM content calculated from all composite samples collected
during the trial was used to adjust the DMI of each animal.
Lambs were weighed every 21 days, early in the morning after
fasting (withdrawal of feed for 12 h). ADG was calculated by a linear
regression of BW during the experimental days on the weighing dates
(days 0, 21, 42, 63, and 90), as described by Arthur et al. (2001).
The DMI, ADG, and mid-test BW were used for the calculation of the
following traits: DMI relative to BW (DMIBW; % BW/day), calculated as
the ratio between DMI (kg/day) and mid-test BW (kg), multiplied by
100; DMI relative to MMW (DMIMMW; % MMW/day), calculated as the
ratio between DMI (kg/day) and MMW (MMW = mid-test BW0.75),
multiplied by 100; feed conversion ratio (FCR; kg DM/kg gain), calcu­
lated as described by Arthur et al. (2001); relative growth rate (RGR;
%/day), calculated as proposed by Fitzhugh Junior and Taylor (1971);
the Kleiber ratio (KR; g gain/kg0.75), calculated as described by Kleiber
(1947); residual feed intake (RFI; g DM/day), calculated as the differ­
ence between DMI (g/day) and DMIEST (g/day) from the MMW and ADG
(Koch et al., 1963); and residual body weight gain (RG; g BW/day),
calculated as the difference between ADG and ADGEST from the MMW
and DMI (Koch et al., 1963). The DMIEST and ADGEST were calculated for
each lamb using the following equations:
DMIEST = − 142.09434 + 65.17440 MMW + 2.20423 ADG R2 = 0.87 and
ADGEST = − 12.85143 − 2.52869 MMW + 0.18160 DMI R2 = 0.71
The residual intake and body weight gain (RIG) index was calculated
as proposed by Berry and Crowley (2012), using the formula:
RIG = [( − 1) × RFI ] + RG
Before the calculation, RFI and RG were standardized to a variance of
1 such that both traits had equivalent weights in the composition of RIG.
2.3. Ingestive behavior traits
Ingestive behavior was assessed by visual observation for 24 h on
days 9, 37, 71, and 86 of the trial. Behavioral evaluations were made by
eight trained observers, who were divided into four pairs; each pair
evaluated the lambs for 6 h. Feeding, rumination, and idleness activities
were identified and recorded every 10 min, according to the method of
Martin and Bateson (1993). Artificial lighting was used inside the barn
during the night and at dawn, and lambs were adapted to this condition
one week prior to the assessments of ingestive behavior. The results are
expressed as a percentage of the day (24 h) spent performing each
activity.
The intake rate (IR) and rumination rate (RR) were calculated in the
manner proposed by Cardoso et al. (2006). The IR was calculated as the
ratio between DMI (g/day) and the time spent feeding (min/day), and
RR was calculated as the ratio between DMI (g/day) and the time spent
in rumination (min/day). The results of both traits are expressed in g
DM/min.
Dietary particle selection was evaluated in the composite samples of
orts from each lamb. Composite samples (approximately 250 g) were
made from the orts collected during the 7 days prior to each evaluation
of ingestive behavior. These samples were fractionated using the Penn
State Particle Separator (PSPS), which is composed of three sequential
sieves with different mesh diameters (19.0, 7.8, and 1.7 mm) and a
3
Small Ruminant Research 192 (2020) 106248
bottom. Thus, four layers were obtained: L1 – particles larger than
19.0 mm; L2 – particles larger than 7.8 mm and smaller than 19.0 mm;
L3 – particles larger than 1.7 mm and smaller than 7.8 mm; and L4 –
particles smaller than 1.7 mm. Each particle layer was weighed, and
their proportions relative to the fractionated sample were used to
calculate the average particle size (APS, mm) of the orts, according to
the methodology described by Heinrichs and Kononoff (2002). After the
fractionation, the samples were reconstituted and subjected to chemical
analysis following the same procedures used in the analyses of
feedstuffs.
2.4. Longissimus muscle measurements
Backfat thickness (BFTUS, mm) was assessed by ultrasound on the
first and last days of the trial. The images were captured by one trained
individual using a Mindray® ultrasound (model DP-6600 Vet) equipped
with a linear transducer (10 cm in length and frequency of 5 MHz).
Lambs were restrained in a standing position and, after shaving the
lamb’s wool and cleaning the skin on the left side of the region between
the 12th and 13th thoracic vertebrae, the ultrasound images of the
longissimus muscle were captured using the same procedure described by
Rozanski et al. (2017). At the end of the trial, all images were transferred
to a computer and analyzed using the software Corel Draw®, version X8.
The images were analyzed by the same individual who operated the
ultrasound. The gain in BFTUS (mm) was calculated as the difference
between the initial and final BFTUS.
On the last day of the trial, the lambs were fasted (withdrawal of feed
for 14 h), weighed to record the body weight at slaughter (SW), and sent
to a commercial slaughterhouse in the region. The slaughter was con­
ducted in accordance with good animal welfare practices, in which the
animals were stunned, followed by bleeding (performed by severing the
jugular veins and carotid arteries), skinning, and evisceration. After
slaughter, the carcasses were identified, suspended by the metatarsal
joints, and transferred to a cold room at 4 ◦ C, where they remained for
24 h. After cooling, the carcasses were cut into two halves in the caudal-
cranial direction, with the left half sectioned into six commercial cuts
according to the Brazilian standard of lamb carcass cuts (Cezar and
Sousa, 2007): shoulder, leg, loin, ribs, breast + flank, and neck.
In the dorsal portion of the longissimus muscle of the loin, at the cutoff
point between the 13th thoracic vertebra and the 1 st lumbar vertebra,
the minimum and maximum backfat thickness on the loin eye (BFTLE,
mm), the maximum width (LEW, cm) and maximum depth (LED, cm) of
the loin eye, and the loin eye area (LEA, cm2) were measured, as
described by Garcia et al. (2003). The minimum and maximum BFTLE
were obtained using a digital caliper, and both measures were used to
calculate the mean BFTLE. The contour of the transverse section of the
longissimus muscle was traced on a plastic film and scanned to a com­
puter for LEA measurement using QUANT software (Vale et al., 2003).
The LEW and LED were obtained from the contour of the loin eye traced
on plastic film, using a ruler with a scale in millimeters. The ratio be­
tween both measures (W:D ratio) was also calculated.
2.5. Efficiency groups
At the end of the trial, the lambs were classified into three efficiency
groups with the limits established as ± 50 % of the SD for RIG. Thus,
lambs were classified as efficient (RIG > + 0.50 SDRIG ; n = 7), inter­
mediate (− 0.50 SDRIG ≤ RIG ≤ + 0.50 SDRIG ; n = 6), or inefficient
(RIG < − 0.50 SDRIG ; n = 6) for this trait. Due to the increase of endo­
parasite infection, the anthelmintic treatment had to be repeated in one
lamb from the intermediate group and two lambs from the efficient
group at day 17, and in one lamb from efficient group at day 53 of the
trial.
E.M. do Nascimento et al. Small Ruminant Research 192 (2020) 106248

