de Araújo et al.

Journal of Animal Science and Biotechnology (2017) 8:46

DOI 10.1186/s40104-017-0175-3

RESEARCH Open Access

Effects of quantitative feed restriction and

sex on carcass traits, meat quality and meat
lipid profile of Morada Nova lambs
Thiago L. A. C. de Araújo1, Elzânia S. Pereira1, Ivone Y. Mizubuti2, Ana C. N. Campos1, Marília W. F. Pereira1,
Eduardo L. Heinzen1, Hilton C. R. Magalhães3, Leilson R. Bezerra4, Luciano P da Silva1 and Ronaldo L. Oliveira5*

Abstract
Background: An experiment was conducted to evaluate the effects of feed restriction (FR) and sex on the
quantitative and qualitative carcass traits of Morada Nova lambs. Thirty-five animals with an initial body weight
of 14.5 ± 0.89 kg and age of 120 d were used in a completely randomized study with a 3 × 3 factorial scheme
consisting of three sexes (11 entire males, 12 castrated males and 12 females) and three levels of feeding (ad
libitum – AL and 30% and 60% FR).
Results: Entire males presented greater hot and cold carcass weights (P < 0.05), followed by castrated males and
females. However, the hot carcass yield was higher for females and castrated males than for entire males. Luminosity
values were influenced (P < 0.05) by sex, with entire males presenting higher values than castrated males and females.
Females showed higher (P < 0.05) concentrations of linoleic acid and arachidonic acid in the meat of the longissimus
thoracis muscle. The meat of animals submitted to AL intake and 30% FR showed similar (P > 0.05) concentrations, and
the concentrations of palmitic acid, palmitoleic acid, stearic acid, oleic acid and conjugated linoleic acid were higher
(P < 0.05) than those of animals with 60% FR. The meat of females had a higher ω6/ω3 ratio and lower h/H ratio, and
females had greater levels of feeding. The meat of animals on the 60% FR diet had a greater ω6/ω3 ratio, lower h/H
ratio and lower concentration of desirable fatty acids in addition to a greater atherogenicity index (AI) and
thrombogenicity index (TI).
Conclusion: Lambs of different sexes had carcasses with different quantitative traits without total influence on the
chemical and physical meat characteristics. The lipid profile of the meat was less favorable to consumer health when
the animals were female or submitted to 60% feed restriction.
Keywords: Dietary restriction, Fatty acid, Hair sheep, Lean meat, Semi-arid condition

Background Production systems in semi-arid regions are based on

Farming ruminants has an unquestionable importance the use of genetic resources with high adaptability and
to the economic and food security of many regions of heat tolerance, which are heavily influenced by the quali-
the world, especially for tropical semi-arid regions [1], tative and quantitative seasonality of food, but these
where sheep are highly relevant [2, 3]. In these regions, local breeds are now threatened with extinction. Morada
the environment interferes intensely with productive Nova is a prominent genetic group of hair sheep in
management strategies, and making decisions is crucial semiarid regions of Brazil. These sheep are an indigen-
for the success of animal husbandry. ous group used for the production of meat and skin.
They are smaller, have lower mortality rates, produce
lighter carcasses, and are usually late to slaughter. Adult
* Correspondence: [email protected] Morada Nova hair sheep weigh between 45 and 50 kg
5
Department of Animal Science, School of Veterinary Medicine and Animal
Science/Federal University of Bahia, Salvador City, Bahia State 40.170-110,
and can reproduce at approximately 8–9 mons of age
Brazil with approximately 28 kg of body weight (BW) [4].
Full list of author information is available at the end of the article

© The Author(s). 2017 Open Access This article is distributed under the terms of the Creative Commons Attribution 4.0
International License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted use, distribution, and
reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to
the Creative Commons license, and indicate if changes were made. The Creative Commons Public Domain Dedication waiver
(http://creativecommons.org/publicdomain/zero/1.0/) applies to the data made available in this article, unless otherwise stated.
de Araújo et al. Journal of Animal Science and Biotechnology (2017) 8:46 Page 2 of 12

The improvement of animal yield by enhancing supply the nutritional requirements of late maturity
sustainable biodiversity may be a pathway toward greater lambs with a gain of 150 g/d as recommended by the
food supplies. Such sustainable increases may be espe- National Research Council (NRC) [12]. Before beginning
cially important for the 2 billion people reliant on small data collection, the animals were randomly assigned to
farms, many of which are undernourished, yet we know individual boxes provided with feed and water troughs,
little about the efficacy of this approach. where they underwent an adaptive period of 15 d. Total
Some productive strategies affect animal performance mixed rations were provided twice a day (0730 and
as well as the chemical and physical quality of the meat 1600 h), allowing for up to 10% orts only for animals fed
produced. The farmer can, for example, opt to obtain AL. Before each morning feeding, the orts of each
production rates and carcasses with different characteris- animal fed AL were removed and weighed to calculate
tics depending on the sex of the animals [5–7]. In tropical the intake and feeding level of the lambs submitted to
semi-arid regions, animals can be submitted naturally to 30% and 60% FR (300 and 600 g/kg of FR). Thus, the
periods of feed restriction (FR) due to feed supply varia- restrictions were proportionally based on the intake of
tions or due to feed management planning, which is animals fed AL of each sex.
commonly used to save resources and reduce costs [8, 9]. The ingredients used in the total ration and their pro-
Under these conditions, it may be assumed that in portions and composition are described in Table 1.
addition to lower performance, meat products may have Samples of the roughage, concentrated and feed orts
different chemical and physical characteristics if the were taken to determine their chemical compositions
animals are sold [10, 11]. and dry matter intake (DMI) of the lambs. The lambs
In the face of global concerns about the safety and were weighed every fifteen days to calculate BW gain
nutritional quality of foods, it is necessary to understand (BWG). The trial period lasted 120 d. At the end of the
the effects of commonly used production strategies not trial period, the animals were weighed to determine total
only on productivity but also on aspects related to human weight gain (TWG) and average daily gain (ADG).
health, as is the case with lipid quality parameters. Thus,
we simulate the impact of feed restriction in hair sheep of Slaughter, carcass data and meat samples
different sexes in semi-arid regions. We then evaluate After 18 h of fasting, the animals were weighed to deter-
their effects on carcass characteristics, meat quality and mine their BW at slaughter (BWS). The animals were
fatty acid profiles in the meat of Morada Nova lambs. then skinned and eviscerated according to the rules
established in the Regulation of Brazilian Industrial and
Methods Sanitary Inspection of Animal Products. Subsequently,
Animal care and location the lambs were stunned with the proper equipment,
This study was conducted at the Department of Animal bled, skinned, and eviscerated. The viscera were weighed
Science, Federal University of Ceará, located in Forta- when filled, emptied, washed, drained and weighed when
leza, CE, Brazil. Protocols (n° 98/2015) were in accord- empty to determine the contents of the gastrointestinal
ance with the standards established by the Committee of tract and subsequently the empty BW (EBW) of the
Ethics in Animal Research of the Federal University of animals. The carcasses were identified and weighed to
Ceará. obtain the hot carcass weight (HCW) and yield (HCY)
calculated in relation to BWS. After 24 h of cooling at
Animals, experimental design and management 4 °C, the carcasses were weighed to obtain the cold
Experimental lambs were obtained from the Morada carcass weight (CCW). Twenty-four hours post mortem,
Nova sheep breeding facility. The mating season was the pH was measured using a pH meter (HI-99163,
established with the objective of enabling a selection of Hanna® instruments, São Paulo, Brazil) by inserting the
animals with little variation in BW. Thirty-five lambs of meter between the 4th and 5th lumbar vertebrae in the
the Morada Nova breed, including 23 males and 12 longissimus lumborum muscle.
females, were selected. Twelve entire males were The carcasses were sectioned with an electric saw (Ki
randomly assigned to the sexual class of castrated males, Junta®, São Paulo, Brazil) along the spine, and the left
and the males were castrated using the burdizzo castrat- halves of the carcasses were divided into six commercial
ing method. Initially, the lambs had 14.5 ± 0.89 kg of cuts (leg, loin, ribs, lower ribs, neck and shoulder),
BW and 120 d of age. The lambs were distributed in a which were individually weighed. A cross-sectional cut
completely randomized design in a 3 × 3 factorial was made between the 12th and 13th ribs to expose the
scheme. Experimental treatments consisted of three longissimus thoracis (LT) muscle, which measured the
sexes (11 entire males, 12 castrated males and 12 maximum distances between the ends of the muscle in
females) and three quantitative feeding levels (ad libitum the mediolateral direction (A) and dorsal-ventral (B) to
(AL), 30% and 60% FR). The ration was formulated to subsequently calculate the rib eye area (REA) according
de Araújo et al. Journal of Animal Science and Biotechnology (2017) 8:46 Page 3 of 12

