Animal 15 (2021) 100381

Contents lists available at ScienceDirect

Animal
The international journal of animal biosciences

Growth curve, blood parameters and carcass traits of grass-fed Angus

steers
J.A. Carrillo a, Y. Bai b, Y. He c, Y. Li d, W. Cai a, D.M. Bickhart e, G. Liu e, S.M. Barao f, T. Sonstegard g, J. Song a,⇑
a
Department of Animal & Avian Sciences, University of Maryland, College Park, MD 20742, USA
b
College of Life Sciences and Food Engineering, Hebei University of Engineering, Handan 056021, PR China
c
Department of Human Nutrition, Food and Animal Sciences, University of Hawai’i at Manoa, Honolulu, HI 96822, USA
d
College of Animal Science and Technology, South China Agricultural University, Guangzhou 510642, PR China
e
Animal Genomics and Improvement Laboratory, BARC, Agricultural Research Service, USDA, Beltsville, MD 20705, USA
f
Hedgeapple Farm & Market, 3735 Buckeystown Pike, Buckeystown, MD 21717, USA
g
Recombinetics, 3388 Mike Collins Drive, Eagan, MN 55121, USA

a r t i c l e i n f o a b s t r a c t

Article history: The increasing demand for natural products is currently transforming the meat industry, making grass-
Received 13 May 2021 fed and finished beef a valuable option for improving profits. However, the transformation of conven-
Revised 5 September 2021 tional operations to grass-fed systems comprises many modifications, such as logistical, technological,
Accepted 7 September 2021
and financial that could be very complex and expensive, involving economic risk. Therefore, in this study,
Available online 29 October 2021
we analyzed the growth curve, critical economic traits, and carcass quality and finished characteristics
over several consecutive years in closely related grass-fed and finished Angus steers, to reduce the
Keywords:
genetic effect on the results. We found that grass-fed steers require around 188 additional days to reach
Animal welfare
Beef quality
the market weight (approx. 470 kg) and had approximately 70% less average daily gain compared to the
Genomics grain-fed and finished steers. Regression analysis demonstrated an interaction between feed and age
Grass-fed and finished beef (P < 0.01); thus, individual regressions were fitted for each regimen style, obtaining almost perfect linear
Growth development curves for both treatments, which could be straightforwardly used in practical situations due to its sim-
plicity. Six of eight carcass traits were different between grain-fed and grass-fed and finished steers. Hot-
carcass weight, dressing, back fat, and quality grade were superior in grain-fed individuals, contrarily to
yield grade and ribeye area/carcass ratio, which were better in grass-fed and finished steers (P < 0.05).
Interestingly, the meat tenderness was certainly low and similar in both treatments (P = 0.25), indicat-
ing the feasibility of producing tender meat with animals under a grass-fed diet. Nevertheless, according
to the quality grade analysis, grain-fed carcasses were greater ranked compared to grass-fed bodies
(P < 0.01), regardless of their same tenderness. The results will provide valuable information for better
understanding beef cattle in grass-feeding finishing systems, especially from weaning to harvest.
Additionally, the study will expand the knowledge about the quality of meat obtained from animals that
received grass exclusively, becoming relevant information for economic evaluation and management
decisions for grass-based cattle operations.
Ó 2021 University of Maryland. Published by Elsevier B.V. on behalf of The Animal Consortium. This is an
open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).

Implications omega3/omega6 ratio than grain-fed animals, which could poten-

For a long time, there are many arguments about the merits of
organic or natural food. The public often perceives that grain fin-
ishing systems seem unnatural compared to a grass-finishing ani-
mal. The results in this paper demonstrate the merits of beef
quality and animal welfare in a grass-based system style. The
grass-fed Angus steers produce beef with lower total fat and higher
tially benefit human health. Through this study, we will have a bet-
ter understanding of the merits of grass-fed and finished beef,
which will be very important to beef producers, human health,
and the environment.
Introduction

