Livestock Science

Effect of the genetic merit for subcutaneous fat thickness on the meat quality of steers
--Manuscript Draft--

Manuscript Number: LIVSCI-D-24-00015

Article Type: Research Paper

Section/Category: Genetics, Animal Genetic Resources and Breeding

Keywords: Aberdeen Angus, crossbreeding, expected progeny differences, fat, genetic value,
Nellore

Corresponding Author: Thiago Luis Alves Campos de Araújo

Federal University of Ceara
Fortaleza, CE BRAZIL

First Author: Thiago Luis Alves Campos de Araújo

Order of Authors: Thiago Luis Alves Campos de Araújo

Gelson Luis Dias Feijó

Marina de Nadai Bonin Gomes

Elzania Sales Pereira

Gilberto Romeiro de Oliveira Menezes

Ériklis Nogueira

Luiz Orcírio Fialho de Oliveira

Andrei Pereira Neves

Jaqueline Rodrigues Ferreira

Karla Izidio Latta

Douglas Gomes Vieira

Rodrigo da Costa Gomes

Abstract: Understanding subcutaneous adipose tissue deposition can contribute to the

productive efficiency of beef cattle, and the possible impacts of subcutaneous adipose
tissue deposition on meat quality must be studied. The objective of the present study
was to evaluate the effects of the genetic merit for backfat thickness (GMB) and
subcutaneous fat thickness (FT) on the quality of meat from Nellore (N_N) or crossbred
(1/2 Aberdeen Angus + 1/2 Nellore, A_N) steers. The animals were obtained by mating
Nellore cows with Nellore or Aberdeen Angus bulls. Both paternal genetic groups were
selected to provide positive (+, high GMB) or negative (-, low GMB) expected progeny
differences (EPDs) in terms of backfat thickness. Two hundred and fourteen animals
were obtained (N_N+ = 48, N_N- = 53, A_N+ = 51, and A_N- = 62), reared on a
pasture and finished in a feedlot or under semiconfined conditions. The steers were
slaughtered, and the finishing scores, FT, marbling scores and meat color aspects (L*,
a*, and b*) were evaluated for the carcasses (24 h postmortem). In meat samples
without and with maturation (for 14 days), the pH, color, exudation loss, cooking loss,
and shear force were analyzed, and the softening due to maturation was calculated.
The moisture content and intramuscular fat concentrations of the meat were
determined. A second design was proposed to assess the influence of the FT class
(low = 3.66 ± 0.58 mm; medium = 4.99 ± 0.49 mm and high = 6.81 ± 1.06 mm) on the
same variables as in the original study. The GMB influenced (P < 0.05) the FT and
carcass fattening but did not influence (P > 0.05) the physicochemical traits or the fat
content of the meat. Steers with a low, medium, or high FT provided meats with similar
physicochemical attributes (P > 0.05). Thus, the genetic merit for backfat thickness
increases the deposition of subcutaneous adipose tissue without influencing meat
quality attributes. It is not necessary to increase the FT of steers to levels above the
lowest class evaluated (3.66 ± 0.58 mm) to improve the physicochemical attributes of
the meat.

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Highlights

 A genetic merit for backfat thickness does not influence the meat quality in progenies

 Steers with different subcutaneous fat thickness (FT) have similar meat quality attributes

 It is not necessary to increase the FT of steers to improve physicochemical meat attributes.

Manuscript File Click here to view linked References

1 Effect of the genetic merit for subcutaneous fat thickness on the meat quality of steers

12
2
33 Thiago Luís Alves Campos de Araújoa*, Gelson Luís Dias Feijób, Marina de Nadai Bonin Gomesc, Elzania
4
54 Sales Pereiraa, Gilberto Romeiro de Oliveira Menezesb, Ériklis Nogueirab, Luiz Orcírio Fialho de Oliveirab,
6
7
5 Andrei Pereira Nevesd, Jaqueline Rodrigues Ferreirac, Karla Izidio Lattac, Douglas Gomes Vieirac, Rodrigo da
8
9
10 6 Costa Gomesb
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12 7
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14 8 a
Federal University of Ceará, Department of Animal Science, Pici Campus, 810, 60440-900, Fortaleza, CE,
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16 9 Brazil
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18 b
1910 Brazilian Agricultural Research Company, National Beef Cattle Research Center, Radio Maia Av., 830,
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2111 79106-550, Campo Grande, MS, Brazil
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2312 c
Federal University of Mato Grosso do Sul, Faculty of Veterinary Medicine and Animal Science, Senador
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2513 Filinto Muller Av., 2443, 79074-460, Campo Grande, MS, Brazil
26
27 d
2814 Staty University of Londrina, Department of Animal Science, Celso Garcia Cid Highway, Km 380, 86057-
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3015 970, Londrina, PR, Brazil
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3216
33
3417 * Corresponding author: TLACA: thiagotor4@hotmail.com
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36
37
18
38
3919 Abstract
40
4120 Understanding subcutaneous adipose tissue deposition can contribute to the productive efficiency of beef
42
4321 cattle, and the possible impacts of subcutaneous adipose tissue deposition on meat quality must be studied.
44
45
22 The objective of the present study was to evaluate the effects of the genetic merit for backfat thickness (GMB)
46
47
4823 and subcutaneous fat thickness (FT) on the quality of meat from Nellore (N_N) or crossbred (1/2 Aberdeen
49
5024 Angus + 1/2 Nellore, A_N) steers. The animals were obtained by mating Nellore cows with Nellore or
51
5225 Aberdeen Angus bulls. Both paternal genetic groups were selected to provide positive (+, high GMB) or
53
5426 negative (-, low GMB) expected progeny differences (EPDs) in terms of backfat thickness. Two hundred and
55
56
5727 fourteen animals were obtained (N_N+ = 48, N_N- = 53, A_N+ = 51, and A_N- = 62), reared on a pasture and
58
5928 finished in a feedlot or under semiconfined conditions. The steers were slaughtered, and the finishing scores,
60
6129 FT, marbling scores and meat color aspects (L*, a*, and b*) were evaluated for the carcasses (24 h
62
1
63
64
65
 30 postmortem). In meat samples without and with maturation (for 14 days), the pH, color, exudation loss,