Table 1 Ingredient proportion and chemical composition of experimental rations

Ingredient Proportion, % of DM
Tifton 85 grass hay 60.0
Ground corn grain 32.72
Soybean meal 6.30
Dicalcium phosphate 0.06
a
Mineral premix 0.92
Chemical composition, g/kg of DM Total ration Tifton 85 grass hay Ground corn grain Soybean meal
Dry matter 907.72 913.40 892.40 910.00
Crude protein 169.32 172.50 102.80 508.80
Ether extract 30.77 25.57 43.18 19.32
Ash 61.93 73.40 13.30 65.90
Neutral detergent fiber 438.65 668.20 112.54 134.63
NDFap b 418.32 644.85 97.89 110.41
Acid detergent fiber 201.93 317.54 26.31 102.05
Non-fiber carbohydrate 319.67 58.30 728.19 273.30
a
Composition, 1 kg of premix: Calcium 225 g to 215 g; Phosphor 40 g; Sulfur 15 g; Sodium 50 g; Magnesium 10 g; Cobalt 11 mg; Iodine 34 mg; Manganese
1,800 mg; Selenium 10 mg; Zinc 2,000 mg; Iron 1,250 mg; Copper 120 mg; Fluor 400 mg; Vitamin A 37.5 mg; Vitamin D3 0.5 mg and Vitamin E 800 mg
b
Neutral detergent fiber corrected for ash and protein

to Eq. REA = (A / 2 × B / 2) × π. Subcutaneous fat thickness 170 °C. A digital skewer thermometer (Salcasterm 200®,
(SFT) was verified above measure B using a digital caliper. São Paulo, Brazil) was used to monitor the internal
Samples were taken from the LT and longissimus lumborum temperature of the steak until the center reached 71 °C.
(LL) muscles, vacuum packed and stored at −20 °C. Then, each steak was brought to room temperature,
removed from the oven after temperature stabilization,
Physicochemical meat analyses and weighed again. The difference between the initial and
The meat color was evaluated using a transverse cut on final weights of a sample was used to determine the CWL,
the back section, which was exposed to atmospheric air with the value expressed as a percentage.
for 30 min before reading the oxygen myoglobin, which After cooling at room temperature, the samples were
is the primary element that defines meat color [13]. As again wrapped in foil and placed in a refrigerator (Consul
described by Miltenburg et al. [14], the coordinates L*, a* CHB53C®, Salvador, Brazil) for 12 h at 4 °C. Fillets (5 ± 1)
and b* were measured at three different points on the approximately 2 cm long, 1 cm wide and 1 cm high were
muscle, and the triplicates were averaged for each coord- cut from the meat to be evaluated for Warner-Bratzler
inate per animal. These measurements were performed shear force (WBSF). The instrumental texture analysis
using a Minolta CR-10 colorimeter (Konica® Minolta, was performed on a TAXT2 texturometer (Stable Micro
Osaka, Japan) that was previously calibrated with the Systems Ltd., Vienna Court, UK) at 200 mm/min using
CIELAB system using a blank tile, illuminant D65 and 10° standard shear blades (1.016 mm thick with a 3.05-mm
as the standard observation points. L* is related to light- blade). The instrumental texture analysis was performed
ness (L* = 0 black, 100 white); a* (redness) ranges from according to the Research Center for Meat (US Meat
green (−) to red (+); and b*(yellowness) ranges from blue Animal Research Center) and Shackelford et al. [17].
(−) to yellow (+). Measurements were made from a 2° To evaluate lipid oxidation, meat samples of LL
viewing angle using illuminant C. The color saturation muscle stored under fast freezing at −20 °C for three
(chroma, C*) was calculated as (a*2 + b*2)1/2 [15]. months were thawed and crushed. Using the aqueous
Meat samples of LL muscle were processed in a acid extraction method described by Cherian et al. [18], the
crusher to determine the water holding capacity (WHC), 2-thiobarbituric acid reactive substances (TBARS) were
and cooking weight loss (CWL) was determined according measured in mg of malondialdehyde (MDA)/g of tissue.
to the American Meat Science Association (AMSA) Meat samples of LT muscle were evaluated for mois-
[16] using LL meat samples (triplicate) without visible ture, ash and protein contents, following method num-
connective tissue that were previously thawed at 10 °C for bers 930.15, 920.153 and 928.08, respectively [19]. LT
12 h. CWL indicated the difference in the weight of the muscle samples were used to extract and quantify intra-
meat before and after cooking on a preheated grill (George muscular fat (IMF). The fat of meat samples was isolated
Foreman Jumbo Grill GBZ6BW, Rio de Janeiro, Brazil) at and purified using polar solvents (chloroform and
de Araújo et al. Journal of Animal Science and Biotechnology (2017) 8:46 Page 4 of 12