Beef represents a primary source of nutrients in many parts of

⇑ Corresponding author. the world and remains a highly desirable meal, especially in the
E-mail address: songj88@umd.edu (J. Song). U.S. and other western countries. The muscle contains not only

https://doi.org/10.1016/j.animal.2021.100381
1751-7311/Ó 2021 University of Maryland. Published by Elsevier B.V. on behalf of The Animal Consortium.
This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).
J.A. Carrillo, Y. Bai, Y. He et al. Animal 15 (2021) 100381

protein but also provides energy, minerals, and vitamins. Although homogeneous progeny (Lingle et al., 2001). The heifers were trea-
the importance of beef in the human diet is undeniable, its impact ted and raised equally until weaning. After weaning, the heifers
on human health is still controversial. Red meat consumption has were randomly assigned to either grass or grain-fed style. The
been associated with several pathologies such as cancer, coronary Grass-fed steers feeding regimen contained four periods. In the
heart disease, and diabetes (Pan et al., 2013; Xue et al., 2014; period of 50 days of postweaning, they were housed in a pasture
Consolo et al., 2018). According to some studies, the lipid profile with free choice orchard grass and clover. Then, they were offered
and antioxidant contents of the meat are correlated with several high moisture alfalfa haylage for 5–6 months. In the third period,
observed outcomes (Li et al., 2005; Moholisa et al., 2017; they continued to graze orchard grass and clover for 3 months. In
Carabante et al., 2018). Additionally, the concern about the content the fourth period, they were grazed on sudex (Sorghum Sudan-
of toxic metals (zinc, chromium, and nickel) in cow meat was grass) and then switched back to grazing on orchard grass and clo-
raised as a cause of the anthropogenic contamination (Ihedioha ver to finish weights and had access to high moisture hay (Alfalfa/
et al., 2014; Hwang and Joo, 2017). Contrarily, red meat consump- orchard grass) consuming about one bale per week. Bales’ size was
tion could be beneficial, providing high-quality protein that 121.96 cm tall by 182.88 cm wide. The grain-fed steers were fed a
increases satiety and boosts stamina (Campo et al., 2006; diet consisting of corn silage and whole shelled corn, soybean
Johnson et al., 2016; Frank et al., 2017). meal, and minerals in a Total Mixed Ration from weaning to date.
Currently, the beef market is increasingly changing toward They consumed between 9 and 11 kg of DM per head per day. The
grass-fed animals (Zhao et al., 2015; Carrillo et al., 2016; Kerth average seasonal yield for cool-season grasses was around 1.25–
and Rowe, 2016). This adjustment has been generated by con- 2.84–7 tons of DM per hectare.
sumer’s perception that animals raised on pastures and maintained The feed samples were sent to Dairy One Forage Laboratory
in more natural environments have a less detrimental impact on (Ithaca, NY) for nutrient analysis (Table 1) using wet chemistry.
the environment and yield healthier products (Campo et al., Crude protein was determined via combustion in a Leco CN682
2006). Rationally, it is coherent; grazing animals stay on pastures, Carbon/Nitrogen Determinator (Leco Corp., St. Joseph, MI) to obtain
consuming their food ad libitum, and moving freely, exercising nitrogen contents, then multiplying by 6.25 (Feldsine et al., 2002).
their muscles (Pimentel et al., 1980; Zhang et al., 2010). Differently, The acid detergent fiber was determined by acid detergent diges-
feedlot animals received a high-energy diet while they are con- tion in an Ankom A200 Digestion Unit (Ankom Technology, Mace-