131 cooking loss, and shear force were analyzed, and the softening due to maturation was calculated. The moisture
2
332 content and intramuscular fat concentrations of the meat were determined. A second design was proposed to
4
533 assess the influence of the FT class (low = 3.66 ± 0.58 mm; medium = 4.99 ± 0.49 mm and high = 6.81 ± 1.06
6
7
34 mm) on the same variables as in the original study. The GMB influenced (P < 0.05) the FT and carcass fattening
8
9
1035 but did not influence (P > 0.05) the physicochemical traits or the fat content of the meat. Steers with a low,
11
1236 medium, or high FT provided meats with similar physicochemical attributes (P > 0.05). Thus, the genetic merit
13
1437 for backfat thickness increases the deposition of subcutaneous adipose tissue without influencing meat quality
15
1638 attributes. It is not necessary to increase the FT of steers to levels above the lowest class evaluated (3.66 ± 0.58
17
18
1939 mm) to improve the physicochemical attributes of the meat.
20
2140
22
2341 Keywords: Aberdeen Angus, crossbreeding, expected progeny differences, fat, genetic value, Nellore
24
2542
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27
2843 1. Introduction
29
3044 Knowledge about adipose tissue deposition is important for beef production and quality. The
31
3245 presence of subcutaneous fat is usually related to the quality of the carcass and meat, mainly because of its
33
3446 thermoinsulating action that prevents "cold shortening" (Thompson, 2002; Warriss, 2010). Owing to its
35
36
37
47 relevance, covering fat has become a criterion for carcass classification and genetic selection and is mainly
38
3948 represented by subcutaneous fat thickness (FT). However, the pursuit to produce carcasses with a higher FT
40
4149 has given rise to paradoxes related to the amount and composition of fat in the carcass (Webb and O’Neill,
42
4350 2008).
44
45
51 Owing to the inconvenience of having excess fat on the carcass, adipose tissue needs to be trimmed
46
47
4852 during the standardization of cuts, which requires additional labor and reduces commercial yield, impacting
49
5053 the production efficiency (Kelly et al., 2019; Kenny et al., 2020). In addition, compared with that of other body
51
5254 tissues, adipocyte hypertrophy has a high energy cost (Bonnet et al., 2010). This contributes to an increase in
53
5455 the net requirement for gain and a reduction in the feed efficiency (Marcondes et al., 2015), which are more
55
56
5756 obvious as the animal matures (NRC, 2016). Thus, it can be inferred that the excessive deposition of fat in the
58
5957 carcass negatively impacts the production efficiency through a reduction in the feed efficiency and commercial
60
6158 yield.
62
2
63
64
65
 59 For these reasons, the meat industry in several countries delimits ideal margins of the FT in the

160 carcass according to market demands. In Brazil, for example, the margin is 3 to 10 mm (Gomes et al., 2021);
2
361 in Canada, it is between 2 and 7 mm (Bonny et al., 2017); in South Africa, it is between 3 and 7 mm (Bonny
4
562 et al., 2017); and in the USA, it is between 2.5 and 22.5 mm, with adjustments depending on the carcass (Hale
6
7
63 et al., 2013). These criteria encourage the use of tools that predict and control subcutaneous fat deposition in
8
9
1064 a search for production efficiency.
11
1265 Advances in the genetic improvement of beef cattle have leveraged the sector and generated
13
1466 increasingly accurate predictions of expected progeny differences (EPDs). This information has enabled the
15
1667 selection of animals with superior productive traits through the genetic merit of their parents (de Araújo et al.,
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1968 2022; Ferraz et al., 2018; Snijders et al., 2001). In this sense, the genetic merit for backfat thickness (GMB)
20
2169 can aid in the standardization of fattening in bovine carcasses, contributing to production strategies (de Araújo
22
2370 et al., 2022).
24
2571 Information on the genetic value of available genotypes can help the productive sector according to
26
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2872 the needs of each region. In Brazil, the beef cattle industry has joined forces to increase the fattening of the
29
3073 carcasses produced (Gomes et al., 2021), especially in the Nellore breed. On the other hand, the American
31
3274 market has sought to maintain or reduce the FT of carcasses and increase the amount and distribution of
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3475 marbling (Gonzalez and Phelps, 2018). In this context, obtaining divergent phenotypes through the genetic
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76 value of animals is quite convenient; however, it is important to know the extent of these effects on the quality
38
3977 of products from different genotypes.
40
4178 The objective of this study was to evaluate the effect of the GMB on meat quality of Nellore and
42
4379 crossbred (½ Aberdeen Angus + ½ Nellore) steers. Our study contemplated an approach using genetically and
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80 phenotypically divergent populations for the FT, which should further legitimize the effects of this factor on
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4881 meat quality. As a result, the effects of the FT class on meat quality of Nellore and crossbred (½ Aberdeen
49
5082 Angus + ½ Nellore) steers were also evaluated to test the hypothesis that the physicochemical attributes of
51
5283 meat from animals classified as having a high FT were different from those of meat from animals classified as
53
5484 having a medium or low FT.
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5785
58
5986 2. Materials and methods
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61
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 87 The procedures adopted in this research were evaluated and approved by the Ethics Committee on