methanol) according to the procedure of Folch et al. 967.03), ash (method 942.05), crude protein (CP
[20]. Aliquots of the fat extract were reserved and stored method 981.10), and ether extract (EE method 920.29)
at −20 °C for subsequent use in determining the fatty according to the Association of Official Analytical
acid profile. Chemists (AOAC) [27].
The neutral detergent fiber (NDF) content was deter-
Fatty acid profile mined as described by Van Soest et al. [28]. The acid
To determine the fatty acid profile, the fat samples detergent fiber (ADF) contents were determined as
previously extracted from LT muscle were converted to described by Robertson and Van Soest [29]. The NDF
fatty acid methyl esters (FAMEs). The FAMEs were residue was incinerated in an oven at 600 °C for 4 h to
prepared using a solution of methanol, ammonium determine the ash content, and the protein concentra-
chloride and sulfuric acid, following the procedure tion was calculated by subtracting the neutral detergent
described by Hartman and Lago [21]. insoluble protein (NDIP). NDF was corrected for the ash
Samples were analyzed using a chromatograph and protein contents. The Non-fiber carbohydrate
(GC2010, Shimadzu®, São Paulo, Brazil) equipped with a (NFC) content was measured according to Mertens [30]
flame-ionization detector and a biscyanopropyl polydi- and calculated based on the differences in the equation
methylsiloxane capillary column of stationary phase NFC = 100 – NDF – CP – EE – ash.
(SP2560, 100 m × 0.25 mm, df 0.20 μm; Supelco®,
Bellefonte, PA, USA). The column oven temperature Statistical analyses
was as follows: the initial temperature was held for 80 ° Variables were subjected to analysis of variance using
C, increased at 11 °C/min to 180 °C and at 5 °C/min to the GLM procedure of Statistical Analysis System - SAS®
220 °C and then maintained for 19 min. Hydrogen was software [31] and the following equation: Yijk = μ + Si +
used as a carrier gas at a flow rate of 1.5 mL/min, the Rj + Si × Rj + εijk , where Yijk is the dependent or response
split ratio was 1:30, and the injector and detector variable measured in the animal or experimental unit “k”
temperatures were 220 °C. The FAMEs were identified of sexual class “i” at FR “j”; μ is the population mean or
by a comparison of the FAME retention times with those global constant; Si is the effect of sexual class “i”; Rj is
of authentic standards (FAME mix components, Supelco®, the effect of FR “j”; Si × Rj is the interaction between
Bellefont, PA, USA) following the same injection method. effects of sexual class “i” and FR “j”; and ɛijk is
The results were quantified by normalizing the areas of unobserved random error. Tukey-Kramer’s test was used
the methyl esters and converted to mg/100 g of meat to compare the means with a significance level of 5%
using a conversion factor of 0.92 for the contribution of probability (P < 0.05), and the same criterion was adopted
fatty acids in lipids [22]. for interactions between the effects of sex and FR.
The concentrations of saturated fatty acids (SFAs),
unsaturated fatty acids (UFAs), monounsaturated fatty Results
acids (MUFAs), polyunsaturated fatty acids (PUFAs), ω6 Performance and carcass traits
and ω3 were calculated based on the fatty acid profile of There was an interaction (P < 0.05) between sex and FR
the meat. Lipid quality indexes were determined using for ADG, BWS and EBW (Table 2). In sum, females sub-
the sum of the desirable fatty acids [23], the thrombo- jected to AL intake presented similar ADG, BWS and
genicity index (TI), the atherogenicity index (AI) [24], EBW (P > 0.05) to those of entire males and castrated
and the ratio between fatty acids hypocholesterolemic males fed 30% FR (Table 3). Entire males fed AL pre-
acid and hypercholesterolemic acid (h/H) [25]. The sented higher (P < 0.05) ADG, BWS and EBW due to
activity of enzymes involved in lipid metabolism, such as their higher growth (Table 3, Fig. 1).
Δ9 desaturase in C16, Δ9 desaturase in C18 and elon- Except for SFT, the carcass traits that were analyzed
gase, were calculated according to the methods of (HCW, HCY, CCW, CCY and REA) were influenced
Malau-Aduli et al. [26]. (P < 0.05) by sex (Table 4). Entire males showed higher
means of HCW and CCW followed by castrated males
Feed chemical analysis and females. However, females and castrated males had a
To determine the chemical composition of the feed, tripli- higher HCY than did entire males. After cooling and con-
cate samples were dried at 55 °C for 72 h in a forced-air sidering the losses caused by this process, only females
oven, ground with a Willey mill (Tecnal®, São Paulo, had the highest yield (CCY), whereas castrated males did
Brazil) with a 1-mm sieve, and stored in airtight plastic not differ from the other sexes. The level of FR did not
containers (ASS®, São Paulo, Brazil). The samples were influence (P > 0.05) the HCY and CCY, which indicated
then stored in plastic jars with lids (ASS®, São Paulo, that the lower weights due to lower feed intake occurred
Brazil), labeled, and subjected to further laboratory ana- proportionately throughout the bodies of the animals.
lysis to measure the contents of dry matter (DM method HCW, CCW and SFT decreased with increasing FR (60%).
de Araújo et al. Journal of Animal Science and Biotechnology (2017) 8:46 Page 5 of 12

Table 2 Performance of Morada Nova lambs of different sexes subjected to feed restrictions
Variables Sexes Feed restrictions SEM P-value
Ent Cas Fem AL 30% 60% Sex Res Sex × Res
IBW, kg 14.4 14.7 15.5 14.6 14.6 14.4 0.158 0.711 0.860 0.348
b c b c
DMI, kg 0.66ª 0.60 0.52 0.80ª 0.62 0.36 0.009 <.0001 <.0001 0.418
BWS, kg 27.4ª 23.3b 19.9c 28.8a 24.4b 17.3c 0.308 <.0001 <.0001 0.047
b c b c
EBW, kg 21.0ª 18.1 16.1 23.0ª 19.0 13.3 0.214 <.0001 <.0001 0.038
ADG, g 106ª 71.7b 42.5c 116ª 80.8b 23.3c 2.317 <.0001 <.0001 0.029
Ent Entire males, Cas Castrated males, Fem Females, AL ad libitum intake, 30% 30% feed restriction, 60% 60% feed restriction, SEM standard error of the mean, Sex
sexes and Res = feed restriction, IBW initial body weight, DMI dry matter intake, BWS body weight at slaughter, EBW empty body weight, TWG total weight gain,
ADG average daily gain
a b c
Means followed by different letters differ between sexes according to a Tukey-Kramer test (P < 0.05)
a b c
Means followed by different letters differ between feed restrictions according to a Tukey-Kramer test (P < 0.05)