fined to restricted spaces, preventing them from moving quickly don, NY). The neutral detergent fiber was determined by digestion
for more efficient feed conversion. The latest condition has been in a neutral detergent solution with the addition of a-amylase and
considered the traditional way to produce beef in the U.S. since sodium sulfite (Van Soest et al., 1991). Crude fat (or ether extract)
the 1950s. This system allows creating a large number of animals was obtained via extraction with diethyl ether in a Soxtec HT6 Sys-
in small areas, making the process more efficient by reducing costs tem (Foss North America, Eden Prairie, MN) (Feldsine et al., 2002).
such as housing and transportation. Thus, customers have habitu- Non-fiber carbohydrates (NFCs) were calculated by difference
ated to consume feedlot beef, which has specific characteristics [NFC = 100  (% NDF + % CP + % ether extract + % ash)]. Starch
that are difficult to achieve under other production systems was determined by first extracting sugar via water bath incubation
(Roosevelt, 2006). Nevertheless, public consciousness about health, at 40 °C followed by filtration, hydrolyzing starch to dextrose with
environment, and animal welfare are inducing farmers to adjust to glucoamylase enzyme, and analyzing dextrose concentration via
more extensive operations. Accordingly, the meat industry is pro- YSI 2700 Biochemistry Analyzer (YSI Inc. Life Sciences, Yellow
gressively experiencing an expansion of grass-based beef produc- Springs, OH). Net energy values are predicted by Dairy NRC 2001
tion to meet the growing demand (Swan and Boles, 2006; software.
Kenneth Mathews, 2013).
In this study, we hypothesize that diet styles, especially grass-
fed and finished and grain-fed and finished, could result in growth Growth curve
rate, physiological and quality variations of beef. This study aimed
to determine the real divergences between grass-fed and finished We randomly selected 92 Angus heifers from 2011 to 2014 to
and grain-fed and finished animals. Therefore, firstly, we ascer- analyze growth rates. About 20–25 Angus heifers were collected
tained the growth curve and compared several physiological traits every year. The grass-fed and grain-fed clusters consisted of 40
in both groups. Secondly, we explored the beef quality traits to
demonstrate whether a real difference exists between these two
types of meats. Ultimately, our result provides scientific evidence Table 1
Cattle feed component analysis.
for economic assessments and management decisions for beef cat-
tle production system transformation or modifications. Components DM basis (%)
Grain-fed Grass-fed
CP 13.9 24.3
Material and methods Adjusted CP 13.9 24.3
Soluble Protein 47 15.8
Animals and diet description ADF 18.4 32.8
NDF 29.7 36.8
Lignin 2.6 7.23
This experiment was conducted at the Wye Angus farm hosted Non-fiber NFC 47.1 28.7
by the University of Maryland, at the Wye Research & Education Starch 35.6 1.3
Centre, Queenstown, Maryland, USA (38 540 46.1500 N, 76 090 Crude fat 2.9 2.5
Ash 6.47 9.35
06.7500 W, 1 087 m average above sea level). The prevailing climate
TDN 73 63
in Queenstown is known as a local steppe climate, with a mean NEL, Mcal/Lb 0.77 0.65
annual temperature of 10.2 °C in recent ten years. The animals NEM, Mcal/Lb 0.79 0.64
were managed, fed, and harvested according to the University of NEG, Mcal/Lb 0.51 0.37
Maryland Institutional Animal Care and Use Committee. Abbreviations: NE = net energy, NE is broken down into three types: NE used for
The Angus herd in the Wye Angus farm has been closed to any lactation (NEL), NE used for maintenance (NEM), and NE used for BW gain (NEG);
animal importation for almost 70 years. Hence, it produces very NFC = Non-Fiber Carbohydrate; TDN = Total Digestible Nutrients.