188 the Use of Animals of the National Center for Beef Cattle Research (CNPGC) of the Brazilian Agricultural
2
389 Research Corporation (Embrapa) (CEUA/CNPGC Process No. 007-2015).
4
590
6
7
91 2.1. Experimental animals and treatments
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1092 The experimental animals were obtained by mating Nellore (N) cows with Nellore (N) or Aberdeen
11
1293 Angus (A) bulls. The two paternal genetic groups were selected for the prediction of expected progeny
13
1494 differences, namely, positive (+, high GMB) or negative (-, low GMB) for backfat thickness (EPDbft). The
15
1695 criteria and names of the bulls chosen to obtain the experimental animals are described in Table 1. A total of
17
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1996 214 animals from the matings described were used as follows: N_N+ = 48, N_N- = 53, A_N+ = 51, and A_N-
20
2197 = 62, with three experiments (I, II, and III) conducted in different periods and in different systems.
22
2398 Experiment I was conducted with calves born in 2016 (n = 68) in a pasture rearing system and was
24
2599 finished keeping castrated steers under semiconfined conditions. Experiment II was conducted with calves
26
27
100
28 born in 2017 (n = 70) in a pasture rearing system and was finished keeping castrated steers under semiconfined
29
101
30 conditions. Experiment III was carried out with calves born in 2017 (n = 76) in a pasture rearing system;
31
102
32 however, it was finished keeping noncastrated steers in a feedlot.
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34
103 The rearing systems used were similar for the animals in all three experiments; from calving to
35
36
104
37
weaning, the progenies were kept with the sows in Brachiaria spp. pastures and received mineral
38
105
39 supplementation ad libitum. The calves were weaned by definitive separation from the cows at 8.60 ± 0.55
40
106
41 months of age and had a body weight (BW) of 222.43 ± 28.83 kg (adjusted for 240 days).
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43
107
44
45
108 2.2. Rearing and finishing phases
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47
109
48 All the experiments were carried out by keeping the animals in a single lot on a Brachiaria spp. grass
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110
50 pasture under continuous stocking. The steers received a protein-energy supplementation (Appendix A), which
51
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111 was provided at a level of 0.3% of the average BW of the herd, and had free access to water. Every 45 days,
53
54
112 the BW of the steers was measured, and they finished the rearing phase with average BWs of 309.51 and
55
56
113
57 382.27 kg for the N_N and A_N animals, respectively. During this period, the average daily weight gain (ADG)
58
114
59 was 449.89 g/day for the N_N steers and 556.44 g/day for the A_N steers.
60
61
62
4
63
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65
115 Afterward, the animals in Experiments I and II were surgically castrated through a lateral incision

116
1 (Silva et al., 2009) and kept under semiconfined conditions. The steers remained in pastures formed with
2
117
3 Brachiaria brizantha cv. Xaraés, and their diet was supplemented with a concentrated feed (Appendix B)
4
5
118 offered daily at the level of 1.5% of the herd average BW.
6
7
119 The animals in Experiment III finished in a feedlot and received a total mix ration (TMR) twice a
8
9
120
10 day; they were not castrated. The TMR contained 50% of sorghum silage and 50% of a dry matter-based
11
121
12 concentrated feed. The formulation and composition details of the TMR are available in Appendix C.
13
14
122 At the end of the finishing phase, the steers were 26.24 ± 2.36 months old on average. During this
15
16
123 period, the ADG of the N_N animals was 0.98 kg/day, and the final BW was 539.70 kg; the ADG of the A_N
17
18
124
19 animals was 1.20 kg/day, and the final BW was 592.33 kg.
20
125
21
22
23
126 2.3. Slaughter and 24 h postmortem evaluations
24
25
127 The animals were slaughtered following humane procedures regulated by Brazilian legislation
26
27
128
28 (RIISPOA, 2017). The carcasses were individually identified, longitudinally sectioned, weighed and cooled at
29
129
30 2 °C for 24 h. After the cooling period, the visual finishing score was evaluated using a scale from 1 to 5, with
31
130
32 scores assigned in relation to the amount of carcass subcutaneous fat, as recommended by Gomes et al. (2021).
33
34
131 Next, the longissimus thoracis (LT) muscle was sectioned between the 12th and 13th ribs, making it possible
35
36
132
37
to measure the subcutaneous fat thickness (FT) with a digital caliper. The marbling score was calculated using
38
133
39 the standard classification proposed by the USDA (2017). For each degree of marbling, the following
40
134
41 numerical values were assigned in ascending order, including intermediate values: traces (tr- = 1, tr0 = 2 and
42
43
135 tr+ = 3); slight (sl- = 4, sl0 = 5 and sl+ = 6); small (sm- = 7, sm0 = 8 and sm+ = 9); modest (mo- = 10, mo0 =
44
45
136 11 and mo+ = 12); moderate (md- = 13, md0 = 14 and md+ = 15); and abundant (ab- = 16, ab0 = 17 and ab+
46
47
137
48 = 18).
49
138
50 Subsequently, the LT portion between the 10th and 12th ribs of each carcass was removed. After a
51
52
139 blooming period (~30 min), the instrumental color aspects of the meat and clean and unstained surfaces of the
53
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140 subcutaneous fat on the LT muscle were evaluated. A MiniScan XE Plus colorimeter (HunterLab, Reston,
55
56
141
57 USA) was used, with a 10° iris opening angle and a D65 illuminant (similar to sunlight, including the
58
142
59 ultraviolet region), programmed to measure the following parameters of the CIELab system: luminosity (L*
60
61
62
5
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143 24 h postmortem) and intensities of red (a* 24 h postmortem) and yellow (b* 24 h postmortem). The means

144
1 resulting from the measurements taken at three points in the samples were recorded.
2
145
3
4
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146 2.4. Meat quality analysis
6
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147 The LT muscle was used for meat quality evaluations. Three 2.5-cm thick steaks (I, II and III) were
8
9
148
10 removed in the same sequence and anatomical position in the caudal-cranial direction. Steak I of each sample
11
149
12 was subjected to maturation for fourteen days at 4 °C before being frozen (steak I – matured), while the other
13
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150 two steaks (steak II and steak III – unmatured) were vacuum packed and stored at -20 °C as soon as they were
15
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151 removed from the samples.
17
18
152
19 One of the subsamples of the unaged meat (steak III) was ground in a multiprocessor and subjected
20
153
21 to freeze-drying for 38 h in an LJJ freeze-dryer (JJ Científica, São Carlos, SP, Brazil). The moisture amount
22
23
154 removed during freeze-drying was determined by the difference in the weights of the sample before and after
24
25
155 the process. Subsequently, the total moisture content was determined using method 967.03 of AOAC (2016).
26
27
156
28 In lyophilized samples, the total fat content was determined according to the AM 5-04 procedure established
29
157
30 by AOCS (2017) using the ANKOM XT15 system (ANKOM Technology, Macedonia, USA) and petroleum
31
158
32 ether as the solvent.
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159 Other analyses were carried out in both steaks I – matured, and steaks II – unmatured, following the
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36
160
37
guidelines of AMSA (2016). For this purpose, the subsamples were thawed under refrigeration (4 °C), removed
38
161
39 from the packages, and placed on cooking grids. The mass of the exudate contained in the package was
40
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41 carefully weighed on an LS2 precision scale (Marte Científica, MG, Brazil) to obtain the exudation losses.
42
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163 Subsequently, the color aspects (L*, a*, and b*) were measured in each subsample as previously described.
44
45
164 The pH of the meat was measured using an HI99163 potentiometer (Hanna Instruments,
46
47
165
48 Woonsocket, USA). After the weight was recorded, each sample was baked in an LTedesco 6GNs electric
49
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50 oven (Tedesco Stoves, Caxias do Sul, RS, Brazil) at 190 °C with forced air circulation until the temperature in
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167 the geometric center of the subsample, which was individually measured using a digital thermocouple, reached
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168 71 °C. The weight loss was calculated by the weight difference before and after cooking.
55
56
169
57 With the aid of a corer, six cylindrical subsamples with a diameter of 1.27 cm were removed to
58
170
59 determine the shear force (SF) parallel to the direction of the muscle fibers of each cooked subsample (steaks
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171
61 I and II). Each of the cylinders was sheared on a 235 6X analog texturometer (Salter Brecknell, Birmingham,
62
6
63
64
65
172 England) with a 1.18-mm thick Warner–Bratzler blade. The force required for perpendicular rupture of the