There was an interaction (P < 0.05) between sex and reduced (P < 0.05) concentration of IMF. Ash percent-
FR in neck weight (Table 4). However, no clear result age was higher (P < 0.05) in the meat from animals
was evidenced (Table 5). The weights of all commercial subjected to 60% FR.
cuts were influenced (P < 0.05) by sex and by FR
(Table 4). Entire males had heavier cuts, followed by Fatty acid profile
castrated males and females, which reflected the effects It was not possible to separate the peaks of conjugated
observed with the CCW. The weights of the commercial linoleic acid (CLA) isomers normally identified in meats
cuts decreased due to the reduction in feeding supply. from ruminants. Thus, the nomenclature used covered
all isomers (CLA). There was an interaction (P < 0.05)
Physicochemical meat quality between sex and FR for elaidic acid (C18:1 t9) and
L* was influenced (P < 0.05) by sex (Table 6). The color behenic acid (C22:0) in the meat of the LT muscle
parameters a*, b* and C* were not affected by sex or FR. (Table 8). However, after adjusting for multiple compari-
Castrated males and females did not differ (P > 0.05), sons, a clear interaction response was only detected for
and they had lower values (P < 0.05) than did entire elaidic acid (Table 9). Probably data set characteristics
males. Animals subjected to 60% FR showed a higher contributed to the absence of significance after the
24 h post-mortem pH in their meat compared to that of Tukey-Kramer test, contradicting the initial result of the
animals subjected to AL intake. ANOVA for the behenic acid. Females submitted to 60%
There was no effect (P > 0.05) of FR and sex on the FR had higher amounts of elaidic acid than did entire
protein content in meat from LT muscle (Table 6). males and castrated males submitted to 60% FR (Table 9).
However, an interaction (P < 0.05) between sex and FR There was an effect (P < 0.05) of sex only on the con-
for this variable was observed. Females fed AL had a centrations of linoleic acid (C18:2 c9c12) and arachi-
higher (P < 0.05) protein content than did females donic acid (C20:4 c5c8c11c14) (Table 8). Females
submitted to 30% FR (Table 7). The moisture content showed higher (P < 0.05) concentrations of these fatty
in the LT muscle of animals with AL intake was lower acids than did entire males and castrated males, which
(P < 0.05) compared to that of animals subjected to 30 showed no difference (P > 0.05) compared to the other
and 60% FR (Table 6). Animals submitted to AL intake categories. Meat of animals submitted to AL intake and
and 30% FR provided similar (P > 0.05) amounts of 30% FR showed similar (P > 0.05) and higher (P < 0.05)
IMF, whereas animals subjected to 60% FR had a values compared to those of animals subjected to 60%

Table 3 Interactions between sexes and feed restriction on performance of Morada Nova lambs
Variables Entire male Castrated male Female
AL 30% 60% AL 30% 60% AL 30% 60%
adA bdBC cdEF aeB bdBCD ceFG afCDE bdeDEF
BWS 35 27 20 28 25 17 23 21 15bfG
adA bdBC cdEF aeB beCD cdEF aeBCD bfDE
EBW 27 21 15 22 19 13 20 17 12ceG
ADG 164adA 110bdBC 47cdDE 114aeB 78beCD 21ceEF 72afD 54afD 5bfF
AL ad libitum intake, 30% 30% feeding restriction, 60% 60% feeding restriction. BWS body weight at slaughter (kg), EBW empty body weight (kg),
ADG average daily gain (g)
abc
; Means followed by different letters in same sexes differ by Tukey-Kramer test (P < 0.05)
def
; Means followed by different letters in same feeding restriction level differ by Tukey-Kramer test (P < 0.05)
ABCDEFG
; Means followed by different capital letters in same line differ by Tukey-Kramer test (P < 0.05)
de Araújo et al. Journal of Animal Science and Biotechnology (2017) 8:46 Page 6 of 12

40
The concentration of PUFAs was greater in the meat
of females and lower in the meat of entire males
35
(Table 10). However, the meat of females provided a
Ent AL
30
Cas AL higher ω6/ω3 ratio and AI in addition to presenting a
25 Ent 30 lower h/H ratio. The activity of the elongase enzyme was
Weight

Cas 30
20 higher in the LT muscle of castrated males. The sum of
Fem AL
15 Fem 30 SFA, UFA and PUFA was similar (P > 0.05) in the meat
10 Ent 60 of animals subjected to AL intake and 30% FR and was
Cas 60
5
higher (P < 0.05) compared to that in animals subjected
Fem 60
to 60% FR. The meat of animals subjected to 60% FR
0
120 130 149 169 189 209 230 240 provided a greater ω6/ω3 ratio, lower h/H ratio and
Age lower concentration of desirable fatty acids, in addition
Fig. 1 Evolution of animal weights during the experimental period. to a greater AI and TI.
Average animal weight (kg) of each treatment in relation to
approximate age in days from the beginning (120 d) to the end of
Discussion
the experiment (240 d); Ent AL = Entire males subjected to ad
libitum intake; Cas AL = Castrated males subjected to ad libitum Performance, carcass traits and physicochemical meat
intake; Fem AL = Females subjected to ad libitum intake; Ent 30 = Entire quality
males subjected to 30% feed restriction; Cas 30 = Castrated males Productions systems in semi-arid regions are based on
subjected to 30% feed restriction; Fem 30 = Females subjected to 30% the use of genetic resources with high adaptability and
feed restriction; Ent 60 = Entire males subjected to 60% feed restriction;
heat tolerance, which are heavily influenced by qualita-
Cas 60 = Castrated males subjected to 60% feed restriction; Fem
60 = Females subjected to 60% feed restriction tive and quantitative seasonality of food [32, 33]. An
example is the Morada Nova, an important indigenous
breed of hair sheep in northeastern Brazil that is used
FR for concentrations of palmitic acid (C16:0), palmitoleic for meat and skin production and is highly valued on
acid (C16:1c9), stearic acid (C18:0), oleic acid (C18:1c9) and the international market [34]. Weight loss has a strong
CLA. The concentration of myristic acid (C14:0) was higher impact on animal productivity [35], compromising the
(P < 0.05) in meat from animals subjected to 30% FR than animal welfare and income of farmers worldwide [33]. In
in meat from animals subjected to 60% FR. Meat from our study, we observed an absence of growth in females
animals subjected to 60% FR showed greater (P < 0.05) with 60% FR and a low ADG in lambs with 30% FR. This
concentrations of arachidonic acid and eicosapentaenoic effect is a response to a lower amount of nutrients [36]
acid (C20:5c5c8c11c14c17 - EPA) than did meat from because lambs have higher energy and protein require-
animals subjected to AL intake and 30% FR. ments for growth [32, 37], which demand higher intakes.