2
J.A. Carrillo, Y. Bai, Y. He et al.
and 52 individuals, respectively. Body weights were recorded at
birth, weaning, and then monthly until harvest.
Genetic background
To check genetic homogenous, the genomic DNA of the 92
Angus steers was extracted from blood using the Wizard Genomic
DNA Purification Kit (Promega, Madison, WI). After extraction and
quality assessment, 66 DNA samples have been adequately
shipped to GeneSeek Company, which performed the genotyping
using the GeneSeek Genomic Profiler Bovine HDTM (GGP-HD)
BeadChip (80,000 SNPs) (Neogen, Lansing, MI). Subsequently, the
data have been analyzed with PLINK, which comprises a whole-
genome association analysis toolset (Purcell et al., 2007).
Carcass trait measurement
Beef Image Analysis Live animal carcass ultrasound was used
with an automated, chute side software system that captures
images and calculates carcass trait measurements on an objective
basis. At the termination of the study, the traits determined by Beef
Image Analysis ultrasound software were back fat thickness and
Ribeye Area (REA). After harvest, the carcasses were weighed and
then spilt longitudinally. The dressing percentage was calculated
as: dressing percentage = (carcass weight/live weight)  100%.
The ratio of Ribeye Area and Carcass weight (REA/CWT) was calcu-
lated. Yield grades were determined by using the following for-
mula: 2.50 + (2.5  adjusted fat thickness in inches) + (0.
2  percent kidney, heart, and pelvic fat) + (0.0038  hot carcass
weight) - (0.32  ribeye area at harvest in square inches). External
fat is measured in terms of thickness of fat over the ribeye muscle,
measured three-fourths of the length of the ribeye from its chine
bone end. The fat thickness was adjusted to reflect unusual
amounts of fat (or leanness) on other parts of the carcass. Kidney,
heart, and pelvic fat, or internal fat was evaluated subjectively and
expressed as a percent of the hot carcass weight. Ribeye area at
harvest was measured between the 12th and 13th ribs (where
marbling is also evaluated). Quality grades were determined by
marbling and overall maturity. The quality grade was designated:
Prime, Choice, Select, Standard, Commercial, Utility, Cutter, and
Canner, based on USDA marbling standards.
Warner-Bratzler shear force
After carcasses were maintained for 14 days in 4 °C chambers
for maturation, steaks from the Longissimus dorsi muscle were
cut at the level of the 12th intercostal space, vacuum packaged,
and immediately frozen at 20 °C. Later, to assess the Warner-
Bratzler shear force (WBSF), the steaks were thawed at 4 °C and
then cooked to an internal temperature of 70 °C employing a
George Foreman grill. An Oakton digital thermometer (Temp JKT
Acorn series) was used to measure the core temperature of the
steaks. After the samples cooled down to room temperature, six
cores of tissue (1.27 cm in diameter and parallel to the direction
of the fibers) were cut employing a hand-held coring device. The
WBSF was measured individually in each core utilizing an Instron
#5442 Test Machine (Norwood, MA). The WBSF value for each
sample was obtained, averaging the measurements of six centers
from the same steak.
Blood analysis
In 2014, five 14-month-old steers from each group were
selected at random to undergo blood analysis. Three 10 ml tubes
of blood were collected through jugular puncture; the first and sec-
ond tubes were coated with ethylenediaminetetraacetic acid and
3
Animal 15 (2021) 100381
heparin, respectively, the last one contained no additive. All sam-
ples were immediately stored in ice and shipped to the lab. The
samples were analyzed in a commercial lab (BioReliance, Rockville,
MD) following the standard procedures for typical hematological
parameters, plus carbon dioxide and glucose content.
Statistical analysis
The statistical analysis was performed in R (free software envi-
ronment for statistical computing and graphics) employing the
‘nlme’ package, which can fit and compare Gaussian linear and
non-linear mixed-effect models (Pinheiro and Bates, 1996;
Pinheiro et al., 2020). Treatment results are reported as least
squared means with a significance threshold of P < 0.05 and
P < 0.01. For quality grade analysis, a Fisher test was employed
to compare the number of individuals assigned to different cate-
gorical groups.
Results
Feed component analysis
As expected, the content of NFC and starch was much higher in
the grain diet, providing more total digestible nutrients and avail-
able energy (Table 1). Contrarily, proteins, lignin, fiber matter, and
ash were more abundant in the grass than in the grain diet. All
these differences were anticipated. However, the magnitude of
those differences should be known and considered in this study.
Genetic background
To minimize the influence of the genetic background, we com-
pleted genetic background cluster analysis between Wye Angus
and external Angus animals based on SNP genotyping. The genetic
analysis performed in 66 animals suggested no genetic differences
between the two groups (grain-fed and grass-fed). Fig. 1a shows
the principal component analysis (PCA) result from the 66 exam-
ined animals. There is no grouping of animals regarding diet style,
suggesting that the randomization process immediately after
weaning was successful in removing any of the possible genetic
differences between groups. Fig. 1b represents the result of the
same PCA analysis but including 58 external Angus animals. It
demonstrates that Wye Angus animals are genetically very homo-
geneous and cluster separately from the newly added individuals.
This detail remains critical for performing epigenetic studies,
which also is in our scope of interest.
Growth curve
At weaning, the animals showed a random difference of 13 kg
between groups with 174.5 and 187.5 kg for grain-fed finished
and grass-fed finished steers, respectively. After correction, the
weaning weight adjusted at 205 days (WW205) was also signifi-
cantly higher for grass-fed finished animals, averaging 218.1 kg
against the 206.7 kg observed in grain-fed finished individuals
(P = 0.009). The final weight for grass-fed finished animals was
marginally lower (459.6 kg) than for grain-fed finished steers
(472.4 kg) (P > 0.05). Nevertheless, the variation of weight at termi-
nation within grass-fed finished animals and grain-fed finished
individuals was similar; both groups showed a SD of approxi-
mately 35 kg. The maximum and minimum final weights with
564.7 kg at 286 days postweaning and 392.8 kg at 448 days post-
weaning correspond to a grain-fed finished, and a grass-fed fin-
ished steer, respectively. Grain-fed finished animals
demonstrated around 70% greater average daily gain (ADG) than
J.A. Carrillo, Y. Bai, Y. He et al. Animal 15 (2021) 100381