2.6. Statistical analyses
Data were analyzed by analysis of variance (ANOVA) with respect to
the efficiency groups based on RIG (PROC GLM). The analyses were
performed following a completely randomized design, including initial
BW as a covariate (Model 1), according to the model:
Yij = μ + Gi + βBW + εij
where Yij = the value of the dependent variable from the jth animal in
the ith group adjusted for the initial BW of jth animal; μ = the mean
value of dependent variable (constant); Gi = the effect of the ith group; β
= the regression coefficient for initial BW; BW = the initial BW of the jth
animal; and εij = random error. When initial BW was not significant
(P > 0.05), this covariate was withdrawn from the model and the data
were analyzed by ANOVA only with respect to the efficiency groups
(Model 2). The means were fitted to the statistical models and compared
between efficiency groups by Tukey–Kramer test (PROC LSMEANS).
A Pearson correlation analysis (PROC CORR) was performed be­
tween RIG and all the evaluated characteristics. When the initial BW was
significant in the ANOVA, the analysis was performed using the partial
option to adjust the correlation coefficients for this covariate.
A significance level of 0.05 was adopted for all statistical analyses,
which were all carried out in Statistical Analysis System software,
version 9.0 (SAS Institute, 2002).
3. Results
3.1. Performance and feed efficiency
The RFI, RG, and RIG had mean values of 0.0, and SDs of 51.1 g DM/
day, 14.7 g BW/day and 1.8, respectively. The RIG had a high correla­
tion with RFI (r = -0.90) and RG (r = 0.90; Table 2); the minimum and
maximum values observed for these traits were -89.4 and 81.2 g DM/day
for RFI; -32.7 and 21.1 g BW/day for RG; and -3.3 and 3.2 for RIG.
Given that RIG was highly correlated with RFI and RG, both these
traits differed among RIG groups (Table 2). The RFI value was the lowest
in the efficient group and the highest in the inefficient group (-50.6 vs.
55.3 g DM/day), and RG values were the lowest and highest in the
inefficient and efficient groups, respectively (-16.4 vs. 10.7 g BW/day).
Given that RIG was used to define the efficiency groups, this trait pre­
sented the lowest value in the inefficient group and the highest value in
the efficient group (-2.2 vs. 1.7). The intermediate group differed from
the other groups in terms of RFI and RIG, with values of 3.8 g DM/day
and 0.2, respectively. However, the RG of the intermediate group (3.9 g
BW/day) was statistically similar to the efficient group.
The initial and final BW, MMW, ADG, and DMI had no correlation
with RIG and did not differ among RIG groups (Table 2), with mean
values of 23.04 kg, 34.57 kg, 12.41 kg0.75, 128.1 g/day, and 952.2 g/
day, respectively. The DMIBW tended to present a negative and moderate
correlation with RIG (r = -0.45; P = 0.0547); it also tended to differ
(P = 0.0614) among RIG groups, with the lowest and highest values in
the efficient and inefficient groups, respectively (3.13 vs. 3.43 % BW/
day; Table 2). The DMIMMW had no correlation with RIG, but it was
lower in the efficient group (7.19 % BW0.75/day) compared with the
intermediate and inefficient groups, which presented similar values
(7.89 % BW0.75/day on average; Table 2).
RIG had a high correlation with the FCR (r = -0.74), which decreased
by around 15.70 % between the inefficient group (8.47 kg DM/kg gain)
and efficient and intermediate groups (7.14 kg DM/kg gain on average;
Table 2). Thus, efficient and intermediate lambs consumed 1.33 kg DM/
kg gain less than the inefficient lambs. The RGR and KR had no corre­
lation with RIG, and they did not differ among RIG groups (Table 2); the
mean values for these traits were 0.20 %/day and 10.27 g gain/kg0.75,
respectively.
3.2. Ingestive behavior
For ingestive behavior, the proportion of time spent feeding, in
rumination, and in idleness had no correlation with RIG; they also did
not differ among RIG groups (Table 3); the mean values for these traits
were 22.18, 37.89, and 39.93 %/day, respectively. The IR and RR had no
correlation with RIG, but the RR was lower in the efficient group (1.61 g
DM/min) compared with the intermediate and inefficient groups, which
presented similar values (1.82 g DM/min on average; Table 3). The IR
did not differ among RIG groups (Table 3); it presented a mean value of
3.01 g DM/min.
With regard to the ort characteristics, the proportion of particles in
each layer obtained from the PSPS and the APS had no correlation with
RIG and did not differ among RIG groups (Table 4). The mean values for
the proportions of particles found in each layer were 45.06 % for par­
ticles larger than 19.0 mm, 12.91 % for particles ranging from
7.8–19.0 mm, 30.14 % for particles ranging from 1.7 to 7.8 mm, and
11.90 % for particles smaller than 1.7 mm; the mean APS was 10.87 mm.

Table 2
Performance and feed efficiency traits of crossbred ½ Dorper × ½ Santa Inês lambs in the three efficiency groups based on residual intake and body weight gain.
P-value
Pearson correlation RIG group
Variablea Model 1 Model 2

rRIG P-value Inefficient Intermediate Efficient IBW RIG group RIG group

RIGc – – − 2.2 ± 0.3 c 0.2 ± 0.3 b 1.7 ± 0.3 a 0.9018 <0.0001 <0.0001
RFI (g DM/day)c − 0.90 <0.0001 55.3 ± 10.5 a 3.8 ± 10.5 b − 50.6 ± 9.7 c 0.2087 <0.0001 <0.0001
RG (g BW/day)c 0.90 <0.0001 − 16.4 ± 3.7 b 3.9 ± 3.7 a 10.7 ± 3.5 a 0.3956 0.0004 0.0002
IBW (kg)c − 0.12 0.6155 23.30 ± 1.33 23.89 ± 1.33 22.09 ± 1.23 – – 0.5997
FBW (kg)b 0.42 0.0851 33.37 ± 0.85 35.66 ± 0.86 34.68 ± 0.81 <0.0001 0.1974 0.4101
MMW (kg0.75)b 0.43 0.0767 12.21 ± 0.14 12.60 ± 0.14 12.43 ± 0.13 <0.0001 0.1741 0.4901
ADG (g/day)b 0.42 0.0851 114.8 ± 9.5 140.2 ± 9.6 129.4 ± 8.9 0.0365 0.1974 0.2017
DMI (g/day)b − 0.16 0.5133 962.3 ± 36.5 994.0 ± 36.9 900.4 ± 34.4 0.0004 0.2072 0.1486
DMIBW (% BW/day)c − 0.45 0.0547 3.43 ± 0.09 3.36 ± 0.09 3.13 ± 0.08 0.1170 0.0283 0.0614
DMIMMW (% MMW/day)c − 0.40 0.0900 7.89 ± 0.21 a 7.88 ± 0.21 a 7.19 ± 0.20 b 0.6941 0.0677 0.0420
FCR (kg DM/kg gain)c − 0.74 0.0003 8.47 ± 0.33 a 7.15 ± 0.33 b 7.12 ± 0.31 b 0.9834 0.0196 0.0148
RGR (%/day)c 0.40 0.0880 0.18 ± 0.01 0.21 ± 0.01 0.20 ± 0.01 0.5868 0.2796 0.2721
KR (g gain/kg0.75)c 0.39 0.1005 9.42 ± 0.64 11.06 ± 0.64 10.32 ± 0.59 0.7959 0.2472 0.2203