Table 4 Carcass characteristics and commercial cuts weight of Morada Nova lambs of different sexes subjected to feed restriction
Variables Sexes Feed restrictions SEM P-value
Ent Cas Fem AL 30% 60% Sex Res Sex × Res
HCW, kg 11.5ª 10.3b 8.99c 12.7ª 10.6b 7.51c 0.139 <.0001 <.0001 0.110
b
HCY, % 42.1 44.1ª 45.2ª 44.4 43.6 43.4 0.325 0.002 0.474 0.967
CCW, kg 11.4ª 10.18b 8.88c 12.58ª 10.51b 7.42c 0.138 <.0001 <.0001 0.106
CCY, % 41.7b 43.6ab 44.7ª 43.9 43.2 42.9 0.315 0.002 0.435 0.965
SFT, mm 1.32 1.10 1.12 1.55a 1.25ª 0.75b 0.060 0.283 <.0001 0.185
REA, cm2 11.0ª 10.3ab 9.36b 11.7ª 10.1b 8.88b 0.205 0.010 <.0001 0.649
a b c a b
Leg, kg 1.84 1.64 1.48 2.02 1.69 1.26c 0.022 <.0001 <.0001 0.109
a b b a b c
Loin, kg 0.58 0.48 0.42 0.65 0.50 0.32 0.011 <.0001 <.0001 0.102
Neck, kg 0.40a 0.35b 0.27c 0.40a 0.33b 0.28c 0.007 <.0001 <.0001 0.010
a b c a b c
Shoulder, kg 1.01 0.93 0.83 1.12 0.93 0.70 0.011 <.0001 <.0001 0.383
Rib, kg 1.07a 0.86b 0.78b 1.11a 0.98a 0.62b 0.024 0.0002 <.0001 0.329
a a b a b c
Lower rib, kg 0.85 0.79 0.63 0.96 0.80 0.51 0.019 0.0002 <.0001 0.485
Ent Entire males, Cas Castrated males, Fem Females, AL ad libitum intake, 30% 30% feed restriction, 60% 60% feed restriction, SEM standard error of the mean, Sex
sexes and Res = feed restriction, HCW hot carcass weight, HCY Hot carcass yield, CCW cold carcass weight, CCY cold carcass yield, SFT subcutaneous fat thickness,
REA Rib eye area
a b c
Means followed by different letters differ between sexes according to a Tukey-Kramer test (P < 0.05)
a b c
Means followed by different letters differ between feed restrictions according to a Tukey-Kramer test (P < 0.05)
de Araújo et al. Journal of Animal Science and Biotechnology (2017) 8:46 Page 7 of 12

Table 5 Interactions between sexes and feed restriction on the weight of commercial cuts of Morada Nova lambs
Variables Entire male Castrated male Female
AL 30% 60% AL 30% 60% AL 30% 60%
Neck 0.51adA 0.36bBC 0.33bdBC 0.42adAB 0.33bBC 0.29bdCD 0.28abeCD 0.32aBCD 0.21beD
AL ad libitum intake, 30% 30% feeding restriction, 60% 60% feeding restriction
abc
; Means followed by different letters in same sexes differ by a Tukey-Kramer’s test (P < 0.05)
def
; Means followed by different letters in same feeding restriction level differ by a Tukey-Kramer test (P < 0.05)
ABCD
; Means followed by different capital letters in same line differ by a Tukey-Kramer test (P < 0.05)

The study of van Harten et al. [38] showed that in animals Dietary restrictions have reduced the accumulation of
under FR, lipids are mobilized and transformed into energy. the body stores of fat and protein, which resulted in
This occurs to meet the net energy requirements for main- lighter carcasses and commercial cuts. Nutritional limi-
tenance. In addition to genetic and nutritional factors, sex tations reduce cell proliferation and differentiation in
is variable and impacts animal productive responses. tissues in response to reduced local production of
Effects of interactions between sex and FR showed that insulin-like growth factor-1 (IGF-1), as signaled by the
females performed similarly to entire males and state of relative resistance of growth hormone (GH) [45].
castrated males with 30% FR. Such effects reflect the In addition, in the post-absorptive state, non-esterified
empty cup weight gain, when females generally present fatty acids, glycerol, alanine and glycine are oxidized,
lower rates of protein deposition in the empty body [39]. which supplies part of the energy demand [46]. Thus, in
A sex effect is evident in the regulation of adiposity and situations of lower nutritional intake, some of the body
muscularity and has been attributed to sexual steroid energy reserves are consumed. This situation was
hormones [40] because testosterone promotes an increase observed in animals submitted to the levels of FR used
in body mass [41]. These effects were observed in this in this research.
study, in which entire males had higher carcass weights and The 24 h post-mortem pH was higher in the meat from
commercial cuts. Castrated males [6] and females [42] animals subjected to 60% FR, which may be related to
present higher HCY attributed to increased fat deposit the lower content of muscle glycogen caused by lower
during weight gain [43]. Furthermore, the higher central or feed intake and intense mobilization of reserves during
intra-abdominal accumulation of fat in male individuals the development of these animals. However, the pH
[44] contributed to greater proportions of non-carcass remained between 5.5 and 5.8, which is desirable for
components and consequently had a lower carcass yield. meat [47]. An increase in the pH of meat can increase