173
1 cylinders was recorded. The mean of the six measurements performed to obtain the SF value was subsequently
2
174
3 calculated. The softening promoted by meat maturation was calculated by the difference between the SFs of
4
5
175 matured (steak I) and unmatured (steak II) samples.
6
7
176 A second design was proposed to evaluate the effects of the FT class on the same parameters as those
8
9
177
10 in the original study. After slaughter and evaluation of the FT in the progenies, the steers in each experiment
11
178
12 and breed were classified into three classes according to their individual FT values, regardless of their paternal
13
14
179 EPD for this trait. Animals with an FT above 0.5 standard deviations (FT > 0.5 σ, n = 82) in relation to the
15
16
180 general mean of each breed and system were considered to be in the high class; individuals with the SSE
17
18
181
19 between -0.5 and 0.5 standard deviations (-0.5 σ < SSE < 0.5 σ, n = 66) were considered to be in the middle
20
182
21 class; and animals with an FT lower than -0.5 σ (FT < -0.5 σ, n = 62).
22
23
183
24
25
184 2.5. Statistical analysis
26
27
185
28 A randomized block design was used, with a factorial arrangement of 2 × 2 (paternal breeds × GMB)
29
186
30 and 2 × 3 (paternal breeds × FT classes) according to the following mathematical models: 𝑌𝑖𝑗𝑙𝑚𝑛 = 𝜇 + 𝑃𝑖 +
31
32
187 𝐺𝑗 + 𝐿𝑙 + 𝐸𝑚 + (𝑃 × 𝐺)𝑖𝑗 + 𝜀𝑖𝑗𝑙𝑚𝑛 and 𝑌𝑖𝑘𝑙𝑚𝑛 = 𝜇+𝑃𝑖 + 𝐶𝑘 + 𝐿𝑙 + 𝐸𝑚 + (𝑃 × 𝐶)𝑖𝑘 + 𝜀𝑖𝑘𝑙𝑚𝑛 , where
33
34
188
35 Yijlmn = dependent variable observed in animal "n", paternal breed "i", GMB "j", lot "l" and experiment "m";
36
189
37 Yiklmn = dependent variable observed in animal "n", paternal breed "i", FT class "k", lot "l" and experiment
38
190
39 "m"; μ = population mean or global constant; Pi = effect of paternal race "i"; Gj = effect of GMB "j"; Ck =
40
41
191 effect of FT class "k"; Ll = effect of lot "l"; Em = effect of experiment "m"; (P x G)ij = interaction between
42
43
192
44
paternal breed "i" and GMB "j"; (P x C)ik = interaction between paternal breed "i" and FT class "k"; and ɛijlmn
45
193
46 and ɛiklmn = random error not observed.
47
194
48 The data were subjected to restricted maximum likelihood analysis using the PROC MIXED
49
50
195 procedure of SAS On Demand for Academics, considering the lot and experiment as random effects. The
51
52
196 random lot effect included individual variables such as age and weight at slaughter, while the random
53
54
197
55 experiment effect included variations in origin, year of birth and finishing system. Because there were more
56
198
57 than two levels, the Tukey–Kramer test was applied between the means of the FT classes. A significance level
58
59
199 of 5% was used.
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61
62
7
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200

201
1 3. Results
2
202
3 3.1. Effects of the GMB and paternal breed
4
5
203 The paternal mean EPDbft, as well as the FT and finishing scores of the animals, are shown in Table
6
7
204 2. The effects and interactions between the factors for these variables were discussed in a previous publication
8
9
205
10 (de Araújo et al., 2022).
11
206
12 There was no interaction effect (P > 0.05) between the paternal breed and GMB on the variables
13
14
207 evaluated 24 h postmortem. There was no effect (P > 0.05) of the GMB on the variables evaluated 24 h
15
16
208 postmortem (Table 2). Thus, animals with high and low GMB provided meats with similar marbling scores
17
18
209
19 and color aspects, regardless of the paternal breed (Nellore or Aberdeen Angus). There was an effect (P <
20
210
21 0.05) of the paternal breed for most of the 24-h postmortem evaluations, except for b* of meat and L* of
22
23
211 subcutaneous fat. The Nellore animals produced meat with lower marbling scores, a higher luminosity (L*)
24
25
212 and a lower red color intensity (a*), while the A_N steers produced fat on the LT muscle with higher red (a*)
26
27
213
28 and yellow (b*) intensities.
29
214
30 There was an interaction effect (P < 0.05) between the paternal breed and GMB on the aspects of the
31
215
32 color of the matured meat (Table 3). The L* and b* values of the N_N animals differed from those of the A_N
33
34
216 animals for the matured meat, with the first genetic group showing greater averages for these traits. There was
35
36
217
37
no effect (P > 0.05) of the GMB on the instrumental quality or fat and moisture contents of the meat. There
38
218
39 was an effect (P < 0.05) of the paternal breed on most of the instrumental quality variables and on the fat
40
219
41 content of the meat.
42
43
220 Regarding the instrumental aspects of color, the N_N animals provided meat with higher luminosity
44
45
221 (L*) and yellow intensity (b*) than did the A_N steers, both in the matured and unmatured meat (Table 3). In
46
47
222
48 the same way, regardless of the maturation process, the meat of the A_N animals lost more weight during
49
223
50 cooking; however, the SF of the meat of these animals was lower. With maturation, the SF of the N_N meat
51
52
224 decreased more than that of the A_N steer meat. While the SF of the A_N meat decreased by 12.6 Newtons
53
54
225 upon maturation, that of the N_N meat decreased by 17.30 Newtons. However, the same result was not
55
56
226
57 obtained for the relative value (%) of the shear force before and after maturation.
58
227
59
60
228
61 3.2. Effect of the subcutaneous fat thickness class
62
8
63
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65
229 The paternal means of EPDbft, as well as the FT and the finishing scores of the animals, are presented