Means in the same row followed by different lowercase letters differ by Tukey-Kramer test (P < 0.05).
a
DM: dry matter; BW: body weight; RIG: residual intake and body weight gain; RFI: residual feed intake; RG: residual body weight gain; IBW: initial BW; FBW: final
BW; MMW: mean metabolic weight; ADG: average daily gain; DMI: dry matter intake; DMIBW: DMI relative to the BW; DMIMMW: DMI relative to the MMW; FCR: feed
conversion ratio; RGR: relative growth rate; KR: Kleiber ratio.
b
Pearson correlation coefficient adjusted to IBW, and means and standard errors adjusted to the Model 1.
c
Pearson correlation coefficient non-adjusted to IBW, and means and standard errors adjusted to the Model 2.

4
E.M. do Nascimento et al. Small Ruminant Research 192 (2020) 106248

Table 3
Characteristics of ingestive behavior of crossbred ½ Dorper × ½ Santa Inês lambs in the three efficiency groups based on residual intake and body weight gain.
P-value
Pearson correlation RIG groupa
a
Variable Model 1 Model 2

rRIG P-value Inefficient Intermediate Efficient IBWa RIG group RIG group

Feeding (%/day)c − 0.24 0.3245 23.44 ± 1.05 22.08 ± 1.05 21.01 ± 0.97 0.7841 0.2752 0.2632
Rumination (%/day)c 0.30 0.2172 37.07 ± 0.99 37.96 ± 0.99 38.64 ± 0.92 0.6626 0.5037 0.5180
Idleness (%/day)c − 0.02 0.9196 39.49 ± 1.34 39.96 ± 1.34 40.35 ± 1.24 0.9131 0.9107 0.8954
IR (g DM/min)b − 0.02 0.9283 2.90 ± 0.17 3.13 ± 0.17 3.01 ± 0.16 0.0063 0.6398 0.5265
RR (g DM/min)b − 0.37 0.1184 1.81 ± 0.06 a 1.82 ± 0.06 a 1.61 ± 0.06 b 0.0003 0.0414 0.0522

Means in the same row followed by different lowercase letters differ by Tukey-Kramer test (P < 0.05).
a
RIG: residual intake and body weight gain; IBW: initial body weight; IR: intake rate; RR: rumination rate; DM: dry matter.
b
Pearson correlation coefficient adjusted to IBW, and means and standard errors adjusted to the Model 1.
c
Pearson correlation coefficient non-adjusted to IBW, and means and standard errors adjusted to the Model 2.

Table 4
Characteristics of orts from the diet provided to crossbred ½ Dorper × ½ Santa Inês lambs in the three efficiency groups based on residual intake and body weight gain.
P-value
Pearson correlation RIG groupa
Variablea, b
Model 1 Model 2

rRIG P-value Inefficient Intermediate Efficient IBWa RIG group RIG group

Particles size
L1 (%) − 0.11 0.6618 49.78 ± 7.63 39.54 ± 7.63 45.85 ± 7.06 0.2831 0.5548 0.6399
L2 (%) − 0.12 0.6227 13.03 ± 1.59 13.84 ± 1.59 11.86 ± 1.47 0.1462 0.4420 0.6591
L3 (%) 0.24 0.3200 26.17 ± 4.61 32.93 ± 4.61 31.31 ± 4.27 0.2398 0.5314 0.5651
L4 (%) − 0.03 0.9036 11.03 ± 3.79 13.69 ± 3.79 10.98 ± 3.50 0.8925 0.8440 0.8428
APS (mm) − 0.01 0.9566 11.69 ± 1.90 9.50 ± 1.90 11.41 ± 1.76 0.3320 0.5798 0.6771
Chemical composition
CP (g/kg DM) 0.40 0.0932 130.0 ± 13.0 161.3 ± 13.0 157.0 ± 12.0 0.7339 0.2310 0.2096
EE (g/kg DM) 0.13 0.5947 16.3 ± 2.0 18.3 ± 2.0 18.4 ± 1.9 0.5135 0.7394 0.7147
NDF (g/kg DM) − 0.27 0.2605 685.2 ± 43.6 648.7 ± 43.6 613.5 ± 40.4 0.5962 0.4718 0.4978
ADF (g/kg DM) − 0.21 0.3938 325.8 ± 25.7 309.8 ± 25.7 290.2 ± 23.8 0.8207 0.6067 0.6031
Lignin (g/kg DM) − 0.32 0.1749 71.3 ± 5.0 67.2 ± 5.0 60.9 ± 4.7 0.9094 0.3847 0.3383
NFC (g/kg DM) 0.21 0.3809 82.3 ± 31.2 87.0 ± 31.2 129.0 ± 28.9 0.4338 0.4157 0.4882
Ash (g/kg DM) − 0.19 0.4356 86.1 ± 2.6 84.8 ± 2.6 82.2 ± 2.4 0.0608 0.6823 0.5331
TDN (g/kg DM) 0.37 0.1179 591.1 ± 10.1 593.9 ± 10.1 615.3 ± 9.4 0.4048 0.2756 0.1860
DE (Mcal/kg DM) 0.35 0.1360 2.19 ± 0.07 2.27 ± 0.07 2.34 ± 0.06 0.8085 0.3147 0.2674
ME (Mcal/kg DM) 0.35 0.1362 1.80 ± 0.05 1.86 ± 0.05 1.92 ± 0.05 0.8023 0.3182 0.2705
a
RIG: residual intake and body weight gain; IBW: initial body weight; L1: particles larger than 19 mm; L2: particles larger than 7.8 mm and smaller than 19.0 mm;
L3: particles larger than 1.7 mm and smaller than 7.8 mm; L4: particles smaller than 1.7 mm; APS: average particles size; CP: crude protein; EE: ether extract; NDF:
neutral detergent fiber; ADF: acid detergent fiber; NFC: non-fiber carbohydrates; TDN: total digestible nutrients; DE: digestible energy; ME: metabolizable energy; DM:
dry matter.
b
Pearson correlation coefficient non-adjusted to IBW, and means and standard errors adjusted to the Model 2.

