Meat Science 183 (2022) 108599

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Meat Science
journal homepage: www.elsevier.com/locate/meatsci

Feeding strategies impact animal growth and beef color and tenderness
Juan F. Morales Gómez a, Daniel S. Antonelo b, Mariane Beline a, Bruna Pavan a,
Danilo B. Bambil a, Paulo Fantinato-Neto a, Arlindo Saran-Netto a, Paulo Roberto Leme a, Rodrigo
S. Goulart a, David E. Gerrard c, Saulo L. Silva a, *
a
Department of Animal Science, College of Animal Science and Food Engineering, University of São Paulo, Pirassununga, SP 13635-900, Brazil
b
Department of Animal Nutrition and Production, College of Veterinary Medicine and Animal Science, University of São Paulo, Pirassununga, SP 13635-900, Brazil
c
Department of Animal and Poultry Sciences, Virginia Tech, Blacksburg, VA 24061, USA

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

Keywords: The impact of growth rate (GR) and finishing regime (FR) on growth and meat quality traits of Angus x Nellore
Grain-fed crossbred steers, harvested at a constant body weight (530 ± 20 kg) or time on feed (140 days), was evaluated.
Grass-fed Treatments were: 1) feedlot, high GR; 2) feedlot, low GR; 3) pasture, high GR and 4) pasture, low GR. Live body
Feeding regime
composition, carcass and meat quality traits were evaluated. High GR had greater impact on muscle and fat
Meat aging
Muscle growth
deposition in feedlot-finished, but not in pasture-finished animals. Feedlot animals had higher Longissimus muscle
area, backfat thickness, meat luminosity and tenderness when compared to pasture groups. Moreover, pasture-
and feedlot-finished animals with similar GR did not differ in the chromatic attributes of non-aged meat,
regardless of endpoint. Thus, GR appeared to be the main factor driving beef chromatic parameters, while FR had
a major impact on achromatic attributes and tenderness of meat.

1. Introduction
Low input cattle feeding systems, such as pasture-based feeding or
other forage-based feeding approaches form the basis for producing
ruminant meat throughout the world. These systems result in the pro­
duction of older animals because growth rates are generally slower but
carcasses are leaner at harvest (Koch, Pavan, Andrae, & Duckett, 2018;
Koch, Pavan, Long, Andrae, & Duckett, 2019; Nian, Kerry, Prendiville, &
Allen, 2017). Conversely, high input or intensive meat animal produc­
tion through the use of cereal grains, such as the case for most feedlot
systems, increases growth rate and feed efficiency, and results in car­
casses of high quality by reducing age at harvest and increasing fat
deposition (Koch et al., 2018, 2019; Wicks et al., 2019).
In all finishing systems, growth rate, age at harvest and body weight
affect muscle characteristics and energy metabolism. These unique
features of muscle can drive postmortem metabolism and ultimately
consumer acceptability of the final product either through purchasing
decisions or positive eating experiences (Wicks et al., 2019). In this
sense, Gagaoua, Picard, Soulat, and Monteils (2018) reported that more
tender beef is associated with the decrease of age at harvest and forage
proportion, and to the increase of fatness degree and concentrate level.
Likewise, age at harvest, final weight and degree of fatness impact beef
color traits (Gagaoua, Picard, & Monteils, 2018). In general, beef cattle
finished in feedlot systems (cereal grain fed) is brighter red in color and
produces a more tender cooked product than those cattle finished in
extensive systems (pasture-based). In contrast, cattle produced in more
extensive-based systems produce fresh beef that is often darker in
appearance and less tender for consumption (Apaoblaza et al., 2020;
Torrecilhas et al., 2021; Wicks et al., 2019). However, finishing regime
(grain versus forage) and growth rate are often confounded because
pasture or high forage finishing diets energetically result in lower
growth rate, while cereal grain-based diets used in most feedlots support
a more rapid growth rate. Furthermore, finishing system and growth
rate have profound impacts on carcass characteristics and subsequently
meat quality development depending on which harvest endpoint strat­
egy is selected. Specifically, harvesting cattle at equivalent final body
weights (BW), days on feed or degree of fatness can result in quite
different end products because lean and fat tissue accretion is
confounded with animal age and final weight.
Several studies have evaluated the influence of the finishing strategy
on carcass characteristics and meat quality development, however dif­
ferences in growth rate (Apaoblaza et al., 2020; Koch et al., 2019), age at
harvest (Apaoblaza et al., 2020; Koch et al., 2018; Manni, Rinne,
Huuskonen, & Huhtanen, 2018), and final BW (Koch et al., 2019; Nian

* Corresponding author.
E-mail address: saulo.luz@usp.br (S.L. Silva).

https://doi.org/10.1016/j.meatsci.2021.108599
Received 29 December 2020; Received in revised form 1 June 2021; Accepted 4 June 2021
Available online 6 June 2021
0309-1740/© 2021 Elsevier Ltd. All rights reserved.
J.F. Morales Gómez et al. Meat Science 183 (2022) 108599

et al., 2017) across these studies complicate the issue (Manni et al., Table 1
2018). Therefore, the overall aim of this study was to understand more Dietary ingredients and chemical composition (DM basis) of finishing diets and
precisely how finishing cattle, namely growth rate and feeding regime supplements.
and harvest endpoints affect beef quality. Item Feedlota Pastureb Pasturec Supplementd