230
1 in Table 4. As recommended, the FT and the finishing scores that differed (P < 0.05) between the FT classes
2
231
3 and effects of the paternal breed, as observed in Study I, were repeated in Study II.
4
5
232 There was no interaction effect (P > 0.05) between the paternal breed and FT class on the variables
6
7
233 evaluated 24 h postmortem (fresh meat - Table 4). Similarly, there was no effect (P > 0.05) of the FT class on
8
9
234
10 marbling or on instrumental color attributes evaluated in fresh meat, except for b*. The yellow fluorescence
11
235
12 intensity was greater in the fresh meat of animals with a high FT than in those with a low FT, while this
13
14
236 parameter in animals with a medium FT did not differ from that of the others.
15
16
237 There was no interaction effect (P > 0.05) between the paternal breed and FT class on the
17
18
238
19 physicochemical attributes of the matured and unaged meat (Table 5). Except for the intramuscular fat content,
20
239
21 the FT class did not influence the physicochemical attributes of the meat quality. The intramuscular fat content
22
23
240 of the meat of the animals classified as having a high FT was greater than that of the other groups (classified
24
25
241 as having a medium or low FT), and the two groups of meat did not differ from each other.
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27
242
28
29
243
30 4. Discussion
31
244
32 4.1. Effects of the GMB and paternal breed
33
34
245 The results showed that there were few and inconclusive interactions between the GMB and paternal
35
36
246
37
breed factors on the physicochemical attributes of meat. As a result, it can be inferred that the factors studied
38
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39 are independent in relation to most of the variables analyzed.
40
248
41 Our research revealed that animals with divergent GMB expressed this trait in their phenotypes,
42
43
249 which resulted in individuals with distinct FT and carcass finish times, indicating that there is a distinction in
44
45
250 lipid metabolism between these animals (de Araújo et al., 2022). However, even the animals with low GMB
46
47
251
48 did not present with FT values lower than the minimum of 3 mm, as recommended to avoid damage caused
49
252
50 by rapid cooling (Prado et al., 2008), which may explain the absence of an effect of the GMB on the
51
52
253 physicochemical attributes of the meat.
53
54
254 The absence of an effect of the GMB on the marbling score and fat content of the meat indicates that
55
56
255
57 the genetic mechanism that modulates lipid metabolism in subcutaneous adipose tissue (SAT) does not
58
256
59 influence the rate of fat deposition in intramuscular adipose tissue (IAT). It is well documented that simple
60
257
61 genetic correlations are low between FT and marbling, parameters related to fat deposition in SAT and IAT,
62
9
63
64
65
258 respectively (Bertrand et al., 2001; Londoño-Gil et al., 2022). The divergent biochemical mechanisms between

259
1 these tissues are evident, with an emphasis on the primary substrates for lipogenesis, namely, glucose in IAT
2
260
3 and acetate in SAT (Rhoades et al., 2007; Smith and Crouse, 1984). However, genetic mechanisms that
4
5
261 regulate adipogenesis and the interdependencies between different tissues are still being elucidated (Urrutia et
6
7
262 al., 2020).
8
9
263
10 When analyzing RNA sequencing data from the IAT, SAT, and perirenal adipose tissue of cattle,
11
264
12 Sheng et al. (2014) revealed that there was a pattern involving 110 differentially expressed genes between the
13
14
265 tissues. The authors also observed a greater occurrence of alternative splicing events in SAT, which denotes a
15
16
266 greater dynamism in the translation of genetic information into biological functions. Hudson et al. (2020)
17
18
267
19 reinforced the wide divergence of gene expression between the SAT and IAT and highlighted that the main
20
268
21 differences were related to the metabolism of carbohydrates, cholesterol, and retinoic acid. This evidence may
22
23
269 be key to explaining how the GMB modulates lipid metabolism only in SAT. However, in our study, the gene
24
25
270 expression parameters necessary to prove these hypotheses were not evaluated.
26
27
271
28 The results observed among the studied breeds have been widely described in the literature,
29
272
30 indicating that crossbred animals (1/2 Aberdeen Angus + 1/2 Nellore, A_N) provide meats with greater
31
273
32 marbling and intramuscular fat (Chardulo et al., 2013; Oliveira et al., 2011), greater cooking loss (Oliveira et
33
34
274 al., 2011), a lower shear force (Aroeira et al., 2017; Barcellos et al., 2017; Miguel et al., 2014; Oliveira et al.,
35
36
275
37
2011), and lower luminosity and yellow intensity (Aroeira et al., 2017; Miguel et al., 2014) than do Nellore
38
276
39 animals. Our study reinforces the benefits of physicochemical parameters for meat quality through the use of
40
277
41 Aberdeen Angus as a paternal breed in adopted production systems. In addition, such benefits can be observed
42
43
278 even in a population of individuals selected for divergent fattening carcass traits.
44
45
279
46
47
280
48 4.2. Effects of subcutaneous fat thickness class
49
281
50 Several studies, including a meta-analysis (Boito et al., 2017), empirical studies (Maggioni et al.,
51
52
282 2012; Pflanzer and de Felício, 2011), and a multivariate principal component analysis (Baldassini et al., 2017),
53
54
283 have evaluated the impact of FT on beef meat quality. In the present research, we evaluated the effects of FT
55
56
284
57 on the qualitative attributes of meat via controlled genetic, environmental and maturity factor experiments,
58
285
59 with a large and comprehensive sample population.
60
61
62
10
63
64
65
286 The highest intensity of the color attributes a* and b* in fresh meat (24 h postmortem) of the animals