Furthermore, the chemical components of orts had no correlation with DM), ADF (308.6 g/kg DM), and lignin (66.5 g/kg DM) were higher; the
RIG and did not differ among efficiency groups (Table 4). Regarding the contents of NFC (99.4 g/kg DM), TDN (600.1 g/kg DM), DE (2.27 Mcal/
mean values for these components, the contents of NDF (649.1 g/kg kg DM), and ME (1.86 Mcal/kg DM) were lower; and the contents of CP

Table 5
Backfat thickness assessed by ultrasound and characteristics of longissimus muscle of crossbred ½ Dorper × ½ Santa Inês lambs in the three efficiency groups based on
residual intake and body weight gain.
P-value
Pearson correlation RIG groupa
Variablea Model 1 Model 2
a
rRIG P-value Inefficient Intermediate Efficient IBW RIG group RIG group

Initial BFTUS (mm)b 0.36 0.1467 1.07 ± 0.06 1.14 ± 0.06 1.18 ± 0.05 0.0304 0.3930 0.5823
Final BFTUS (mm)b 0.41 0.0923 1.42 ± 0.08 1.58 ± 0.08 1.58 ± 0.07 0.0023 0.2657 0.4492
Gain in BFTUS (mm)c 0.07 0.7601 0.36 ± 0.10 0.46 ± 0.10 0.38 ± 0.09 0.1509 0.8236 0.7502
LEW (cm)c − 0.07 0.7717 5.95 ± 0.18 5.70 ± 0.18 5.72 ± 0.18 0.1416 0.4974 0.5452
LED (cm)c − 0.09 0.7140 3.05 ± 0.12 3.35 ± 0.12 2.92 ± 0.12 0.0774 0.0766 0.0578
W:D ratioc 0.08 0.7611 1.96 ± 0.09 1.71 ± 0.09 1.98 ± 0.09 0.6039 0.1336 0.0992
BFTLE-MIN (mm)b 0.49 0.0459 1.11 ± 0.39 2.10 ± 0.39 2.06 ± 0.39 0.0164 0.1595 0.2447
BFTLE-MAX (mm)b 0.33 0.1894 2.31 ± 0.43 2.86 ± 0.43 3.00 ± 0.43 0.0047 0.5035 0.6555
BFTLE-MEAN (mm)b 0.42 0.0893 1.71 ± 0.40 2.48 ± 0.40 2.53 ± 0.40 0.0071 0.2912 0.4316
LEA (cm2)b − 0.02 0.9461 12.35 ± 0.44 12.31 ± 0.44 12.00 ± 0.44 0.0026 0.8291 0.6715
a
RIG: residual intake and body weight gain; IBW: initial body weight; BFTUS: backfat thickness assessed by ultrasound; LEW: maximum width of loin eye; LED:
maximum depth of loin eye; W:D: ratio between LEW and LED; BFTLE: backfat thickness measured on the loin eye; LEA: loin eye area.
b
Pearson correlation coefficient adjusted to IBW, and means and standard errors adjusted to the Model 1.
c
Pearson correlation coefficient non-adjusted to IBW, and means and standard errors adjusted to the Model 2.