Fig. 1. Principal component analysis (PCA) grass-fed against grain-fed beef. (a) PCA in grass and grain-fed Wye Angus steers. (b) PCA obtained after adding 58 external Angus
individuals.

the grass-fed finished counterpart, ranging from 0.889 to 1.406 kg.

The behavior of the three most significant traits, as shown in
Table 2, was expected. However, similar final weights and distinct
weaning weights adjusted to 205 days between animals from dif-
ferent feeding styles were unanticipated.
The growth curve for each feeding style in four consecutive
years can be observed in Fig. 2. The year effect is observed; animals
from the years 2011 and 2014 are at the top and bottom of both
clusters correspondingly. However, the slopes of the curves within
each feeding system are very similar. The shortest period of termi-
nation corresponds to the grain-fed finished steers in 2013, with
only 254 days after weaning. Interestingly, the grass-fed finished
group from the same year shows the most extended finishing per-
iod with 448 days postweaning.
The result of a linear model fitting indicates an interaction
Fig. 2. Growth curves of grass and grain-fed Angus steers in four consecutive years.
between feeding style and age of animals (P < 0.01), thus both fac-
tors should be considered in the model for maximizing the expla-
nation of variation in the population (R2 = 0.92). The intercept and
fitted simple linear regressions extend from 0 to 286 and 461 days
coefficient were 385.34 and 1.355 for the grass-fed group, and
postweaning for the grain and grass-fed clusters correspondingly.
were 377.71 and 2.514 for the grain-fed group, respectively. The

Table 2
Statistical descriptors of economic traits in grass and grain-fed Angus steers.

Traits Feed-Style Minimum Median Mean Maximum P-value

Weaning weight (kg) Grain-Fed 135.6 173 174.6 ± 21.28 224.1 0.0054
Grass-Fed 137.4 186 187.5 ± 21.83 244.9
Weaning weight at 205 days (kg) Grain-Fed 144.7 208.2 206.7 ± 20.65 250.8 0.0095
Grass-Fed 185.5 217 218.1 ± 19.71 269.4
Final weight (kg) Grain-Fed 402.8 467.2 471.1 ± 36.49 564.7 0.1353
Grass-Fed 392.8 452.7 459.6 ± 35.87 544.3
Age at termination (days) Grain-Fed 394 459 456.5 ± 24.31 499 3.54e16
Grass-Fed 619 644 644.1 ± 20.47 713
Postweaning days at termination (days) Grain-Fed 254 282 273.9 ± 12.79 286 <2.2e16
Grass-Fed 448 454 454.2 ± 5.13 461
Average daily gain (kg) Grain-Fed 0.888 1.056 1.075 ± 0.12 1.406 1.38e14
Grass-Fed 0.453 0.635 0.639 ± 0.09 0.861