287
1 with a high FT is related to a slight variation in the internal temperature of the muscle during cooling in
2
288
3 response to the thermal insulation effect obtained with the higher finish in the carcass of these animals
4
5
289 (MacDougall, 1982). Rigor mortis settles more rapidly at higher chamber temperatures (Young et al., 1999),
6
7
290 and during this process, the myofibrils become more reflective (MacDougall, 1982). However, the color
8
9
291
10 attributes measured during a short postslaughter period are not definitive parameters of the color appearance
11
292
12 and have a low correlation with the meat color in the following days or weeks (Young et al., 1999); this was
13
14
293 evidenced by the similarity of the color attributes of the unmatured and matured meat between the FT classes.
15
16
294 The higher intramuscular fat content in the meat of steers with a high FT observed in this study is an
17
18
295
19 indication that these animals could be in transition to an increase in adipocyte differentiation in the IAT.
20
296
21 According to Du et al. (2013), with the development of the animal, preadipocytes present in various body
22
23
297 tissues mature to adipocytes by hyperplasia, and the IAT is the last in the order of priority of differentiation.
24
25
298 However, greater fat deposition was not visually perceptible since the marbling score was not influenced. It is
26
27
299
28 important to emphasize that the progenies were slaughtered relatively young (26.24 ± 2.36 months, 0 to 2
29
300
30 permanent teeth), and these effects could be more obvious with the advance of maturity in these animals.
31
301
32 This study reinforces that an FT above the minimum value (2 to 3 mm) does not greatly influence
33
34
302 the physicochemical attributes of meat quality, even in populations that are genetically divergent for
35
36
303
37
subcutaneous fat deposition. Considering the productive and economic impacts of promoting high fat
38
304
39 deposition on carcasses, it can be inferred that it would not be necessary to increase the FT much above 2 to 3
40
305
41 mm to improve the physicochemical attributes. In this sense, the increase in the proportion of fat in the carcass
42
43
306 at medium and high levels is justified only for meeting the demands for cuts with greater fat thickness, such
44
45
307 as those intended for the preparation on grills or under dry heat, such as in barbecues.
46
47
308
48
49
309
50 5. Conclusion
51
52
310 Animals with divergent GMB values provide meats with similar physicochemical attributes.
53
54
311 However, in populations with divergent GMB values, the physicochemical attributes of meat quality are
55
56
312
57 improved when the Aberdeen Angus breed is used for Nellore dams. The FT class did not influence most of
58
313
59 the physicochemical attributes of the meat, indicating that it would not be necessary to increase the FT much
60
314
61 above 2 to 3 mm for meat protection.
62
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315