1 2 3
2. Material and methods
Ingredients, %
Corn silage 20.3 – – – – –
All experimental procedures involving animal care were conducted Ground corn grain 71.3 – – 49.5 74.8 76.3
in accordance with the Institutional Animal Care and Use Committee Soybean meal 45% 4.6 – – 40.0 20.0 20.0
Guidelines of the College of Animal Science and Food Engineering at Limestone 0.5 – – – – –
Potassium chloride 0.4
University of São Paulo (protocol approval number 13.1.541.74.0). – – – – –
Urea 1.4 – – 3.0 1.5 1.5
Mineral mixturee 1.5 – – 2.5 1.3 0.8
2.1. Animals and treatments Mineral salt – – – 5.0 2.5 1.5
Chemical
composition, %
Seventy-two contemporary ½ Angus x ½ Nellore crossbred animals
Dry matter (% as 62.7 24.3 38.4 98.0 98.5 98.2
were weaned (eight months-old) and surgically castrated (ten months- fed)
old). Approximately 45 d after castration, steers (330 ± 30 kg BW; 12 Crude protein 14.4 7.5 6.9 30.8 19.4 19.5
± 1 month-old) were subjected to a completely randomized experi­ Ash 4.2 8.4 8.0 – – –
mental design consisting of four treatments (n = 18 per treatment): 1) Neutral detergent 20.9 64.8 70.0 11.2 11.8 11.9
fiber
feedlot finished, high growth rate (F-H; average daily gain [ADG] pro­ Acid detergent fiber – 36.6 39.9 5.3 4.4 4.5
jected at 1.5 kg/d); 2) feedlot finished, low growth rate (F-L; ADG Ether extract 2.7 1.5 1.5 2.4 3.0 3.1
project at 0.9 kg/d); 3) pasture finished, high growth rate (P-H; ADG Non‑nitrogen 58.9 55.9 55.3 – – –
projected at 0.9 kg/d); and 4) pasture finished, low growth rate (P-L; extract
Total digestible 79.0 66.7 71.6 72.7
ADG projected at 0.6 kg/d). Additionally, half of the steers on each – –
nutrientsf, %
treatment were randomly assigned prior to study for harvest at BW of
a
530 kg. The other half were harvested after 140 d on feed, regardless of Feedlot diets were formulated using RLM software (Integra Software, Pira­
the feeding regime (Fig. 1). cicaba, São Paulo, Brazil) to meet the nutrient requirements of growing Angus x
Nellore crossbred steers at a rate of 1.5 kg of ADG as specified by the Cornell Net
Carbohydrate and Protein System (CNCPS; Fox et al., 1992).
2.2. Pasture system b
Chemical composition of a pool of 4 pasture samples collected in the rainy
season.
Thirty-six steers were pasture finished in 12 pens (three steers/pen; c
Chemical composition of a pool of 4 pasture samples collected in the dry
six pens/harvest endpoint criteria; three pens/gain rate) containing season.
Marandu grass (Brachiaria brizantha cv. Marandu), supplied with auto­ d
Chemical composition of supplements offered to pasture-fed animals during
matic waterers and feed bunks for ad libitum mineral supplementation. the dry season.
e
During the rainy season (September–May), cattle were supplemented Minerthal 160MD (Minerthal) calcium, 208 g; cobalt, 148 mg; copper 2664
with ad libitum access to salt (Minerthal 80, Minerthal). Growth rate was mg; sulfur, 64 g; fluorine (Max), 1600 mg; phosphor, 160 g; iodine, 141 g;
controlled by adjusting stocking densities in each pen every two weeks manganese, 2200 mg; selenium, 37 mg; zinc, 7992 mg; monensin sodium, 4000
mg.
using off-study cattle as diluters. During the dry season (May–Sep­ f
Estimated by the formula of Weiss et al. (1992).
tember), growth rate of steers harvested at a constant BW endpoint was
controlled every week by protein-energy supplementation (Table 1)
with the amount of supplement varying according to the pasture avail­ individual feed intakes. All feed bunks were covered and cattle had ad
ability, to ensure targeted growth rates. libitum access to water. Steers were provided a 31-d adaptation period to
the facilities and diet (Table 1). Adaptation period was not counted in
the time on feed for groups harvested on a days of feed endpoint.
2.3. Feedlot system
Experimental diets were formulated using RLM software (Integra Soft­
ware, Piracicaba, São Paulo, Brazil) to meet the nutrient requirements of
Thirty-six feedlot-fed steers were housed in four pens (n = 9 steers/
growing Angus x Nellore crossbred steers at a rate of 1.5 kg of ADG. To
pen; two pens/for each growth rate) equipped with electronic gates
attain the targeted growth rate of each treatment, the F-H treatment was
(American Calan Inc., Northwood, NH, USA), which allowed control of
fed ad libitum and the F-L treatment received 70% of the daily fed of the
F-H.

2.4. Growth traits and ultrasound measurements

On the first (beginning of trial) and last (before harvest) weighing

Fig. 1. Study design, animals in feedlot with high (F-H) and low (F-L) growth
rate, and animals in pasture with high (P-H) and low (P-L) growth rate harvest
at BW of 530 kg or after 140 d on feed.
days, cattle were subjected to a 16 h feed withdraw, while all other
weights were collected without fasting. Ultrasound measurements to
determine the Longissimus muscle area (LMAU) and backfat thickness
(BFTU) between the 12th and 13th ribs and the rump fat thickness
(RFTU) on the Biceps femoris muscle were taken at 1, 28, 56, 116, 134,
178, 214, 262 and 292 d of study, using an Aloka® equipment, model
SSD 500 Micrus (Aloka Co. Ltd., Zug, Switzerland), with a linear probe
3.5 mHz and 172 mm in length. Images were recorded and then inter­
preted using Lince® software (M&S Consultoria Agropecuária, Piras­
sununga, SP, Brazil).