5
E.M. do Nascimento et al.
(149.4 g/kg DM), EE (17.7 g/kg DM), and ash (84.4 g/kg DM) were close
to those of the ration provided to lambs (Table 1).
3.3. Longissimus muscle
For the initial and final BFTUS, the gain in BFTUS and the charac­
teristics of the longissimus muscle, only the minimum BFTLE had a pos­
itive and moderate correlation with RIG (r = 0.49), but it did not differ
among RIG groups; all other traits also did not differ among RIG groups
(Table 5). The mean values for initial, final, and gain in BFTUS were 1.13,
1.53, and 0.40 mm, respectively. The mean values for LEW, LED, W:D
ratio, minimum, maximum, and mean BFTLE, and LEA were 5.79 cm,
3.11 cm, 1.88, 1.76 mm, 2.72 mm, 2.24 mm, and 12.22 cm2,
respectively.
The final BFTUS was 0.71 mm lower than the mean BFTLE (1.53 vs.
2.24 mm), but there was a positive and moderate correlation between
these traits (r = 0.63; P = 0.0050). Thus, the same response was verified
for the final BFTUS and the mean BFTLE with respect to RIG, with no
differences among the efficiency groups (Table 5).
4. Discussion
4.1. Performance and feed efficiency
High phenotypic correlations of RIG with RFI and RG have also been
reported in other studies with growing cattle (Berry and Crowley, 2012;
Favero, 2014; Fernandes, 2014), finishing cattle (Takeda et al., 2018),
and lambs (Lima et al., 2017). Particularly in the latter study, the au­
thors evaluated crossbred ½ Dorper × ½ Santa Inês lambs and found
correlation coefficients for RIG with RFI and RG (rRFI = -0.92 and
rRG = 0.92) that are within the same magnitude as those verified in the
present study (rRFI = -0.90 and rRG = 0.90; Table 2). The RIG is an index
derived from RFI and RG, a fact that justifies the high phenotypic cor­
relation between these traits.
Despite the similarity of the correlation coefficients for RIG with RFI
and RG, Lima et al. (2017) reported higher values for these traits in
efficient and inefficient groups (2.0 vs. -2.4 for RIG; -80 vs. 90 g DM/day
for RFI; 20 vs. 30 g BW/day for RG) than those verified for the same
groups in the present study (Table 2). These results are related to the
higher DMI, ADG, and final BW (1350 g/day, 290 g/day, and 40.86 kg
on average, respectively) of the lambs evaluated by Lima et al. (2017),
which led to greater variation in these traits during the trial, as well as in
the RFI, RG and RIG values.
One of the main advantages of RFI and RG is their independence
from BW, which was also noted for RIG, because this trait had no
phenotypic correlation with final BW and MMW (Table 2). A similar
relationship has been verified in other studies with growing cattle (Berry
and Crowley, 2012; Fernandes, 2014; Grion et al., 2014) and lambs
(Lima et al., 2017).
The improvement in the efficiency for RIG leads to a reduced DMI
and increased ADG (Berry and Crowley, 2012; Nascimento et al., 2016);
however, both traits had no phenotypic correlation with RIG and were
similar among RIG groups (Table 2). These results may be related to the
low amplitude of RFI, RG, and RIG between the efficient and inefficient
groups, which corresponded to 105.9 g DM/day, 27.1 g BW/day, and
3.9, respectively. In lambs, Lima et al. (2017) and Carneiro et al. (2019)
also found no reduction in DMI, but they observed an increase in ADG
from inefficient to efficient group for RIG. Regarding the study of Lima
et al. (2017), the authors verified an amplitude of 170 g DM/day, 50 g
BW/day, and 4.4 for RFI, RG, and RIG, respectively. Associated with our
findings, these results suggest that the amplitude of RFI should be higher
than 170 g DM/day, whereas the amplitude of RG should be ≥ 50 g
BW/day in crossbred ½ Dorper × ½ Santa Inês lambs to cause a decrease
in DMI and an increase in ADG between the inefficient and efficient RIG
groups in statistical terms.
Lambs had the same initial and final BW, MMW, ADG, and DMI
6
Small Ruminant Research 192 (2020) 106248
among RIG groups, but DMIBW and DMIMMW were lower in the efficient
group (Table 2). Similar results for both traits have been reported in
other studies with growing cattle (Fernandes, 2014; Nascimento et al.,
2016) and lambs (Montelli et al., 2019); in the present study, this
discrepancy is likely explained by the numerical variations of DMI and
MMW among RIG groups. DMI decreased by 61.9 g/day (962.3 vs.
900.4 g/day) and MMW increased by 0.22 kg0.75 (12.21 vs. 12.43 kg0.75)
from the inefficient to the efficient group (Table 2), in which the latter
was derived from the mid-test BW. Thus, the DMIBW decreased from the
inefficient to the efficient group, and this response was reinforced by the
negative and moderate correlation of this trait with RIG (Table 2).
Besides affecting DMIBW and DMIMMW, the improved efficiency in
terms of RIG reduced the FCR (Table 2); this response has also been
reported in other studies with growing cattle (Berry and Crowley, 2012;
Fernandes, 2014) and lambs (Lima et al., 2017; Carneiro et al., 2019).
These authors found that the FCR decreased by 19.55–21.64 % and
23.08–23.28 % with an increase of RIG in lambs and growing cattle,
respectively. Thus, in the present study, the percentage variation for the
FCR between the efficient and inefficient groups (15.70 %) was lower
than those verified by Lima et al. (2017) and Carneiro et al. (2019).
Differences in the FCR among RIG groups are attributed to numerical
variations in DMI and ADG; the latter increased by 14.6 g/day from the
inefficient to efficient group (Table 2). Moreover, DMI and ADG were
moderately to highly correlated (P < 0.05) with FCR (rDMI = -0.51; rADG
= -0.89). This feed efficiency trait is derived from DMI and ADG, a fact
that justifies the phenotypic correlations with these performance traits.
When the results for FCR are associated with those for MMW and
DMIBW (Table 2), it is noted that the improved efficiency in terms of RIG
does not affect the body size, but it does reduce feed intake, resulting in
better utilization of the feed consumed by lambs. This pattern was also
verified in growing cattle by Fernandes (2014), who described that
better feed utilization is determined by the lower maintenance re­
quirements of growing bulls efficient for RIG. The same seems to occur
with lambs efficient for RIG, because they may present reduced organ
masses (rumen, liver, lung, and kidney; Zhang et al., 2017) and low
visceral fat deposition (Carneiro et al., 2019). These changes in body
composition lead to a reduced metabolic rate, which affects the energy
partition during the metabolization of nutrients and energy from diet. In
this case, a greater proportion of energy is directed to meet the net re­
quirements for gain, which is translated to better FCR associated or not
associated with reduction in feed intake (Redden et al., 2013; Zhang
et al., 2017; Carneiro et al., 2019).
The lack of correlation of RIG with RGR and KR and, hence, the
similar values of these traits between RIG groups (Table 2), are not
consistent with the literature. When ADG increases and MMW slightly
increases or remains unchanged with RIG, RGR and KR should increase
from the inefficient to the efficient group in terms of this trait (Berry and
Crowley, 2012; Fernandes, 2014; Lima et al., 2017). However, ADG had
no correlation with RIG and presented a slight, but not significant, dif­
ference among RIG groups, leading to the same RGR and KR for the
efficient and inefficient lambs.
4.2. Ingestive behavior
The similarities of the time spent feeding, in rumination, and in
idleness, as well as the IR among RIG groups (Table 3), have also been
reported by Favero (2014) for growing cattle. These feeding behavior
traits and the RR had no phenotypic correlation with RIG (Table 3);
however, Lima (2016) found a moderate correlation of the time spent
feeding, in rumination, and of IR with RIG (0.39, -0.34, and -0.42,
respectively). In this case, the time spent feeding increased, whereas the
time spent in rumination and the IR decreased with an increase in RIG,
which may be related to the higher manipulation of food and the small
bite size taken by efficient lambs. In the present study, there was no
evidence that corroborated these findings.
The lowest RR in the efficient group indicates that the amount of DM
E.M. do Nascimento et al.
ruminated per minute by efficient lambs was lower than those verified in
the intermediate and inefficient lambs, in terms of the same rumination
time among RIG groups (Table 3). The most digestible fractions of the