316
1 CRediT authorship contribution statement
2
317
3 Thiago L. A. C. de Araújo: Formal analysis, Investigation, Writing – original draft, Writing – review
4
5
318 & editing. Gélson L. D. Feijó: Formal analysis, Methodology, Resources. Marina de N. B. Gomes:
6
7
319 Methodology, Resources, Writing – original draft. Elzania S. Pereira: Supervision, Resources. Gilberto R.
8
9
320
10 de O. Menezes: Conceptualization, Investigation, Writing – review & editing. Ériklis Nogueira:
11
321
12 Conceptualization. Luiz O. F. de Oliveira: Formal analysis, Project administration. Andrei P. Neves: Formal
13
14
322 analysis. Jaqueline R. Ferreira: Formal analysis. Karla I. Latta: Formal analysis. Douglas G. Vieira:
15
16
323 Formal analysis. Rodrigo da C. Gomes: Conceptualization, Resources, Formal analysis, Writing – review &
17
18
324
19 editing, Supervision, Project administration.
20
325
21
22
23
326 Funding
24
25
327 National Council for Scientific and Technological Development (CNPq): process: 420641/2016-5
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27
328
28
29
329
30 Conflict of interest
31
330
32 All authors declare no conflicts of interest.
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331
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37
Acknowledgements
38
333
39 Embrapa for enabling the development of the project, assisting financially and with field structure and
40
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41 laboratories. The Federal Universities of Ceará (UFC) and Mato Grosso do Sul (UFMS) for their support to
42
43
335 students. The authors thank the National Council for Scientific and Technological Development (CNPq) for
44
45
336 financial assistance to the project (process: 420641/2016-5) and to the students. To the owners and
46
47
337
48 administrators of the Primavera/Panorama, São Bento do Abobral and São Miguel da Catechesis farms, for
49
338
50 their fundamental contribution in obtaining the animals and conducting part of the research. The authors would
51
52
339 like to thank Connan Animal Nutrition for technical collaboration and supply of nutritional supplements. To
53
54
340 the Naturafrig Alimentos for its contribution to the collection of data and samples.
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341
57
58
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59 References
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462 Table 1
463 Criteria used for the selection of bulls and information on the selected bulls used for the production of the
464
1 animals studied
2 Product1 %Year3 Acc4 %Bft5 Acc4 Summary2
3 N+ <30% >30% <5% >30% ANCP (2014)
4 N- <30% >30% 99% >30% ANCP (2014)
5
6 A+ <30% >30% <10% >30% AAA (2014)
7 A- <30% >30% >65% >30% AAA (2014)
8 Bull name %Year 3
Acc 4
%Bft 5
EPDbft 6
Acc4 NOS7
9
N+ 48
10
11 REM Quilano 4 0.93 0.1 1.09 0.88 9
12 Relevo do Golias 25 0.82 1 0.50 0.73 7
13 Quarto JHV 4 0.87 2 0.44 0.77 14
14 Maximus da Matinha 2 0.79 2 0.36 0.74 9
15
16 Samaritano JHV 3 0.84 2 0.38 0.73 9
17 N- 53
18 Golias FIV Col 7 0.71 99 -0.54 0.68 12
19
Nougan do API 6 0.77 99 -0.37 0.68 8
20
21 Faracatu JB da Gur 1 0.78 99 -0.33 0.64 11
22 Imperioso Col 3 0.74 99 -0.22 0.66 15
23 Donato de Navirai CSCC 3 0.93 99 -0.28 0.79 7
24
A+ 51
25
26 Sitz Gunslinger 11612 3 0.36 3 2.21 0.33 15
27 EF Complement 8088 5 0.82 1 2.13 0.64 8
28 Sitz RLS Undertaker 11582 5 0.36 3 1.98 0.34 8
29
Connealy Final Product 5 0.93 2 1.93 0.68 8
30
31 SAV Plainsman 0468 15 0.33 2 1.91 0.33 12
32 A- 62
33 Connealy Lead On 30 0.96 99 -2.34 0.72 4
34
SAV Blacksmith 2815 3 0.38 97 -1.30 0.34 16
35
36 Quaker Hill Rampage 0A36 1 0.34 95 -1.14 0.39 11
37 HA PROGRAM 5652 17 0.91 95 -1.14 0.66 19
38 Circle A Incentive 10 0.67 85 -0.64 0.70 12
39
40
Total 214
1
465
41 Product: N+ = Nellore bull with a high genetic merit for backfat thickness; N- = Nellore bull with a low
466
42 genetic merit for backfat thickness; A+ = Aberdeen Angus bull with a high genetic merit for backfat thickness;
467
43 A- = Aberdeen Angus bull with a low genetic merit for backfat thickness; 2 Summary: Information was
44
468 obtained from the bull summaries of the National Association of Breeders and Researchers and the American
45
469 Angus Association (ANCP, 2014; AAA, 2014); 3 %Year = Percentile of bulls for yearling weight; 4 Acc =
46
470 Accuracy of the trait; 5 %Bft = Percentile of bulls for subcutaneous fat thickness; 6 EPDbft = Expected progeny
47
471 difference for backfat thickness (in millimeters); 7 NOS = Number of offspring used in the study
48
472
49
50
51
52
53
54
55
56
57
58
59
60
61
62
18
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473 Table 2
474 Effects of paternal breed and genetic merit for backfat thickness on marbling and the color of fresh meat (24 h
475
1 postmortem) and subcutaneous fat of steers
2 Paternal1 GMB2 P-Value
3 Variable Angus Nellore Low High SEM3
4 Paternal1 GMB2 Interaction
(n=113) (n=101) (n=115) (n=99)
5
6 EPD4 backfat thickness (mm) - - -0.79 1.32 0.08 - <.0001 <.0001
7 Backfat thickness (mm) 5.26 4.64 4.49 5.40 0.11 0.0071 <.0001 0.3893
8 Finishing score 2.55 2.58 2.44 2.69 0.04 0.6392 0.0002 0.1276
9 Marbling score 6.07 4.69 5.48 5.28 0.16 <.0001 0.4714 0.9348
10
11 L* fresh meat (%) 34.63 35.89 35.02 35.50 0.16 <.0001 0.0952 0.1689
12 a* fresh meat (%) 19.80 18.79 19.27 19.32 0.16 0.0002 0.8197 0.4612
13 b* fresh meat (%) 15.55 15.38 15.42 15.52 0.14 0.5128 0.6577 0.9454
14 L* subcutaneous fat (%) 73.90 74.71 74.05 74.26 0.21 0.2525 0.5960 0.6171
15
16 a* subcutaneous fat (%) 9.92 8.94 9.28 9.57 0.15 0.0029 0.3283 0.4522
17 b* subcutaneous fat (%) 22.34 20.93 21.56 21.71 0.23 0.0008 0.6695 0.8265
18 Angus Nellore
19 Factors interaction
Low High Low High
20 4 d a c
21 EPD backfat thickness (mm) -1.17 2.05 -0.35 0.54b
1
476
22 Paternal breed of progenies; 2 Genetic merit for backfat thickness; 3 Standard error of the mean; 4 Expected
23
477 difference in progeny; L* = Luminosity. a* = red color intensity. b* = yellow color intensity; abc Means with
24
478 distinct letters on the same lines differ by the Tukey–Kramer test at 5% significance.
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
19