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J.F. Morales Gómez et al. Meat Science 183 (2022) 108599

2.5. Harvest, carcass traits and meat quality
After reaching pre-established endpoints, steers were fasted for 16 h
then transported (1 km and 4 km for feedlot- and pasture-fed animals,
respectively) to the abattoir located at the University of São Paulo,
Pirassununga, SP, Brazil. To minimize day to day variation, personnel
and facility remained constant across the four days of harvest for those
cattle harvested at on a constant weight endpoint basis. Cattle harvested
at a constant time on feed were all harvested on the same day. Animals
were harvested according to humane procedures as required by Brazil­
ian law. Hot carcass weights (HCW) and dressing percentages (DP) were
determined immediately after evisceration. Afterward, kidney, pelvic
and inguinal fat (KPIF) were collected and weighed.
After a 24 h chill (0 to 2 ◦ C), carcasses were ribbed between 12th and
13th rib and an image of the Longissimus muscle surface was taken using
a digital camera coupled to stainless steel (15 × 20 cm) device and
posterior determination of Longissimus muscle area (LMAC) and backfat
thickness (BFTC) was completed using Lince® software. In addition,
samples were excised from the Longissimus thoracis (LT) muscle at the
12th and 13th rib level, immediately snap frozen in liquid nitrogen, and
stored at − 80 ◦ C for pH measurements. Three 2.5 cm thick LT samples
from each carcass were collected, vacuum packed and aged (0 to 2 ◦ C)
for 0, 7 and 14 d for further instrumental analyses of color (L*, a* and
b*) and Warner-Bratzler shear force (WBSF).
mm/min. The WBSF values were determined by averaging the
maximum peak force of six replicate cores per steak.
2.9. Statistical analysis
Data were analyzed in a completely randomized design using PROC
MIXED procedure of SAS 9.4 (SAS Institute Inc., Cary, NC) and each
harvest endpoint was analyzed separately. Treatment was included as
fixed, and pen and animal were used as experimental unit for pasture
and feedlot treatments, respectively. Plots of residuals and the W sta­
tistic (Shapiro and Wilk, 1965) were evaluated to determine normality
for all data, and outliers (> 3 and < − 3 standard deviation) were
excluded. The BW and ultrasound measurements data were analyzed as
repeated measurements over time, covariance structures were modeled
and the structure with the best fit for every trait was used. Color char­
acteristics and SF data were analyzed in a split-split plot design with
treatment (F-H, F-L, P-H and P-L) as whole plot and aging time (0, 7, and
14 d) as subplot. The least squares means (LSMEANS) statement was
used to calculate adjusted means for treatment and means were
compared by Student’s t-test. Differences were considered statistically
significant when P ≤ 0.05, and values ranging from 0.05 < P ≤ 0.10
were considered as trends.
3. Results

2.6. pH measurement
The pH samples were prepared using the iodoacetate method as
described by Bendall (1973). Briefly, four volumes of homogenate were
removed from the reaction vessel and placed in a new centrifuge tube
containing one volume of 25 mM sodium acetate and 750 mM KCL (pH
7.0). Samples were centrifuged at 13,000 rpm for 5 min at room tem­
perature, equilibrated to 25 ◦ C and measured directly using a portable
digital pH meter (Hanna Instruments - model HI99163, São Paulo,
Brazil).
2.7. Total intramuscular lipid
Fresh muscle samples (~50 g) were collected and homogenized in a
processor (Mixer Walita Model RI1364 with microprocessor, Phillips of
Brazil LTDA., Brazil). Approximately 3.0 g of the sample were weighed
in duplicate. Lipids were extracted by homogenizing the sample with a
chloroform:methanol:distilled water (2:1:0.8) solution. Sodium chloride
was added to 1.5% so that the lipids were isolated and determined by
gravimetry, according to the methodology proposed by Bligh and Dyer
(1959).
Actual ADG for each treatment was very close to those originally
targeted (Table 2). However, P-H steers harvested on a final BW
endpoint grew slower (0.15 kg/d) than originally planned. In groups
harvested after a constant day on feed endpoint, F-H and P-L steers, had
the highest and lowest (P < 0.001) final BW, respectively, while F-L and
P-H had intermediate final BW, with no difference between the two
treatments (Table 2).
3.1. Body weight and carcass traits evaluated by ultrasound
A treatment × time interaction (P < 0.001) was observed for BW in
both harvest endpoint criteria (Fig. 2A and B). No differences in the
initial BW among treatments were evident. The F-H steers were heavier
(P < 0.01) after 28 d on feed and this disparity continued to increase
with time on feed until harvest, regardless of endpoint. The P-H group
Table 2
Least square means, standard error mean (±) and probabilities (P-value) of
growth traits used to characterize the treatments in both harvest endpoint
criteria.
Traits Treatment P-value