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J.A. Carrillo, Y. Bai, Y. He et al. Animal 15 (2021) 100381

Carcass traits Table 3

Quality grade of cattle carcasses regarding feeding regimen.

We found the carcass weight in grain-fed animals was 290.3 kg, Quality Grade Grain-fed Grass-fed
which was 35.6 kg heavier than the carcasses from grass-fed indi- CH+ 9 1
viduals. Interestingly, the mass of the carcasses was relatively CH 7 1
homogeneous within both groups, showing a similar SD of approx- CH 14 9
imately 23 kg. The dressing percent of steers ranged from 51.1 to Sel+ 7 6
Sel 1 6
66.5 and 58.8 to 65.8 in the grass and grain-finished animals corre- Sel 2 17
spondingly. Overall, the dressing of animals that received grain
was more homogeneous and 6.4% greater than the grass-fed indi- Fisher test (P < 0.01). CH = choice; Sel = Select.

viduals. Regarding back fat, grain-fed finished carcasses presented

0.71 cm more than grass-fed animals.
Interestingly, the REA was similar for both groups (P = 0.41); the
minimum value was 7 in2 for a grass-fed steer, and the maximum
score corresponded to a grain-fed animal with 11.8 in2. However,
given the heavier grain-fed carcasses and similar REA in both
groups, the REA/CWT was greater for grass-fed carcasses, with a
mean of 1.7 ± 0.20 against the 1.54 ± 0.16 average ratio observed
in the grain-fed cluster. The results from the carcass traits evalua-
tion are summarized in Fig. 3. Carcasses from grass-fed finished
animals showed lower yield grades, which generally were 1.22
points lower than the grain-fed finished carcasses that averaged
3.86 ± 0.58. As a reminder, USDA yield grades are ranked numeri-
cally from 1 to 5. Yield grade 1 represents the greatest yielding car-
cass and Yield grade 5, the lowest. Considering quality grades as
shown in Table 3, a more significant proportion of grain-fed car-
casses (30 of 40) were classified within USDA Choice, comparing
to the fraction of grass-fed carcasses (29 of 40) that were assigned
to USDA Select categories (P < 0.01).
Warner-Bratzler shear force
The analysis of measurements acquired from the WBSF test
indicates no difference in the tenderness of beef from animals
receiving grass or grain diet from weaning to termination
(P > 0.25). However, Longissimus dorsi muscles from grass-fed ani-
mals were slightly tender than the grain-fed individuals, requiring
0.16 less kg force to cut the samples, as shown in Fig. 4. The neces-
sary force to cut the grain-fed strips was 2.27 kg on average. Gen-
erally, the steaks from both groups were acceptably tender, with a
WBSF maximum value of 3.39 and 3.44 for the grain-fed and grass-
fed groups, respectively.
Blood analysis
The routine laboratory blood analysis yielded typical values for
all parameters. It showed no difference between animals under the
grass or grain diet, except for carbon dioxide and glucose with a
corresponding P-value of 0.04 for both components, as shown in
Table 4.
Discussion
In this study, we evaluated the grass-fed and finished beef pro-
duction in steers from a closed herd to minimize the genetic effect
while maximizing the diet effect in the results. The nutritional
analysis confirms our expectations regarding nutrient disparities
and quantifies the corresponding differences between diets. It is
well known that nutritional requirements change with the age
and physiological stage of the animal. However, the periodic

Fig. 3. Carcass trait evaluation in Angus steers under different feeding systems. REA = ribeye area; CWT = carcass weight.