cell wall tend to be less degraded and digested when DMI increases (Van
Soest, 1994). Hence, a lower amount of short chain fatty acids is pro­
duced from fiber fermentation in the rumen, reducing the efficiency of
dietary energy utilization and leading lambs to increase the DMI to meet
their ME energy requirements (NRC, 2007; Ríos-Rincón et al., 2014).
Given that DMI decreased slightly (P = 0.2072) from the inefficient to
the efficient group (Table 2), and the ration provided to the lambs had a
high roughage:concentrate ratio (64:36; Table 1), the reduction of the
RR may be a physiological adaptation of the efficient lambs to increase
the digestibility of food and improve the dietary energy utilization. This
hypothesis is reinforced by the findings of Montelli et al. (2019), who
verified that DM digestibility increased and fecal output decreased with
an increase in RIG, in which both traits had a low to moderate pheno­
typic correlation with RIG (0.27 and -0.44, respectively).
The reduction of RR in lambs efficient for RIG may also be an indirect
effect of RFI, because its variation is influenced by 2% for feeding pat­
terns and by 10 % for food digestibility (Herd and Arthur, 2009). When
evaluating the feeding behavior and diet digestibility in Nellore heifers
with divergent efficiency for RFI, Magnani et al. (2013) found that the
digestibility of DM and NDF increased by 8.29 and 13.57 %, respec­
tively, from the inefficient to the efficient group in terms of RFI; those
authors also verified a moderate negative phenotypic correlation be­
tween RFI and the digestibility of NDF (r = -0.55). Furthermore, Zhang
et al. (2017) reported that the length of the duodenum and ileum in
low-RFI lambs was longer than that in high-RFI lambs; these findings
demonstrated that an increased capacity of nutrient absorption is a
consequence of improvement of feed efficiency based on RFI. Although
this efficiency trait is strongly correlated with RIG (Table 2), further
studies should be carried out to understand the relationship of ingestive
behavior and nutrients digestibility with RIG in growing lambs.
The improvement of RIG efficiency did not affect the proportion of
particles in each layer of the PSPS and the APS of the orts (Table 4); these
data demonstrated that lambs classified as efficient, intermediate, and
inefficient in terms of RIG selected similar diets. No studies have re­
ported a pattern of diet selection in lambs and cattle with divergent
efficiencies for RIG. However, using the PSPS, Fernandes (2014) eval­
uated the proportion of different particle sizes and the APS of orts from
growing cattle classified as efficient, intermediate, and inefficient for
RFI. This author noted a higher proportion of particles larger than
19.0 mm (17.33 vs. 12.14 %), a lower proportion of particles ranging
from 1.7 and 7.8 mm (29.64 vs. 36.60 %), and a higher APS (9.35 vs.
7.89 mm) in the orts of efficient animals, compared with those of inef­
ficient animals; there was also a low to moderate phenotypic correlation
of these traits with RFI (-0.27, 0.33 and -0.30, respectively). These
findings indicate that efficient animals selected diets with a higher
proportion of concentrate than inefficient animals, in terms of RFI.
Given that RIG is derived from RFI, and that sheep are more selective
than cattle during feeding, a similar response had been expected in
lambs efficient for RIG, but this was not confirmed in the present study.
Following the aforementioned results, the chemical composition of
the orts had no relationship with RIG (Table 4). However, the differences
between the chemical composition of ration and orts suggest that lambs
selected different ingredients from the ration in a similar way among the
efficiency groups. Due to the higher proportion of fibrous components in
the orts compared with the ration (Table 1), the lambs had a preference
for ingredients contained in the concentrate feed, refusing a greater
amount of hay during feeding. This result is a consequence of the
increased demand for energy and nutrients, mainly protein, until body
maturity is reached (NRC, 2007). Indeed, lambs can select different
foods in order to meet their nutritional requirements for growth, espe­
cially increasing the intake of feedstuffs rich in protein (Forbes, 2007;
Dikmen et al., 2009). This was observed in the present study, because
the concentrate feed had a CP content that was almost three times higher
7
Small Ruminant Research 192 (2020) 106248
(27.08 vs. 9.56 % DM), with an ME content that was 40 % higher (2.58
vs. 1.86 Mcal/kg DM) than Cynodon hay.
4.3. Longissimus muscle
Given that the measurements obtained in vivo by ultrasound or
directly from the longissimus muscle have a high correlation with carcass
composition (Garcia et al., 2003; Silva, 2017), the lack of a correlation of
these traits (except for minimum BFTLE; Table 5) with RIG suggests that
the lambs’ carcass composition is not influenced by this efficiency trait.
Similar results have been reported for growing cattle (Favero, 2014;
Fernandes, 2014) and lambs (Lima et al., 2017), which showed no
phenotypic correlation of RIG with BFTUS and LEA measured by ultra­
sound. There is a positive and high correlation between LEA measured
by ultrasound and LEA measured directly on the longissimus muscle
(Cartaxo et al., 2011; Cacere et al., 2014; Silva, 2017), findings that
justify the lack of phenotypic correlation between the latter with RIG in
the present study.
The underestimation of BFTUS measured at the end of the trial in
relation to the mean BFTLE (Table 5) has also been verified in other
studies with lambs (Rozanski et al., 2017; Nascimento et al., 2018) and
hoggets (Cacere et al., 2014). Even though a trained person performed
the ultrasonographic evaluation, some factors, such as a slight change in
the transducer position over the site of measurement, the pressure
applied on transducer over the skin, a smaller subcutaneous fat thickness
and difficulty in distinguishing the skin:fat interface in sheep compared
to other species (cattle and pigs), may have led to an underestimation of
BFTUS (Silva, 2017). Due to the positive and moderate correlation
verified between the final BFTUS and the mean BFTLE (r = 0.63), both
traits had the same response to RIG.
The morphology of the loin eye (characterized by LEW, LED, and the
W:D ratio) was not influenced by RIG. This outcome is explained by the
similarity of LEA among the efficiency groups (Table 5). Furthermore,
despite the positive and moderate correlation between RIG and the
minimum BFTLE (r = 0.49), this trait, as well as the maximum and mean
BFTLE, was similar among efficiency groups, indicating that the thick­
ness of fat covering the carcass was not influenced by RIG. The lack of
effect of this efficiency trait on the characteristics of the longissimus
muscle may be related to the similar BW at slaughter among the effi­
ciency groups (Table 2), which seems to have a stronger effect than RIG
on the carcass traits of growing lambs (Carneiro et al., 2019).
According to Herd and Arthur (2009), differences in body composi­
tion account for 5% of the variation in RFI, in which low-RFI animals
have greater muscle development and lower fat deposition than
high-RFI animals during the growing phase. RIG is derived from RFI;
hence, it had been expected that changes in RIG efficiency could be
related to differences in the body composition of lambs. However, in
terms of the results obtained for LEA and BFT in the present study and
those obtained by Lima et al. (2017), who evaluated crossbred ½ Dorper
× ½ Santa Inês lambs heavier than those in this study, an improvement
of RIG efficiency seems to not influence the muscle development and fat
deposition in the carcass of these animals during the growth and fin­
ishing phases.
5. Conclusions
An improvement in the efficiency for RIG in ½ Dorper × ½ Santa Inês
lambs leads to better feed utilization, with reduced feed intake relative
to body weight and feed conversion ratio but no effects on the charac­
teristics of the longissimus muscle. Furthermore, the better feed effi­
ciency seems to be related to the reduction of rumination rate in efficient
lambs, although our results are not fully conclusive.
Acknowledgments
The authors wish to acknowledge the financial support, granted as a
E.M. do Nascimento et al. Small Ruminant Research 192 (2020) 106248