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479 Table 3
480 Effects of paternal breed and genetic merit for backfat thickness on meat quality of steers
1 Paternal1 GMB2 P-Value
2 Variable Angus Nellore Low High SEM3
3 Paternal1 GMB2 Interaction
(n=113) (n=101) (n=115) (n=99)
4 pH 5.61 5.60 5.60 5.60 0.01 0.4615 0.9078 0.8851
5
6 L* (%) 35.65 37.19 36.40 36.44 0.17 <.0001 0.9202 0.2857
7 a* (%) 12.93 13.07 13.18 12.82 0.13 0.5362 0.0886 0.2145
8 b* (%) 12.07 12.58 12.41 12.25 0.08 0.0019 0.3039 0.1805
9 Exudating loss (%) 8.53 9.44 9.06 8.97 0.30 0.1538 0.8515 0.8382
10
11 Cooking loss (%) 23.21 24.35 23.79 23.77 0.34 0.0749 0.9832 0.7342
12 SF4 (N) 54.61 71.91 62.89 63.62 1.40 <.0001 0.7525 0.9113
13 Moisture (g/100g meat) 81.08 80.89 80.95 81.02 0.40 0.8169 0.9182 0.2307
14 Fat (g/100g meat) 1.40 1.13 1.32 1.22 0.04 0.0045 0.2314 0.6393
15 5
16 Maturated meat
17 pH 5.70 5.67 5.69 5.68 0.01 0.1035 0.8523 0.4032
18 L* (%) 37.36 39.21 38.24 38.34 0.17 <.0001 0.7404 0.0061
19 a* (%) 12.09 12.28 12.18 12.19 0.13 0.4287 0.9380 0.2460
20
21 b* (%) 12.22 12.92 12.58 12.56 0.08 <.0001 0.8679 0.0024
22 Exudating loss (%) 9.54 10.17 9.60 10.11 0.25 0.1936 0.2358 0.5956
23 Cooking loss (%) 23.67 23.70 23.80 23.57 0.34 0.9637 0.6583 0.8783
24 SF4 (N) 35.71 48.32 40.61 43.41 1.02 <.0001 0.1004 0.8866
25 5
26 Softening by maturation (N) -17.87 -25.04 -21.30 -21.61 1.11 0.0018 0.8786 0.2711
27 Softening by maturation5 (%) -32.06 -31.90 -32.55 -31.40 1.26 0.9533 0.6191 0.2561
28 Angus Nellore
29 Factors interaction
Low High Low High
30
31 L* of maturated meat5 (%) 36.90c 37.81bc 39.58a 38.86ab
32 b* of maturated meat5 (%) 12.03c 12.41bc 13.14a 12.71ab
1
33
481 Paternal breed of progenies; Genetic merit for backfat thickness; Standard error of the mean; 4 Shear force;
2 3
34
482 5
Maturation for 14 days; abc Means with distinct letters on the same lines differ by the Tukey–Kramer test at
35
483 5% significance.
36
484
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
20
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15
16
17
18
19
20
485 Table 4
21
486 Effects of the paternal breed and subcutaneous fat thickness (FT) class1 on marbling and the color of fresh meat (24 h postmortem) and subcutaneous fat of steers
22
23 Paternal2 FT class1 P-Value
3
24 Variable Angus Nellore Low Medium High SEM
25 Paternal2 FT1 Interaction
(n=113) (n=101) (n=82) (n=66) (n=62)
26
EPD4 backfat thickness (mm) 0.34 0.08 -0.07c 0.18b 0.51a 0.09 0.1508 0.0214 0.3322
27
c b
28 Backfat thickness (mm) 5.38 4.93 3.66 4.99 6.81a 0.11 0.0003 <.0001 0.7237
29 Finishing score 2.51 2.62 2.40 b
2.50 b
2.81 a
0.03 0.1207 <.0001 0.0321
30
31 Marbling score 6.11 4.75 5.04 5.58 5.68 0.16 <.0001 0.1117 0.5761
32 L* fresh meat (%) 34.59 35.86 35.13 34.91 35.64 0.16 0.0002 0.1386 0.4712
33 a* fresh meat (%) 19.82 18.78 19.10 b
19.09 b
19.73 a
0.16 0.0002 0.0471 0.4605
34 b b a
35 b* fresh meat (%) 15.46 15.39 15.18 15.24 15.85 0.13 0.8111 0.0269 0.4249
36 L* subcutaneous fat (%) 73.85 74.46 73.98 74.10 74.40 0.21 0.1800 0.6774 0.8827
37
38 a* subcutaneous fat (%) 9.92 8.91 9.26 9.52 9.45 0.15 0.0022 0.7454 0.9159
39 b* subcutaneous fat (%) 22.44 20.86 21.60 21.65 21.69 0.23 0.0002 0.9748 0.7665
40
487 1
High = FT > 0.5σ for each paternal breed. Medium = -0.5σ < FT < 0.5σ for each paternal breed. and Low = FT < -0.5σ for each paternal breed; Paternal breed of
2
41
488 progenies; 3 Standard error of the mean; 4 Expected progeny difference; abc Means with distinct letters on the same lines differ by the Tukey–Kramer test at 5%
42
489 significance.
43
490
44
45
46
47
48
49
50
51
52
53
54
55
56
57
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59
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21
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15
16
17
18
19
20
491 Table 5
21
492 Effects of the paternal breed and subcutaneous fat thickness (FT) class1 on the meat quality of steers
22
23 Paternal2 FT class1 P-Value
3
24 Variable Angus Nellore Low Medium High SEM
25 Paternal2 FT1 Interaction
(n=113) (n=101) (n=82) (n=66) (n=62)
26
pH 5.62 5.59 5.61 5.61 5.60 0.01 0.2535 0.8022 0.4415
27
28 L* (%) 35.62 37.21 36.28 36.38 36.58 0.17 <.0001 0.7376 0.6898
29 a* (%) 12.96 13.14 12.77 13.17 13.22 0.14 0.4610 0.1420 0.9316
30
31 b* (%) 12.09 12.61 12.19 12.29 12.58 0.08 0.0020 0.1174 0.7019
32 Exudating loss (%) 8.70 9.47 8.95 9.30 9.01 0.30 0.2004 0.8448 0.6286
33 Cooking loss (%) 23.22 24.26 23.88 22.96 24.37 0.34 0.1084 0.1707 0.1588
34 4
35 SF (N) 55.21 71.15 64.03 62.51 62.99 1.42 <.0001 0.8486 0.2916
36 Moisture (g/100g meat) 81.13 80.90 81.69 81.30 80.07 0.41 0.7870 0.1845 0.5548
37 b b a
38 Fat (g/100g meat) 1.39 1.14 1.17 1.19 1.44 0.04 0.0075 0.0130 0.2489
4
39 Maturated meat
40 pH 5.62 5.59 5.61 5.61 5.60 0.01 0.2535 0.8022 0.4415
41
42 L* (%) 35.62 37.21 36.28 36.38 36.58 0.17 <.0001 0.7376 0.6898
43 a* (%) 12.96 13.14 12.77 13.17 13.22 0.14 0.4610 0.1420 0.9316
44
45
b* (%) 12.09 12.61 12.19 12.29 12.58 0.08 0.0020 0.1174 0.7019
46 Exudating loss (%) 9.54 10.31 9.80 9.98 10.00 0.25 0.1284 0.9025 0.6493
47 Cooking loss (%) 23.65 23.62 23.86 23.44 23.62 0.34 0.9619 0.7933 0.9387
48 4
49 SF (N) 35.57 48.22 42.15 41.24 42.29 1.03 <.0001 0.8702 0.2843
50 Softening by maturation5 (N) -32.67 -32.11 -32.55 -31.22 -33.39 1.28 0.8356 0.7583 0.1356
51
493 1
High = FT > 0.5σ for each paternal breed. Medium = -0.5σ < FT < 0.5σ for each paternal breed. and Low = FT < -0.5σ for each paternal breed; Paternal breed of
2
52
494 progenies; 3 Standard error of the mean; 4 Maturation for 14 days; abc Means with distinct letters on the same lines differ by the Tukey–Kramer test at 5% significance.
53
495
54
55
56
57
58
59
60
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22
63
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Highlights

Declaration of interests

☒ The authors declare that they have no known competing financial interests or personal relationships
that could have appeared to influence the work reported in this paper.

☐The authors declare the following financial interests/personal relationships which may be considered
as potential competing interests:
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