2.8. Instrumental color and shear force F-H F-L P-H P-L

Harvest at 530 kg body weight

Samples were removed from the vacuum package and allowed to Feeding time, d 116 228 262 292 –
bloom for 30 min at 4–6 ◦ C (AMSA, 2012). Meat color was evaluated Initial body 349 ± 8.4 331 ± 8.4 335 ± 329 ± 0.446
weight, kg 11.9 11.9
using the CIE Lab system (CIE, 1986) with a portable spectrophotometer
Final body 534 ± 8.3 526 ± 8.3 535 ± 515 ± 0.558
model CM2500d (Konica Minolta Brazil, São Paulo, Brazil) with stan­ weight, kg 11.7 11.7
dard illuminant D65, observation angle of 10◦ and an aperture of 30 mm. Average daily 1.51a ± 0.94b ± 0.76c ± 0.62d ± <0.001
Steaks were roasted in an oven equipped with a thermostat adjusted to gain, kg/d 0.04 0.04 0.06 0.06
170 ◦ C (Model F130 / L – Electric Furnaces Golden Arrow Industry and Harvested at 140 d on feed
Commerce Ltda., São Paulo, Brazil). Internal meat temperatures were Feeding time, d 140 140 140 140 –
monitored using individual thermocouples. Once steaks reached an in­ Initial body 363 ± 338 ± 335 ± 335 ± 0.2634
weight, kg 9.81 9.81 14.55 14.55
ternal temperature of 40 ◦ C, they were turned and cooked to an internal
Final body 582a ± 456b ± 463b ± 419c ± <0.001
temperature of 71 ◦ C, as recommended by AMSA (2015). Cooked steaks weight, kg 13.5 13.5 20.4 20.4
were allowed to cool to 4 ◦ C for 12 h. Six 1.27 cm diameter core samples Average daily 1.58a ± 0.85b ± 0.91b ± 0.60c ± <0.001
were taken parallel to muscle fiber orientation using a Drill Brench gain, kg/d 0.05 0.05 0.08 0.08
(Model FG-13B, Caracol Trading of Machinery and Tools LTDA, São abcd
Values within a row with different superscripts differ significantly at P <
Paulo, Brazil). The WBSF was determined on cores using the TMS-PRO 0.05. F-H = feedlot-fed with high growth rate; F-L = feedlot-fed with low growth
texture analyzer (Food Technology Corporation, Sterling, Virginia, rate; P-H = pasture-fed with high growth rate; P-L = pasture-fed with low
USA) coupled with a Warner-Bratzler shear device set at a speed of 200 growth rate.

3
J.F. Morales Gómez et al. Meat Science 183 (2022) 108599

A. 650 B. 650 Treat P = 0.010

Time P < 0.001
600 600 Treat x time P < 0.001 *
* *
550 * *
, Body weight, kg

550 *

Body weight, kg
* *
500 500 *
* *
450 * 450

400 Treat P < 0.001 400

Time P < 0.001
350 Treat x time P < 0.001
350

300 300
0 20 40 60 80 100 120 140 160 180 200 220 240 260 280 300 0 20 40 60 80 100 120 140
Feeding time, d Feeding time, d

90
C. 90 D. Treat P = 0.010
85 * 85 Time P < 0.001
*
* *
80 * * 80 Treat x time P < 0.001
*
75 * 75
*

LMAU, cm2
LMAU, cm2

70 * 70
*
65 65
60 60
* *
55 Treat P = 0.004 55
50 Time P < 0.001 50
Treat x time P < 0.001
45 45
40 40
0 20 40 60 80 100 120 140 160 180 200 220 240 260 280 300 0 20 40 60 80 100 120 140
Feeding time, d Feeding time, d
F-H F-L P-H P-L F-H F-L P-H P-L

Fig. 2. Least square means and standard error (ɪ) of body weight and Longissimus muscle area (LMAU) evaluated by ultrasound according to feeding time and
treatments (Treat; F-H = feedlot-fed with high growth rate; F-L = feedlot-fed with low growth rate; P-H = pasture-fed with high growth rate; P-L = pasture-fed with
low growth rate. Figs. A and C represent animals harvest at 530 kg body weight and Figs. B and D represent animals harvested at 140 d on feed.

was heavier (P < 0.05) than F-L at 86 d when harvested in a weight basis,
but no difference in BW was found when steers were harvested on days
on feed. The P-H steers were heavier than P-L steers after 56 d and 134 in
those cattle harvested based on days on feed, and those on a final BW
basis, respectively. There was a treatment × time interaction (P < 0.001)
for LMAU in both harvest endpoints (Fig. 2C and D). Animals finished in
the feedlot (F-H and F-L) had larger (P < 0.05) LMAU at all times when
compared to pasture fed cattle (P-H and P-L) regardless of harvest
endpoint. However, on day 214 F-L and P-H were similar when har­
vested on a constant weight endpoint. The F-H animals had larger LMAU
when compared to F-L animals after 56 d and 86 d of feeding when
harvested at a constant weight and time on feed endpoint, respectively
(P < 0.05). Animals finished on pasture (P-H and P-L) had similar LMAU
at all times regardless of harvest endpoint.
A treatment × time interaction was observed for BFTU (Fig. 3A and
B) and RFTU (Fig. 3C and D) for both endpoints (P < 0.001). Feedlot-fed
steers (F-H and F-L) had greater BFTU (P < 0.05) and RFTU (P < 0.05) on
the first weigh day compared to pasture-fed steers (P-H and P-L).
Moreover, F-H steers had greater BFTU and RFTU compared to pasture-
fed steers (P-H and P-L; P < 0.001) and F-L steers (P < 0.05) after 1 and
28 d of feed, respectively, regardless of endpoint targets. The F-L steers
had greater BFTU and RFTU than P-H (P < 0.05) and P-L (P < 0.05) after
28 d on feed when harvested at a constant weight. However, when
harvested on based on days on feed, F-L steers had greater BFTU (P <
0.05) and RFTU (P < 0.05) than P-H and P-L from the beginning up to 86
and 56 d on feed, respectively. Furthermore, P-H steers had greater BFTU
and RFTU than P-L, after 178 d on feed in cattle harvested on a constant
weight endpoint (P < 0.05), but there was no difference in fat deposition
when steers were harvested at constant days on feed.
3.2. Carcass traits evaluated at slaughter
When steers were harvested on a final BW basis, feedlot-fed steers
had greater KPIF (P < 0.001) than those pasture-fed (Table 3). BFTC (P
= 0.085) tended to differ between treatments, in which F-H animals had
greater BFTC when compared to pasture-fed animals. The F-H steers had
greater total intramuscular lipid (P = 0.044) than pasture-fed , and F-L
group was greater than P-L, but did not differ from P-H. However, HCW,
DP, LMAC and pH24h were not impacted by any treatment.
When steers were harvested based on time on feed, F-H had the
heaviest HCW (P < 0.001) compared to pasture-fed steers, while P-H did
not differ with F-L and P-L. Moreover, feedlot-fed steers had higher DP
(P = 0.011) than pasture-fed cattle. The F-H steers had the greatest
LMAC (P < 0.001), BFTC (P < 0.001) and KPI fat (P < 0 .001) than all
other treatments, while P-H and P-L did not differ. Also, F-H steers had
lower pH24h (P = 0.039) and greater total intramuscular lipids (P =
0.008) than P-H and P-L but did not differ from each other.
3.3. Meat color
A treatment × aging interaction was observed for L*, a*, b* and
Chroma (C*) data regardless of harvest endpoint (Fig. 4; P < 0.05).
When steers were harvested on a constant BW, steaks from F-H steers
had highest L* values than all others (P < 0.05). Non-aged and 7-d aged
steaks from F-L steers had greater L* values than those of P-H and P-L
cattle (P < 0.05), while the latter two treatments did not differ. Non-