scholarship, provided by the Coordination for the Improvement of Gordon, H.M., Whitlock, H.V., 1939. A new technique for counting nematode eggs in
sheep faeces. J. Counc. Sci. Ind. Res. 12, 50–52.
Higher Education Personnel (Coordenação de Aperfeiçoamento de Pessoal
Grion, A.L., Mercadante, M.E.Z., Cyrillo, J.N.S.G., Bonilha, S.F.M., Magnani, E.,
de Nível Superior – CAPES) of Brazil, Finance Code 001; Impextraco Latin Branco, R.H., 2014. Selection for feed efficiency traits and correlated genetic
America Ltd. for the donation of Rumimpex® and Elitox®, both used in responses in feed intake and weight gain of Nellore cattle. J. Anim. Sci. 92, 955–965.
the experimental ration; and the students of the Group of Study on https://doi.org/10.2527/jas.2013-6682.
Hall, M.B., Hoover, W.H., Jennings, J.P., Webster, T.K.M., 1999. A method for
Ruminant Nutrition and Forage Crops (Grupo de Estudos em Nutrição de partitioning neutral detergent-soluble carbohydrates. J. Sci. Food Agric. 79,
Ruminantes e Forragicultura – GENFOR) of UFPR, Palotina Campus, for 2079–2086. https://doi.org/10.1002/(SICI)1097-0010(199912)79:15<2079::AID-
their support in the experimental preparation and data collection during JSFA502>3.0.CO;2-Z.
Heinrichs, J., Kononoff, P., 2002. Evaluating particle size of forages and TMRs using the
the trial. new Penn State forage particle separator. Dairy & Animal Science (DAS) 02–42.
Pennsylvania State University – College of Agricultural Sciences, Pennsylvania, USA.
References Herd, R.M., Arthur, P.F., 2009. Physiological basis for residual feed intake. J. Anim. Sci.
87, E64–E71. https://doi.org/10.2527/jas.2008-1345.
Kleiber, M., 1947. Body size and metabolic rate. Physiol. Rev. 27, 511–541. https://doi.
AOAC, 1990. Official Methods of Analysis, 15th ed. Association of Official Analytical
org/10.1152/physrev.1947.27.4.511.
Chemists, Arlington, VA.
Koch, R.M., Swiger, L.A., Chambers, D., Gregory, K.E., 1963. Efficiency of feed use in
Arthur, P.F., Renand, G., Krauss, D., 2001. Genetic and phenotypic relationships among
beef cattle. J. Anim. Sci. 22, 486–494. https://doi.org/10.2527/jas1963.222486x.
different measures of growth and feed efficiency in young Charolais bulls. Livest.
Lima, N.L.L., 2016. Productive Efficiency in Lambs Classified by Residual Feed Intake
Prod. Sci. 68, 131–139. https://doi.org/10.1016/S0301-6226(00)00243-8.
(RFI) and Residual Intake and Gain (RIG). Thesis (Doctor’s Degree in Animal
Berry, D.P., Crowley, J.J., 2012. Residual intake and body weight gain: a new measure of
Science). Federal University of Minas Gerais, Belo Horizonte, Minas Gerais, Brazil,
efficiency in growing cattle. J. Anim. Sci. 90, 109–115. https://doi.org/10.2527/
p. 144. https://sucupira.capes.gov.br/sucupira/public/consultas/coleta/trabalh
jas.2011-4245.
oConclusao/viewTrabalhoConclusao.jsf?popup=true&id_trabalho=3622145#.
Cacere, R.A.S., Morais, M.G., Alves, F.V., Feijó, G.L.D., Ítavo, C.C.B.F., Ítavo, L.C.V.,
Lima, N.L.L., Ribeiro, C.R.F., Sá, H.C.M., Leopoldino Jr, I., Cavalcanti, L.F.L., Santana, R.
Oliveira, L.B., Ribeiro, C.B., 2014. Quantitative and qualitative carcass
A.V., Furusho-Garcia, I.F., Pereira, I.G., 2017. Economic analysis, performance, and
characteristics of feedlot ewes subjected to increasing levels of concentrate in the
feed efficiency in feedlot lambs. R. Bras. Zootec. 46, 821–829. https://doi.org/
diet. Arq. Bras. Med. Vet. Zootec. 66, 1601–1610. https://doi.org/10.1590/1678-
10.1590/s1806-92902017001000005.
6376.
Liu, M.F., Goonewardene, L.A., Bailey, D.R.C., Basarab, J.A., Kemp, R.A., Arthur, P.F.,
Cardoso, A.R., Carvalho, S., Galvani, D.B., Pires, C.C., Gasperin, B.G., Garcia, R.P.A.,
Okine, E.K., Makarechian, M., 2000. A study on the variation of feed efficiency in
2006. Ingestive behavior of lambs fed with a diet of different levels of fiber in neutral
station tested bulls. Can. J. Anim. Sci. 80, 435–441. https://doi.org/10.4141/A99-
detergent. Cienc. Rural 36, 604–609. https://doi.org/10.1590/S0103-
030.
84782006000200038.
Magnani, E., Nascimento, C.F., Branco, R.H., Bonilha, S.F.M., Ribeiro, E.G.,
Carneiro, M.M.Y., Morais, M.G., Souza, A.R.D.L., Fernandes, H.J., Feijó, G.L.D.,
Mercadante, M.E.Z., 2013. Relationship among residual feed intake, digestibility and
Bonin, M.N., Franco, G.L., Rocha, R.F.A.T., 2019. Residual intake and gain for the
ingestive behavior in Nellore heifers. B. Indústr. Anim. 70, 187–194. https://doi.
evaluation of performance, non-carcass components, and carcass characteristics of
org/10.17523/bia.v70n2p187.
confined crossbred Texel lambs. R. Bras. Zootec. 48, e20180206 https://doi.org/
Martin, P., Bateson, P., 1993. Measuring Behavior, 2nd ed. Cambridge University Press,
10.1590/rbz4820180206.
Cambridge.
Cartaxo, F.Q., Sousa, W.H., Cezar, M.F., Costa, R.G., Cunha, M.G.G., Gonzaga Neto, S.,
Molento, M.B., Tasca, C., Gallo, A., Ferreira, M., Bononi, R., Stecca, E., 2004. Famacha
2011. Carcass traits determined by ultrasonography in real time and after slaughter
guide as an individual clinic parameter for Haemonchus contortus infection in small
of lambs finished in feedlot with different levels of energy in the diet. R. Bras.
ruminants. Cienc. Rural 34, 1139–1145. https://doi.org/10.1590/S0103-
Zootec. 40, 160–167. https://doi.org/10.1590/S1516-35982011000100023.
84782004000400027.
Castilhos, A.M., Branco, R.H., Razook, A.G., Bonilha, S.F.M., Mercadante, M.E.Z.,
Montelli, N.L.L.L., Almeida, A.K., Ribeiro, C.R.F., Grobe, M.D., Abrantes, M.A.F.,
Figueiredo, L.A., 2011. Test post-weaning duration for performance, feed intake and
Lemos, G.S., Garcia, I.F.F., Pereira, I.G., 2019. Performance, feeding behavior and
feed efficiency in Nellore cattle. R. Bras. Zootec. 40, 301–307. https://doi.org/
digestibility of nutrients in lambs with divergent efficiency traits. Small Rumin. Res.
10.1590/S1516-35982011000200010.
180, 50–56. https://doi.org/10.1016/j.smallrumres.2019.07.016.
Cezar, M.F., Sousa, W.H., 2007. Carcaças ovinas e caprinas: obtenção, avaliação e
Nascimento, M.L., Souza, A.R.D.L., Chaves, A.S., Cesar, A.S.M., Tullio, R.R., Medeiros, S.
classificação. Agropecuária Tropical, Uberaba.
R., Mourão, G.B., Rosa, A.N., Feijó, G.L.D., Alencar, M.M., Lanna, D.P.D., 2016. Feed
Detmann, E., Valadares Filho, S.C., Pina, D.S., Campos, J.M.S., Paulino, M.F., Oliveira, A.
efficiency indexes and their relationships with carcass, non-carcass and meat quality
S., Silva, P.A., 2006a. Estimation of ether extract digestibility in diets of ruminants: a
traits in Nellore steers. Meat Sci. 116, 78–85. https://doi.org/10.1016/j.
model under Brazilian conditions. R. Bras. Zootec. 35, 1469–1478. https://doi.org/
meatsci.2016.01.012.
10.1590/S1516-35982006000500029.
Nascimento, U.F.S., Santos, G.R.A., Azevedo, C.S., Macedo, F.A.F., Gonçalves, T.R.,
Detmann, E., Valadares Filho, S.C., Henriques, L.T., Pina, D.S., Paulino, M.F.,
Bomfim, L.E.L.M., Farias, J.S., Santos, A.D.F., 2018. Performance and carcass
Valadares, R.F.D., Chizzotti, M.L., Magalhães, K.A., 2006b. Estimation of nonfiber
characteristics of lambs ½ Dorper + ½ Santa Inês, slaughtered with different
carbohydrates digestibility in cattle using the Lucas test approach under Brazilian
thicknesses of subcutaneous fat. Rev. Bras. Saúde Prod. Anim. 19, 125–135. https://
conditions. R. Bras. Zootec. 35, 1479–1486. https://doi.org/10.1590/S1516-
doi.org/10.1590/s1519-99402018000100012.
35982006000500030.
NRC, 2007. Nutrient Requirements of Small Ruminants: Sheep, Goats, Cervids and New
Detmann, E., Pina, D.S., Valadares Filho, S.C., Campos, J.M.S., Paulino, M.F., Oliveira, A.
World Camelids. National Academies Press, Washington, DC.
S., Silva, P.A., Henriques, L.T., 2006c. Estimation of crude protein digestible fraction
Paula, E.F.E., Monteiro, A.L.G., Souza, D.F., Prado, O.R., Nomura, T.M., Stivari, T.S.S.,
in cattle diets under Brazilian conditions. R. Bras. Zootec. 35, 2101–2109. https://
Silva, C.J.A., Santana, M.H.A., 2013. The residual feed intake and its relationship
doi.org/10.1590/S1516-35982006000700030.
with performance and efficiency measures and in vivo carcass characteristics of
Detmann, E., Valadares Filho, S.C., Henriques, L.T., Pina, D.S., Paulino, M.F.,
lambs. Arq. Bras. Med. Vet. Zootec. 65, 566–572. https://doi.org/10.1590/S0102-
Magalhães, A.L.R., Figueiredo, D.M., Porto, M.O., Chizzotti, M.L., 2007.
09352013000200037.
Reparameterization of the model based on surface law to predict the digestible
Redden, R.R., Surber, L.M.M., Grove, A.V., Kott, R.W., 2013. Growth efficiency of ewe
fraction of neutral detergent fiber in Brazilian cattle. R. Bras. Zootec. 36, 155–164.
lambs classified into residual feed intake groups and pen fed a restricted amount of
https://doi.org/10.1590/S1516-35982007000100019.
feed. Small Rumin. Res. 114, 214–219. https://doi.org/10.1016/j.
Dikmen, S., Ustuner, H., Turkmen, I.I., Organ, M., 2009. Fattening performance and feed
smallrumres.2013.07.002.
source preference of native Awassi lambs fed individually in a cafeteria feeding
Ríos-Rincón, F.G., Estrada-Angulo, A., Plascencia, A., López-Soto, M.A., Castro-Pérez, B.
system. Trop. Anim. Health Prod. 41, 485–491. https://doi.org/10.1007/s11250-
I., Portillo-Loera, J.J., Robles-Estrada, J.C., Calderón-Cortes, J.F., Dávila-Ramos, H.,
008-9212-8.
2014. Influence of protein and energy level in finishing diets for feedlot hair lambs:
Favero, R., 2014. Feed Efficiency in Brahman Cattle and Their Relationships With
growth performance, dietary energetics and carcass characteristics. Asian Australas.
Economically Relevant Traits and Behavioral Variables. Dissertation (Master in
J. Anim. Sci. 27, 55–61. https://doi.org/10.5713/ajas.2013.13307.
Animal Science). State University of Londrina, Londrina, Paraná, Brazil, p. 117. htt
Rozanski, S., Vivian, D.R., Kowalski, L.H., Prado, O.R., Fernandes, S.R., Souza, J.C.,
p://www.bibliotecadigital.uel.br/document/?code=vtls000191199&print=y.
Freitas, J.A., 2017. Carcass and meat traits, and non-carcass components of lambs
Fernandes, S.R., 2014. Feed Efficiency and Their Relationships with Blood Biochemical
fed ration containing increasing levels of urea. Semina: Ciênc. Agrár. 38, 1587–1604.
Profile, Pattern of Diet Selection and Carcass Traits of Young Purunã Bulls. Thesis
https://doi.org/10.5433/1679-0359.2017v38n3p1587.
(Doctor’s Degree in Veterinary Science). Federal University of Paraná, Curitiba,
SAS Institute, 2002. Statistical Analysis System (SAS) User’s Guide for Windows:
Paraná, Brazil, p. 119. http://hdl.handle.net/1884/37000.
Statistics, Version 9.0. SAS Institute Inc., Cary, NC, USA.
Fitzhugh Junior, H.A., Taylor, St. C.S., 1971. Genetic analysis of degree of maturity.
Silva, S.R., 2017. Use of ultrasonographic examination for in vivo evaluation of body
J. Anim. Sci. 33, 717–725. https://doi.org/10.2527/jas1971.334717x.
composition and for prediction of carcass quality of sheep. Small Rumin. Res. 152,
Forbes, J.M., 2007. Voluntary Food Intake and Diet Selection in Farm Animals, 2nd ed.
144–157. https://doi.org/10.1016/j.smallrumres.2016.12.020.
CAB International, Wallingford.
Takeda, M., Uemoto, Y., Inoue, K., Ogino, A., Nozaki, T., Kurogi, K., Yasumori, T.,
Garcia, C.A., Monteiro, A.L.G., Costa, C., Neres, M.A., Rosa, G.J.M., 2003. Objective
Satoh, M., 2018. Evaluation of feed efficiency traits for genetic improvement in
measurements and tissue composition of carcass of lambs fed with different energy
Japanese Black cattle. J. Anim. Sci. 96, 797–805. https://doi.org/10.1093/jas/
levels in creep feeding. R. Bras. Zootec. 32, 1380–1390. https://doi.org/10.1590/
skx054.
S1516-35982003000600013.