Table 6 Physicochemical quality of Morada Nova lamb meat of different sexes subjected to feed restriction
Variables Sexes Feed restrictions SEM P-value
Ent Cas Fem AL 30% 60% Sex Res Sex × Res
L* 39.7ª 38.2b 37.5b 38.2 38.4 38.8 0.244 0.003 0.657 0.659
a* 22.2 21.8 21.9 21.9 22.3 21.7 0.233 0.767 0.541 0.689
b* 9.83 9.43 8.80 9.36 9.93 8.77 0.220 0.197 0.126 0.866
C* 24.3 23.8 23.6 23.8 24.4 23.4 0.320 0.386 0.252 0.771
WHC, % 35.9 34.5 35.0 34.7 32.8 37.9 1.045 0.859 0.154 0.511
CWL, % 38.6 38.5 37.4 37.4 38.5 38.6 0.403 0.443 0.408 0.937
pH 5.61 5.60 5.66 5.57b 5.62ab 5.69ª 0.019 0.354 0.044 0.667
Shear force, N 42.9 46.1 39.2 44.0 41.4 42.8 1.623 0.236 0.808 0.702
TBARS 0.72 0.75 0.85 0.82 0.64 0.86 0.041 0.395 0.073 0.534
b
Moisture, % 76.5 75.9 75.8 75.4 75.9ª 76.9ª 0.146 0.167 0.001 0.064
Protein, % 18.6 18.9 19.0 18.9 18.5 19.1 0.117 0.332 0.139 0.018
Fat, % 3.99 4.32 4.18 4.74ª 4.70ª 3.05b 0.107 0.468 <.0001 0.680
Ash, % 0.92 0.90 0.93 0.92b 0.87b 0.98a 0.009 0.327 <.0001 0.155
Ent Entire males, Cas Castrated males, Fem Females, AL ad libitum intake, 30% 30% feed restriction, 60% 60% feed restriction, SEM standard error of the mean, Sex
sexes and Res = feed restriction, Color parameter L* lightness, 0 black and 100 white, Color parameter a* redness, ranges from green (−) to red (+), Color parameter
b* yellowness, ranges from blue (−) to yellow (+), Color parameter C* chroma; C* = (a*2 + b*2)1/2(MacDougall and Taylor, 1975), WHC water holding capacity, CWL
Cooking weight losses, TBARS 2-thiobarbituric acid reactive substance, MDA malondialdehyde
a b
Means followed by different letters differ between sexes according to a Tukey-Kramer test (P < 0.05)
a b
Means followed by different letters differ between feed restrictions according to a Tukey-Kramer test (P < 0.05)
de Araújo et al. Journal of Animal Science and Biotechnology (2017) 8:46 Page 8 of 12

Table 7 Interactions between sexes and feed restriction on Meat composition of Morada Nova lambs
Variables Entire male Castrated male Female
AL 30% 60% AL 30% 60% AL 30% 60%
Protein 18.24AB 18.70AB 18.89AB 18.57AB 18.66AB 19.46AB 19.99A 18.22B 18.93AB
AL ad libitum intake, 30% 30% feeding restriction, 60% 60% feeding restriction
abc
; Means followed by different letters in same sex differ by a Tukey-Kramer test (P < 0.05)
def
; Means followed by different letters in same feeding restriction level differ by a Tukey-Kramer test (P < 0.05)
AB
; Means followed by different capital letters in same line differ by a Tukey-Kramer test (P < 0.05)

the activity of cytochrome oxidase by reducing the (2.56 mg/g of meat) and reported a similar effect of
uptake of oxygen by myoglobin, which results in a sex on L*, recording 39.80% for females and 41.26%
purplish red color [48]. However, the higher pH in meat for entire males. More myoglobin may explain the
from animals subjected to 60% FR was not enough to color variation in the meat of female animals
influence the color and other quality parameters analyzed observed in the present study.
in this study. Testosterone, besides providing an anabolic effect [41],
L* was greatly influenced by the pigment contents, acts on plasma glucose levels in males but does not alter
especially those of hematin, myoglobin and their the phosphorylation of AMP-activated protein kinase in
forms [49]. Chromophores, such as myoglobin and muscle [40], which influences the concentration of IMF
hemoglobin, absorb visible light by increasing light in males to a small degree. A similar result was observed
penetration and consequently decreasing reflectance in this study because the percentage of meat fat was
[50]. Sañudo et al. [51] observed more myoglobin in similar between the sexes and did not reflect the effects
females (2.90 mg/g of meat) than in entire males of higher HCW in entire males.

Table 8 Fatty acids profile in the meat of the longissimus thoracis muscle of Morada Nova lambs of different sexes subjected
to feed restriction
Fatty acids, Sexes Feed restrictions SEM P-value
mg/100 g of meat
Ent Cas Fem AL 30% 60% Sex Res Sex × Res
C10:0 4.70 6.56 6.05 7.18 5.24 4.89 0.700 0.540 0.392 0.913
C12:0 32.8 59.7 43.6 44.9 35.1 56.1 6.476 0.254 0.432 0.648
C14:0 87.3 89.8 108 96.0ab 110a 78.5b 4.568 0.156 0.027 0.937
C14:1c9 8.51 10.3 9.95 10.9 9.64 8.26 0.610 0.447 0.255 0.436
C15:0 22.6 29.3 33.0 23.6 26.72 34.6 1.984 0.122 0.084 0.572
a a b
C16:0 912 914 966 1069 1080 642 30.20 0.711 <.0001 0.816
C16:1c9 41.2 40.3 43.8 49.0a 48.0a 18.3b 1.660 0.685 <.0001 0.551
C17:0 44.4 47.2 48.6 48.9 46. 9 44.4 1.047 0.277 0.245 0.314
C17:1c10 18.9 16.8 17.3 18.9 17.37 16.8 0.901 0.635 0.606 0.300
a a b
C18:0 620 687 649 695 701 561 19.03 0.377 0.008 0.770
C18:1c9 1474 1639 1474 186a 1814a 911b 41.65 0.192 <.0001 0.604
C18:1 t9 36.0 36.1 44.5 41.1 38.4 37.1 1.819 0.106 0.666 0.032
C18:2c9c12 70.1b 83.1ab 91.4a 82.8 79.1 82.8 2.388 0.005 0.768 0.582
CLA 3.96 4.92 4.54 5.29a 4.99a 3.14b 0.289 0.410 0.016 0.397
C18:3c9c12c15 18.5 21.6 21.0 22.2 20.5 18.4 0.701 0.201 0.109 0.510
b ab a b a
C20:4c5c8c11c14 31.2 37.3 39.2 33.0 33.2 41.6 1.236 0.045 0.010 0.862
C20:5c5c8c11c14c17 7.57 8.73 8.75 7.74b 7.39b 9.92a 0.216 0.061 <.0001 0.171
C22:0 8.08 9.13 9.02 8.24 8.51 9.48 0.253 0.201 0.143 0.042
Unidentified 550 583 565 618 617 474 - - - -
Ent Entire males, Cas Castrated males, Fem Females, AL ad libitum intake, 30% 30% feed restriction, 60% 60% feed restriction, SEM standard error of the mean, Sex
sexes and Res = feed restriction, C10:0 Capric acid, C12:0 Lauric acid, C14:0 Myristic acid, C14:1c9 Myristoleic acid, C15:0 Pentadecylic acid, C16:0 Palmitic acid,
C16:1c9 Palmitoleic acid, C17:0 Margaric acid, C17:1c10 Heptadecenoic acid, C18:0 stearic acid, C18:1c9 Oleic acid, C18:1 t9 Elaidic acid, C18:2c9c12 Linoleic acid,
C18:3c9c12c15 α-Linolenic acid, C20:4c5c8c11c14 Arachidonic acid, C20:5c5c8c11c14c17 Eicosapentaenoic acid, C22:0 Behenic acid, CLA Conjugated linoleic acids:
rumenic acid and their isomers
a b
Means followed by different letters differ between feed restrictions according to a Tukey-Kramer test (P < 0.05)
de Araújo et al. Journal of Animal Science and Biotechnology (2017) 8:46 Page 9 of 12