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J.F. Morales Gómez et al. Meat Science 183 (2022) 108599

12 Treat P < 0.001 12

A. Time P < 0.001
B. Treat P < 0.001
Time P < 0.001
10
Treat x time P < 0.001
Treat x time P < 0.001
*
10
*
*
8 8
BFTU, mm

BFTU, mm
* * *
6 6
* *
4 * 4 *
*
2 * *
2
*
0 0
0 20 40 60 80 100 120 140 160 180 200 220 240 260 280 300 0 20 40 60 80 100 120 140
Feeding time, d Feeding time, d

Treat P<0.001
C. Time P<0.001 D. Treat P < 0.001
Treat x time P<0.001 Time P < 0.001
10 10 Treat x time P < 0.001
*
*
8 * * 8
* *
* *
RFTU, mm

RFTU, mm
6 6

* * *
4 * 4

* *
2 2

0 0
0 20 40 60 80 100 120 140 160 180 200 220 240 260 280 300 0 20 40 60 80 100 120 140
Feeding time, d Feeding time, d
F-H F-L P-H P-L F-H F-L P-H P-L

Fig. 3. Least square means and standard error (ɪ) of backfat thickness (BFTU) and rump fat thickness (RFTU) evaluated by ultrasound according to feeding time and
treatments (Treat; F-H = feedlot-fed with high growth rate; F-L = feedlot-fed with low growth rate; P-H = pasture-fed with high growth rate; P-L = pasture-fed with
low growth rate). Figs. A and C represent animals harvest at 530 kg body weight and Figs. B and D represent animals harvested at 140 d on feed.

aged steaks from F-H steers had the highest a*, b* and C* values than all
other treatments (P < 0.05). However, non-aged steaks from F-L steers
did not differ for a*, b* and C* values when compared from those P-H
and P-L cattle. No difference was observed for a* and C* values when
steaks from F-H steers were compared to other treatments aged for 7 and
14 d, respectively. Beef from F-H cattle had higher b* values than P-H (P
< 0.001) and P-L (P < 0.05), while those from F-L cattle had greater b*
values than P-H steers (P < 0.05), when aged for 7 and 14 d.
When steers were harvested on a constant feeding duration, non-
aged and 14 d aged steaks from feedlot-fed steers had higher L* values
than those from pasture-fed cattle, while 7-d aged steaks from F-L steers
had similar L* values compared to other treatments. Steaks from F-H
cattle had higher a*, b* and C* values than those from P-H (P < 0.05)
and P-L (P < 0.05) cattle in all aging periods. Also, 7 and 14 d aged
steaks from F-H steers had higher a* (P < 0.001), b* (P < 0.05) and C* (P
< 0.05) values than those from F-L cattle. Moreover, steaks from F-L and
P-H steers did not differ in a*, b* and C* values at any aging time. Non-
aged and 7-d aged steaks from P-H, and P-L did not differ for a*, b* and
C* values. However, steaks from P-H cattle aged for 14 d had higher a*,
b* and C* values (P < 0.05) than those from P-L.
3.4. Shear force
A treatment × aging interaction was observed for WBSF of LT steaks
using both endpoint criteria (Fig. 5; P < 0.05). When steers were har­
vested on a final weight basis, non-aged steaks from feedlot-fed steers
had lower WBSF values than those from pasture-fed cattle (P < 0.05).
The 7-d aged steaks from F-H steers had lower WBSF than those from P-
H (P < 0.05) and P-L (P < 0.001) cattle, but there were no differences
between those of F-L and P-H cattle. Steaks aged for 14 d did not differ
with treatment. On the other hand, when steers were harvested on days
on feed endpoint, non-aged meat from F-H steers had lower WBSF than
those from P-H (P = 0.009) and P-L (P = 0.025) cattle, while the two did
not differ. Non-aged steaks from P-H steers had higher WBSF values than
those from F-L cattle (P = 0.046). In addition, 7 and 14 d aged steaks
from feedlot-fed steers had lower WBSF values than those from pasture-
fed cattle (P < 0.001).
4. Discussion
4.1. Growth and carcass traits
Growth rate had a greater impact on overall growth, muscle and fat
in feedlot cattle than pasture-fed steers. Greater BW, LMAU, BFTU, and
RFTU were observed in F-H steers after 28-d on feed compared to F-L;
differences that continue to increase with time to harvest, regardless of
the endpoint criteria. Likewise, high growth rate of feedlot-fed steers
harvested based on feeding duration increased LMAC KPI fat and BFTC.
However, differences between P-H and P-L were less evident and not
consistent throughout the study. Growth rate of pasture-fed steers did
not impact carcass traits (HCW, DP, KPIF, LMAC, BFTC and pH24h)
regardless of endpoint.
When comparing F-H and P-L animals, a higher in LMAU, BFTU,
RFTU, LMAC, BFTC, KPI fat and DP was observed in F-H steers, mainly
when harvested based on feeding duration. According to Koch et al.
(2018), feedlot-fed animals have greater growth rates and are harvested
younger, regardless of the final BW. Moreover, Frylinck, Strydom,
Webb, and du Toit (2013) reported that pasture-fed steers have lower
carcass weights than those fed in feedlots, principally due to the fact that
low growth rate results from pasture finishing systems. In addition,

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J.F. Morales Gómez et al. Meat Science 183 (2022) 108599

Table 3 (Ponnampalam, Hopkins, Bruce, Li, & Baldi, 2017).