8
E.M. do Nascimento et al.
Vale, F.X.R., Fernandes Filho, E.I., Liberato, J.R., 2003. QUANT: a software plant disease
severity assessment. In: 8th International Congress of Plant Pathology. Proceedings…
Christchurch, New Zealand,.
Van Soest, P.J., 1994. Nutritional Ecology of the Ruminant, 2nd ed. Cornell University
Press, New York.
Van Soest, P.J., Robertson, J.B., Lewis, B.A., 1991. Methods for dietary fiber, neutral
detergent fiber, and no starch polysaccharides in relation to animal nutrition.
J. Dairy Sci. 74, 3583–3597. https://doi.org/10.3168/jds.S0022-0302(91)78551-2.
Small Ruminant Research 192 (2020) 106248
Weiss, W.P., Conrad, H.R., Pierre, St. N.R., 1992. A theoretically-based model for
predicting total digestible nutrient values of forages and concentrates. Anim. Feed
Sci. Technol. 39, 95–110. https://doi.org/10.1016/0377-8401(92)90034-4.
Zhang, X., Wang, W., Mo, F., La, Y., Li, C., Li, F., 2017. Association of residual feed intake
with growth and slaughtering performance, blood metabolism, and body
composition in growing lambs. Sci. Rep. 7, 12681 https://doi.org/10.1038/s41598-
017-13042-7.