5
J.A. Carrillo, Y. Bai, Y. He et al. Animal 15 (2021) 100381

Fig. 4. Warner-Bratzler shear force (WBSF) measurements in grass-fed and grain-fed beef. (a) WBSF by animal and feeding style; (b) Means of WBSF in Longissimus dorsi
muscle from animals receiving grass or grain diets.

adjustments made in highly efficient production systems to
increasing daily gain are difficult to achieve in grass-fed finished
production systems. The nutrient composition of grass depends
on many factors such as pasture species and habits, developmental
stage of grass at harvest, season, region, soil type, and climate con-
ditions, among others. Therefore, our study comprises cycles of
production in four consecutive years, following the same treatment
from birth to harvest to assess some of these possible sources of
variation.
As expected, the most significant differences between diets
were CP, fiber, and energy content. The high dietary protein level
is critical for development, and early growth; however, high energy
level is particularly crucial during the finishing period. Those dif-
ferences are reflected in more protracted birth to harvest periods
for grass-fed and finished individuals, ranging from 448 to 461 days
in comparison with the 254 to 286 days postweaning required by
the grain-fed finished steers to achieve similar final weights. The
ADG for grass-fed steers was 0.639 kg, a reasonable value com-
pared to the 0.490 kg found in a study using several breeds but
much lower than the 1.04 kg reported in Bonsmara steers
(Tatum et al., 1988). Animals that received grain gained 0.440 kg
more weight daily than grass-fed counterparts, a value comparable
to a previously found difference of 0.480 kg (Tatum et al., 1988).
The reason for the differential ADG between the grass-fed and
grain-fed systems is primarily related to the level of energy sup-
plied in the diet and the energy expenditure that animals suffer
in both conditions (Kaufmann et al., 2011). Usually, grass-fed fin-
ished animals are more exposed to adverse climate conditions,

Table 4
Blood parameters in grain-fed and grass-fed cattle.

Test Grain Grass Unit P-value

White Blood Cells 11.13 ± 2.13 8.35 ± 4.58 103 cells/lL 0.25
Neutrophils 3.23 ± 1.59 2.48 ± 0.50 103 cells/lL 0.34
Lymphocytes 6.68 ± 1.52 6.61 ± 1.41 103 cells/lL 0.94
Monocytes 0.54 ± 0.18 0.49 ± 0.09 103 cells/lL 0.63
Eosinophils 0.47 ± 0.49 0.45 ± 0.20 103 cells/lL 0.92
Basophils 0.11 ± 0.02 0.08 ± 0.02 103 cells/lL 0.06
Large Unstained Cells 0.08 ± 0.06 0.03 ± 0.01 103 cells/lL 0.11
% Neutrophils 28.60 ± 9.67 24.60 ± 3.95 % 0.41
% Lymphocytes 60.36 ± 10.16 65.00 ± 5.15 % 0.38
% Monocytes 4.84 ± 1.12 4.90 ± 0.91 % 0.92
% Eosinophils 4.42 ± 4.50 4.34 ± 1.83 % 0.97
% Basophils 1.06 ± 0.42 0.67 ± 0.37 % 0.16
% Large Unstained Cells 0.72 ± 0.39 0.36 ± 0.16 % 0.09
Red Blood Cells 8.19 ± 0.49 8.29 ± 0.36 106 cells/lL 0.72
Hemoglobin 12.34 ± 1.71 12.58 ± 1.00 g/dL 0.79
Hematocrit 34.38 ± 4.71 34.84 ± 2.62 % 0.85
Mean Corpuscular Volume 41.88 ± 4.26 42.02 ± 2.74 fL 0.95
Mean Corpuscular Hemoglobin 15.04 ± 1.54 15.20 ± 1.06 pg 0.85
Mean Corpuscular Hemoglobin Concentration 35.92 ± 0.31 36.14 ± 0.23 g/dL 0.24
Red Cell Distribution Width 18.62 ± 0.67 19.44 ± 0.88 % 0.13
Platelets 510.60 ± 134.34 384.20 ± 187.40 103 cells/lL 0.25
Mean Platelet Volume 5.94 ± 0.27 6.98 ± 2.36 fL 0.35
CO2 21.94 ± 2.78 25.38 ± 1.69 mmol/L 0.04
Glucose 101.60 ± 15.20 85.00 ± 3.87 mg/dL 0.04