Table 9 Interactions between sexes and feed restriction on fatty acids profile in the meat of the longissimus thoracis muscle of
Morada Nova lambs
Fatty acids, mg/ Entire male Castrated male Female
100 g
AL 30% 60% AL 30% 60% AL 30% 60%
of meat
AB AB B AB AB B AB AB
C18:1 t9 38.6 41.8 28.6 45.5 34.6 28.1 39.2 39.9 54.5A
C22:0 8.2 8.6 7.4 9.1 8.0 10.2 7.3 8.9 10.8
AL ad libitum intake, 30% 30% feeding restriction, 60% 60% feeding restriction
AB
; Means followed by different capital letters in same line differ by Tukey-Kramer test (P < 0.05)

Variations in meat fat concentration occur mainly due meat in response to concentrations of plasmalogens
to changes in balance between dietary energy and nutri- [53]. The effects observed on the concentration of lino-
ent requirements [52]; thus, the least amount of energy leic and arachidonic acids in meat of females indicate
consumed by animals subjected to 60% FR decreased the that this incorporation can also occur in lamb meat but
deposition of IMF. FR of 60% meets the net energy was not observed due to the need for a more detailed
needs for maintenance and obviously does not prioritize analysis. The effect of the interaction between sex and
nutrients for the deposition of adipose tissue. However, FR on elaidic acid concentration shows that under 60%
30% FR did not cause significant changes to this balance FR, females accumulate a greater amount of this acid. It
or to the concentration of IMF in LT muscle. becomes important to know the unique physiological
functions of specific isomers as well as their origin. A
Fatty acid profiles portion of the trans-11 C18:1 isomer produced by rumi-
Studies on cattle have indicated increased incorporation nal microbes is converted into cis-9, trans-11 C18: 2 by
of long chain fatty acids into the phospholipids of heifer tissue desaturase [54]; however, this cannot occur with

Table 10 Fatty acid classes, ratios, indexes and enzyme activity in meat of longissimus thoracis muscle of Morada Nova lambs of
different sexes subjected to feed restriction
Index, Sexes Feed restrictions SEM P-value
mg/100 g
Ent Cas Fem AL 30% 60% Sex Res Sex × Res
of meat
ΣSFA 1731 1837 1853 1988 a
2007 a
1425 b
50.12 0.579 <.0001 0.770
ΣUFA 1707 1896 1748 2131 a
2069 a
1151 b
44.52 0.210 <.0001 0.602
ΣMUFA 1578 1741 1585 1981a 1926a 997b 44.10 0.248 <.0001 0.569
ΣPUFA 129 b
155 ab
163a
150 142 154 4.222 0.008 0.504 0.716
Σω6 99.0b 120a 131a 116 110 124 3.338 0.003 0.212 0.777
Σω3 26.10 30.3 28.8 29.9 27.9 27.4 0.909 0.188 0.515 0.628
ω6/ω3 3.84b 3.99b 4.53a 3.90b 3.96b 4.50a 0.076 0.003 0.009 0.784
h/H 1.60ab 1.77a 1.54b 1.74a 1.65ab 1.52b 0.027 0.008 0.015 0.661
a a
DFA 2328 2583 2397 2826 2769 1713b 56.65 0.185 <.0001 0.661
b b a
TI 1.78 1.70 1.81 1.64 1.72 1.93 0.031 0.351 0.003 0.741
AI 0.75b 0.72b 0.84a 0.70b 0.74b 0.87a 0.016 0.020 <.0001 0.706
Δ desaturase C16
9
4.26 4.17 4.28 4.36 4.28 4.07 0.124 0.925 0.631 0.872
Δ9 desaturase C18 69.8 69.0 67.9 72.6a 72.1a 61.9b 0.556 0.428 <.0001 0.431
b a b
Elongase 68.9 70.9 67.4 69.6 69.1 68.4 0.273 <.0001 0.240 0.179
Ent Entire males, Cas Castrated males, Fem Females, AL ad libitum intake, 30% 30% feed restriction, 60% 60% feed restriction, SEM standard error of the mean, Sex
sexes and Res = feed restriction
SFA saturated fatty acids (ΣC10:0, C12:0, C14:0, C15:0, C16:0, C17:0, C18:0, C22:0); UFA unsaturated fatty acid (ΣC14:1c9, C16:1c9, C17:1c9, C18:1c9, C18:1 t9,
C18:2c9c12, C18:2t9t12, C18:2c9t11, C18:3c6c9c12, C20:4c5c8c11c14, C20:5c5c8c11c14c17), MUFA monounsaturated fatty acid (ΣC14:1c9, C16:1c9, C17:1c9,
C18:1c9, C18:1 t9), PUFA polyunsaturated fatty acid (ΣC18:2c9c12, C18:2t9t12, C18:2c9t11, C18:3c6c9c12, C20:4c5c8c11c14, C20:5c5c8c11c14c17), ω6 (ΣC18:2c9c12,
C18:2t9t12, C20:4c5c8c11c14), ω3 (C18:3c9c12c15, C20:5c5c8c11c14c17), ω6:ω3 relation (Σω6/Σω3), h/H hypocholesterolemic hypercholesterolemic relation
(ΣC18:1c9, C18:2c9c12, C18:3c6c9c12, C20:4c5c8c11c14, C20:5c5c8c11c14c17 / ΣC14:0, C16:0), DFA Desirable fatty acids (ΣMUFA, PUFA, C18:0), TI Thrombogenicity
index [(ΣC14:0, C16:0, C18:0) / Σ(0,5xΣMUFA), (0,5xΣω6), (3xΣω3), (Σω3/Σω6)], AI Atherogenicity index {[ΣC12:0,(4xC14:0), C16:0] / [(Σω3,ω6) + (C18:1c9) + (Σ other
MUFA)]}; Enzyme activity Δ9 desaturase on fatty acid with 16 carbons {100[(C16:1c1) / (ΣC16:1c1, C16:0)]}; Enzyme activity Δ9 desaturase on fatty acid with 18
carbons {100[(C18:1c1) / (ΣC18:1c1, C18:0)]}; Elongase activity enzyme {100[(ΣC18:0,C18:1c9) / (ΣC16:0,C16:1c9,C18:0,C18:1c9)]}
a b
Means followed by different letters differ between sexes according to a Tukey-Kramer test (P < 0.05)
a b
Means followed by different letters differ between feed restrictions according to a Tukey-Kramer test (P < 0.05)
de Araújo et al. Journal of Animal Science and Biotechnology (2017) 8:46 Page 10 of 12