Least square means, standard error mean (±) and probabilities (P-value) of the In the present study, P-L steers harvested on a constant feeding
effect of treatment on carcass traits in both harvest endpoint criteria. duration produced beef with a high pH24h, which can be considered as
Traits Treatments P-value dark-cutting or dark, firm and dry (DFD) meat (pH > 6.0; Ponnampalam
F-H F-L P-H P-L
et al., 2017). According to Ponnampalam et al. (2017), consumers tend
to reject dark-cutting meat as it is perceived to be from old or poorly-
Harvest at 530 kg body weight
handled animals and is described as being less tender, possessing an
Hot carcass weight, 303 ± 294 ± 298 ± 282 ± 0.172
kg 5.3 5.3 7.5 7.5 undesirable flavor, and having a short shelf-life. Dark-cutting meat is
Dressing, % 56.7 ± 56.0 ± 55.8 ± 54.6 ± 0.274 generally associated with a lack of glycogen, oxidative muscle meta­
0.59 0.59 0.84 0.84 bolism and(or) insufficient carcass fat (Ponnampalam et al., 2017; S. Wu
Kidney, pelvic and 16.3a ± 14.4a ± 9.3b ± 8.7b ± <0.001 et al., 2020). To that end, development of dark-cutting beef in this study
inguinal fat, kg 0.92 0.92 1.30 1.30
Longissimus muscle 81.6 ± 74.1 ± 80.4 ± 74.5 ± 0.185
from P-L steers, especially those harvested on constant days on feed, may
area, cm2 2.56 2.56 3.63 3.63 be related to insufficient carcass fatness or an increase in oxidative
Backfat thickness, 6.4a ± 5.1ab ± 3.2b ± 3.2b ± 0.085 muscle metabolism, because they were not stressed, at least from an
mm 0.79 0.79 1.12 1.12 overexertion standpoint. Further, once the glycogen levels in the muscle
pH24h 5.76 ± 5.83 ± 5.86 ± 5.76 ± 0.693
of the P-L group reached 46.9 μmol/g, it was well within the range
0.06 0.06 0.08 0.08
Total intramuscular 6.06a ± 4.95ab ± 3.14bc ± 2.27c ± 0.044 (45–55 μmol/g; Ferguson & Gerrard, 2014) necessary for muscle to
lipids, % 0.76 0.76 1.07 1.07 reach a normal beef pH. Other authors have reported a similar scenario
where ultimate muscle pH differed with finishing regime (Frylinck et al.,
Harvested at 140 d on feed
Hot carcass weight, 327a ± 258b ± 245bc ± 225c ± <0.001 2013; Ponnampalam et al., 2017). However, Apaoblaza et al. (2020) and
kg 9.2 8.5 12.9 12.9 Nian et al. (2017) failed to detect such difference for ultimate pH.
Dressing, % 56.2a ± 56.4a ± 53.1b ± 53.4b ± 0.011
0.62 0.58 0.89 0.89
4.2. Meat color
Kidney, pelvic and 19.0a ± 9.9b ± 3.8c ± 3.3c ± <0.001
inguinal fat, kg 1.18 1.18 1.67 1.67
Longissimus muscle 83.8a ± 75.1b ± 62.0c ± 58.4c ± <0.001 Non-aged meat from feedlot-fed steers had higher L* values than
area, cm2 2.91 2.70 4.12 4.12 those from pasture-fed regardless of growth rate, which suggests that
Backfat thickness, 8.5a ± 3.6b ± 2.1bc ± 1.4c ± <0.001 beef luminosity was greatly affected by feeding regime, although exac­
mm 0.76 0.70 1.07 1.07
pH24h 5.59c ± 5.72bc ± 5.89ab ± 6.04a ± 0.039
erbated by increases in growth rate. In contrast, Purchas et al. (2002)
0.08 0.08 0.12 0.12 found no difference in L* values between high and low growth rate but
Total intramuscular 4,24a ± 3.26b ± 1.92c ± 1.90c ± 0.008 this was in predominately high forage feeding regimes. Luminosity
lipids, % 0.40 0.37 0.56 0.56 changes have been attributed to three main factors: overall muscle
abc
Values within a row with different superscripts differ significantly at P < metabolism, intramuscular fat and muscle fiber characteristics (Hughes,
0.05. F-H = feedlot-fed with high growth rate; F-L = feedlot-fed with low growth Clarke, Purslow, & Warner, 2020; Ramanathan, Suman, & Faustman,
rate; P-H = pasture-fed with high growth rate; P-L = pasture-fed with low 2020). Pasture-fed steers have been reported to have a more oxidative
growth. metabolism and lower L* values, resulting in darker meat when
compared with feedlot-fed steers (Apaoblaza et al., 2020). Others argue
McIntyre et al. (2009) observed that feedlot-fed animals experiencing a that high L* values in steaks from feedlot fed cattle results from higher
high growth rate (1.22 kg/d ADG) had higher BFT and greater marbling marbling scores and lipid content of the muscle (Mancini & Hunt, 2005).
scores than pasture-fed animals with a low growth rate (0.67 kg/d ADG), Indeed, greater intramuscular fat deposition alters the muscle structure
concluding that growth rate has a significant effect on carcass charac­ (Valenzuela, Lloyd, Mastaglia, & Dawkins, 2020), which is a major
teristics. However, in the present study, although F-L and P-H cattle had factor determining the achromatic attributes of the lean color (Hughes
similar ADG, F-L steers had greater muscle (LMAU) and fat depositions et al., 2020). The fact that feedlot cattle in this study possessed greater
(BFTU, RFTU and KPI fat), which indicated that the deposition of muscle intramuscular fat than pasture-fed supports the latter construct.
and fat is mainly determined by the finishing system, and to a lesser Regardless of harvest endpoint, non-aged meat from F-H steers had
extent by growth rate, despite the fact the effects of these factors on higher a* and b* values than those of P-H and P-L treatments, but
carcass characteristics may be accumulative. interestingly, F-L and P-H were not different. These results may suggest
The effect of finishing system observed in this work may be related to that different growth rates across finishing systems may be the main
the fact that feedlot-fed animals were subjected to a high-density energy factor responsible for altering beef chromatic attributes. Davis, Cole,
diet, which may have provided a greater substrate availability leading to Backus, Melton, and Al (1981) reported lighter beef in animals with ad
an increase of propionate in the rumen and therefore glucose produc­ libitum access to feed when compared with restricted fed or pasture fed.
tion, which are related to the muscle and fat deposition (Ladeira et al., The same authors also reported no difference for beef color between
2016; Wicks et al., 2019). In contrast, nutrient intakes are often below pasture- and feedlot-fed animals with similar ADG, consistent with our
animal requirements for maximal growth in pasture system, which af­ observations. Growth rate directly impacts muscle metabolism, where
fects animal growth and lean tissue and fat deposition (Mwangi et al., the lower the growth rate, the more oxidative and darker the muscle is
2019; Wicks et al., 2019). Pasture finished cattle harvested at the same (Picard & Gagaoua, 2020). Apaoblaza et al. (2020) reported that
weight endpoint had a lower BW between 116 and 134 d on feed, which pasture-fed steers (lower growth rate) had a more oxidative muscle
most likely led to losses in BTF and RFT, or a failure to deposit fat and metabolism and greater myoglobin content than feedlot-fed (higher
muscle (LMAU) during this period, which comprised the most critical growth rate) and darker beef. Even though pasture-fed produced darker
time of the dry season of the year in tropical countries (May to beef than feedlot-fed animals, C* values indicated that all the steaks may
September). During this period, there is a shortage of rain and reduced be considered bright red (C* > 20) and would not cause consumer
sunlight, which decreases the amount and nutritional quality of the rejection (MacDougall, 1982).
pasture and, consequently, animals lose BW and body fat as a conse­ Although different growth rates across finishing systems affected
quence of mobilizing reserves for maintenance. As a result of the beef chromatic traits, different growth rates in pasture, regardless of
nutritional deficiency, pasture-fed steers produce leaner carcasses, endpoint criteria, and in feedlot when harvested on days on feed basis,
which may affect the pH and ultimate quality of beef produced did not affect non-aged beef color. Beef from F-H steers had higher L*, a*
and b* values than F-L steers. Therefore, the increased time (112 d)