elaidic acid (C18:1 t9). The quality lipid indexes showed The activity of the elongase enzyme is related to
that the meat of female animals had a lipid profile with concentrations of palmitic, palmitoleic and oleic acids
less desirable characteristics compared to that of meat [11, 70]. Combined concentrations of these fatty acids
from other sexes due to a higher AI and a lower h/H resulted in increased activity of elongase in the muscle
ratio. The higher ω6/ω3 ratio would also indicate lower of castrated male animals. The lower activity of the Δ9
quality meat fat in females [55]; however, the latest desaturase enzyme C18 in animals subjected to 60% FR
recommendations suggest no rational limit for this ratio could be attributed to lower amounts of oleic acid
if the intake of ω6 and ω3 is within the proper range for present in the muscle of these animals [71].
human diets [56].
More feeding can explain the higher concentration of Conclusions
palmitic acid, palmitoleic acid, stearic acid and oleic acid Lambs in different sexes produced carcasses with differ-
in the meat of animals subjected to AL intake and 30% ent characteristics, and except for lightness, sex did not
FR. These effects reflect the results observed with the influence meat quality or chemical composition. How-
IMF content. As 30% FR did not significantly influence ever, females had a fatty acid profile in their meat that
the deposition of IMF, there was no effect on the com- was less favorable to consumer health. FR affected
position of fatty acids. The kinetics of the feed in the carcass traits without influencing the quality of the meat.
rumen may also have influenced the exposure time of IMF content decreased when animals were subjected to
the fatty acids to biohydrogenation [57]. The more 60% FR, but the lipid profile was less favorable to
severe restriction may have resulted in a lower passage consumer health.
rate, higher biohydrogenation, and higher deposition of
SFA. Furthermore, the incorporation of fatty acids syn- Abbreviations
thesized in muscle tissue may have been more effective a*: Redness; ADF: Acid detergent fiber; ADG: Average daily gain;
in animals subjected to AL intake and 30% FR. This is AI: Atherogenicity index; AL: Ad libitum; b*: Yellowness; BW: Body weight;
BWS: Body weight at slaughter; C*: Chroma; CCW: Cold carcass weight;
related to a more lipogenic substrate for de novo synthe- CCY: Cold carcass yield; CP: Crude protein; CWL: Cooking weight losses;
sis in muscle adipocytes [58, 59], especially glucose from DFA: Desirable fatty acids; DM: Dry matter; DMI: Dry matter intake;
the propionate originating from the fermentation of EBW: Empty body weight; EE: Ether extract; FR: Feed restriction; HCW: Hot
carcass weight; HCY: Hot carcass yield; IBW: Initial body weight; L*: Lightness;
carbohydrates in the rumen [60]. LL: longissimus lumborum; LT: longissimus thoracis; MDA: Malondialdehyde;
In our study, the lowest IMF deposit was in the meat MUFA: Monounsaturated fatty acid; NDF: Neutral detergent fiber; NFC: Non-
of animals subjected to 60% FR, which justified the fiber carbohydrate; NRC: National research council; PUFA: Polyunsaturated
fatty acid; REA: Rib eye area; SAS: Statistical analysis software; SFA: Saturated
lower concentration of CLA in meat, as CLA is prefer- fatty acid; SFT: Subcutaneous fat thickness; TBARS: 2-thiobarbituric acid reactive
entially deposited in triglycerides [61, 62]. Similarly, the substances; TI: Thrombogenicity index; TWG: Total weight gain; UFA: Unsaturated
lowest IMF deposits in the meat of animals subjected to fatty acid; WHC: Water holding capacity; ω3: Omega 3; ω6: Omega 6

60% FR may explain the higher concentration of long

Acknowledgements
chain PUFAs (EPA and AA) in meat, which are The authors would like to thank Coordination for the Improvement of
deposited primarily in phospholipids [63, 64]. The IMF Higher Education Personnel (CAPES).
consists of triglycerides deposited in adipocytes and
myofibril cytoplasm droplets, structural phospholipids Funding
This research was supported by the National Council for Scientific and
and cholesterol present in membranes [58]. Triglycerides Technological Development (CNPq-Brazil).
are more mobile, and phospholipids are more stable in
muscle [65]. Availability of data and materials
Based on the results of this study, lambs subjected to Not applicable.
60% FR can be expected to produce meat with fatty acid
concentrations that are less favorable to consumer Authors’ contributions
TA and ES conceived the study and conducted statistical analysis. TA, ES, AC
health and with a lower amount of desirable fatty acids, and MP conducted the laboratory analyses and prepared the manuscript. ES,
a lower h/H ratio, a higher AI and TI and a higher AC and EH managed the animal model and assisted with manuscript
proportion of SFA (55.3%) compared to those of animals preparation. IM and EH assisted with diet formulation, statistical analysis and
manuscript preparation. HM contributed to fatty acid analysis. LB and RO
submitted to AL intake (48.3%) and 30% FR (49.2%). A critically revised the manuscript and edited the language. LP contributed to
lipid profile favorable to the thrombogenicity and ather- conception and statistical analyses. All authors read and approved the final
ogenicity in the meat of animals subjected to 60% FR is manuscript.
related to myristic acid, palmitic acid and stearic acid
Competing interests
concentrations [24]. Values from 0.9 to 1.94 for TI and The authors declare that they have no competing interests.
0.59 to 1.15 for AI have been reported in the literature
[66–69]; the maximum values of TI (1.93) and AI (0.87) Consent for publication
found in this study were within this range. Not applicable.
de Araújo et al. Journal of Animal Science and Biotechnology (2017) 8:46 Page 11 of 12

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