6
J.F. Morales Gómez et al. Meat Science 183 (2022) 108599

Fig. 4. Least square means, standard error (ɪ) and probabilities of meat color traits according to aging and treatments (Treat; F-H = feedlot-fed with high growth rate;
F-L = feedlot-fed with low growth rate; P-H = pasture-fed with high growth rate; P-L = pasture-fed with low growth rate). Fig. A, C, E and G represents animals
harvest at 530 kg body weight and Figs. B, D, F and H represents animals harvested at 140 d on feed. Chroma (C*) values were calculated according to AMSA (2012),
where C* = (a*2 + b*2)0.5. abc Values within each time with different superscripts differ significantly at P < 0.05.

7
J.F. Morales Gómez et al. Meat Science 183 (2022) 108599

A.
Treat P = 0.003
100 Aging P < 0.001
a Treat * Aging P = 0.020
90
a
80

Shear force, N
70 b
b a
60 ab
bc
50 a
c a
40 a a

30

20
0 7 14
Aging, d

B.
100 Treat P < 0.001
a Aging P < 0.001
90 ab a
a Treat * Aging P = 0.025
80 bc
c a a
Shear force, N

70

60

50

40 b b b
b
30

20
0 7 14
Aging, d

F-H F-L P-H P-L

Fig. 5. Least square means, standard error (ɪ) and probabilities of shear force according to aging and treatments (Treat; F-H = feedlot-fed with high growth rate; F-L
= feedlot-fed with low growth rate; P-H = pasture-fed with high growth rate; P-L = pasture-fed with low growth rate). Fig. A represents animals harvest at 530 kg
body weight and Figs. B represents animals harvested at 140 d on feed. abc Values within each time with different superscripts differ significantly at P < 0.05.

required to attain an equivalent weight to F-H cattle affected the beef
color from F-L steers. While the exact reason for this is not readily
evident, this may be related to changes in muscle structure and meta­
bolism, as previously mentioned, or it may be due to increased meat
pigments due to an increase in chronological age, which has been shown
to result in darker beef (Matarneh, Silva, & Gerrard, 2020). On the other
hand, beef color observed in P-H and P-L steers harvested on the same
feeding duration may be somehow related to the biochemistry known to
produce DFD meat. In general, DFD meat has greater water holding
capacity in addition to changes in muscle structure, which reduces light
scattering and leads to a dark appearance (Hughes et al., 2020; Hughes,
Clarke, Purslow, & Warner, 2017). However, Apaoblaza et al. (2020)
argued that dark beef from pasture-fed steers is not recapitulating the
biochemistry undergirding traditional DFD because this is normally
predicated on animal stress, again, not an issue with cattle in this study,
based on residual muscle glycogen concentration.
Besides the antemortem factors of growth rate and feeding regime, the
postmortem aging factor affected beef color development. Beef color
development and stability are vulnerable to changes in muscle tissue
characteristics and metabolism, such as pH, muscle fiber type, antioxi­
dants, metmyoglobin reducing activity, lipid oxidation, oxygen con­
sumption and mitochondrial activity (Ramanathan & Mancini, 2018). In
the present study, increased aging resulted in lighter beef across all
treatments regardless of harvesting endpoint. However, the aging effect
on a* and b* did not behave linearly in all treatments, whose behavior
was observed to be different in non-aged meat than that observed in 7
and 14 d aged meat. According to Ramanathan and Mancini (2018),
aging increases the color development and decreases the color stability
of the meat, as it is mainly related to less metmyoglobin reducing ac­
tivity and less oxygen consumption. Several studies have reported an
increase in the beef luminosity and redness with an increase in aging
period, in addition to relating the development and color stability with
the metabolism of each muscle (Nair, Li, Beach, Rentfrow, & Suman,
2018; Ramanathan & Mancini, 2018).
4.3. Shear force
Feedlot-fed steers produced more tender non-aged beef than pasture-
fed ones, when harvest on a BW basis, regardless of growth rate. How­
ever, WBSF was similar between feedlot- (F-H vs. F-L) and pasture-fed
(P-H vs. P-L) steers differing in growth rates, regardless of harvest
endpoint. Manni et al. (2018) also reported no difference for WBSF
between feedlot finished animals with different growth rates. Therefore,
growth rate, as used in this study, does not seem to be the main factor

8
J.F. Morales Gómez et al.
responsible for differences in meat tenderness between feedlot- and
pasture-fed animals, which is in a good agreement with the results re­
ported by Brown et al. (2007), who found more tender beef in feedlot-
fed steers in comparison with pasture-fed steers, regardless of the
growth rate. Blank, Russell, Lonergan, and Hansen (2017) reported that
dietary fiber content may affect meat tenderness when observing that
steers fed more fibrous diets showed greater variation in the extent of
protein degradation and produced less tender beef.
Despite the lower dietary fiber content, feedlot-fed animals are
subjected to a high-density energy diet, which positively impacts the
growth rate and has been related to the shift in muscle fiber type towards
a more glycolytic metabolism and lower frequency of DFD meats, as well
as presenting a more active lipid metabolism that leads to the increase of
subcutaneous and intramuscular fat (Frylinck et al., 2013; Mwangi et al.,
2019; Wicks et al., 2019). Muscle with a more glycolytic metabolism
(Dang, Buhler, Thornton, Legako, & Matarneh, 2020; Joo, Kim, Hwang,
& Ryu, 2013), greater intramuscular fat (Jung, Hwang, & Joo, 2016;
Valenzuela et al., 2020), and a normal ultimate pH (Wicks et al., 2019)
generally have tender meat. In the present study, feedlot-fed animals,
especially F-H, showed higher subcutaneous and intramuscular fat and a
normal ultimate pH, which supports the lower WBSF values in non-aged
meat of these animals when compared to pasture-fed animals.
Despite the non-aged meat from pasture-fed steers have had higher
WBSF values than those from feedlot-fed steers, there was no difference
between feedlot- and pasture-fed steers harvested at the same ending
weight after 14-d aging similar to that shown by others (Koch et al.,
2019; Nian et al., 2017). However, when cattle were harvested on the
same days of feeding, 14 d of aging was not sufficient to tenderize the
meat from lean pasture-fed steers. Therefore, it can be suggested that
age-related tenderization from pasture- and feedlot-fed steers may be
dependent of endpoint targets. Moreover, the difference observed for
WBSF between feedlot- and pasture-fed steers harvested on feeding
duration may be related to the pH24h, which averaged 5.62 and 5.97,
respectively. Lomiwes, Farouk, Wu, and Young (2014) and Wu, Farouk,
Clerens, and Rosenvold (2014) concluded that ultimate meat 5.8 < pH >
6.2 promoted a faster rate of structural protein degradation, while ul­
timate meat pH between 5.8 and 6.2 promoted the slowest rate of
degradation and, consequently, higher WBSF values. Wu et al. (2014)
also stated that at least 21 d aging was required to mitigate the differ­
ences in WBSF when comparing pH < 5.8 and 5.8 < pH <6.2. Together,
these data argue that feeding strategy can affect the aging needed to
ensure optimal beef quality, though the mechanism underlying this
process requires additional work.
5. Conclusion
Both finishing regime/strategy/system and growth rate impacted the
carcass traits, muscle and fat deposition, and meat quality attributes.
Growth rate appeared to be the main factor responsible for variations in
the beef chromatic attributes, while finishing strategy had a major
impact on differences in beef achromatic attributes and eating quality.
Moreover, 14 d of aging mitigated differences in meat tenderness be­
tween finishing systems when pasture-fed steers were harvested on a
final weight endpoint, regardless of growth rate. Differing endpoint
targets of feeding beef cattle can change these quality-based outcomes.
Declaration of Competing Interest
The authors declare no conflict of interest associated with this
research.
Acknowledgements
This work is supported by the Foundation for Research Support of the
State of São Paulo (FAPESP) [grant # 2018/01434-8; 2018/25320-1]
and the National Council for Scientific and Technological Development
9
Meat Science 183 (2022) 108599
(CNPq) [grant # 430664/2018-4]. This work is supported in part by
A1364 Novel Foods and Innovative Manufacturing Technology [grant
no. 2020-67017-31269] from the USDA National Institute of Food and
Agriculture. Authors would like to thank all the support provided by
Prof. Dr. Valdo Rodrigues Herling, Fernando José Schalch Júnior and
Minerthal®.
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