6
J.A. Carrillo, Y. Bai, Y. He et al.
which could interfere with the period of grazing and increase the
energy demand for thermosregulation, limiting the number of
nutrients available for body growth.
Interestingly, despite the total randomization of animals, the
adjusted weaning weight at 205 days differed between feeding
styles; the grass-fed finished group was an average 11 kg heavier
than grain-fed finished individuals. This involuntary aspect should
be considered during inference. However, the results of this study
are still robust and provide valuable information for the beef
industry and especially for beef producers. Generally, the market
defines the final weight of animals; therefore, we adjusted our
study to the reality of beef production. Animals were sent for har-
vest when they reached the satisfactory market weight and upon
slaughterhouse requirements and schedule. Although the final
weight was similar for both groups (P = 0.13), a slight difference
of 12 kg was observed for the grain-fed finished steers. Grass-fed
finished animals generally need extra 188 days to harvest.
The effect of the year is notorious. Animals from the year 2014
were the inferiors within their correspondent groups, suggesting
that conditions during this period affected the animals’ perfor-
mance negatively. Nevertheless, it is also clear that year effect
was similar in both groups, implying no interaction between feed-
ing style and year. However, the linear regression model indicated
an interaction between feed and days postweaning; thus, individ-
ual regression lines were fitted for both groups. For the two clus-
ters, the growth was linear and constant from weaning to
termination; a similar trend was previously observed in Bonsmara
grass-fed steers in 2 years, in grass-fed tropical-breed steers and
grass-fed buffalo as well (Agastin et al., 2013; Alves and
Franzolin, 2015). Displaying similar deviations from the fitted line,
growth curves from both conditions were reasonably homoge-
neous. Alternatively, other more sophisticated models could
increase the accuracy of prediction. Still, the almost perfect linear-
ity provides a simple method to estimate growth for practical uses
and particularly for management decisions.
Regarding carcass traits, six of the eight measured characteris-
tics were different between animals that received grass or grain.
Grain-fed finished animals produced heavy carcasses (290.25 kg)
with 36 kg more than grass-fed steers. Accordingly, a review of
nine publications, including several cattle breeds, concluded that
grass-fed finished animals produce lighter carcasses than grain-
fed finished individuals (Brewer and Calkins, 2003). Dressing per-
cent reflects the weight relative to the animal’s live weight. Thus,
grain-fed dressing was 6.45% greater than grass-fed, constituting
a relevant economic difference for beef production. Accordingly,
a similar value of 54.2%, compared to the 55.4% observed in this
study, was previously reported in grass-fed tropical breeds
(Agastin et al., 2013). Because the harvest process constitutes a
fixed cost regardless of the weight of animals, high dressing score
is highly appreciated in the beef industry.
Grain-fed carcasses with 1.59 cm had 1.8-fold more back fat
than grass-fed carcasses with 0.87 cm. This variation remains close
to the two-fold change reported earlier in a study that mentioned
the positive association of subcutaneous fat with sarcomere length
and negative correlation with shear force. Additionally, the same
authors suggest that fatter carcasses endure less shortening of
muscle fibers and promote enzymatic proteolysis during the pro-
gression of rigor mortis (Bowling et al., 1977). However, increasing
subcutaneous fat beyond 10.2 mm did not affect sarcomere length,
consequently would not contribute to enhancing beef tenderness
(Bowling et al., 1977). Another study suggested that a thinner layer
of subcutaneous fat in steaks would allow easier access to heat
during cooking, reducing cooking time, and incrementing cooking
loss, affecting beef tenderness (Berry, 1993).
Interestingly, the REA in carcasses from different diet regimens
showed no difference with values of 63.54 cm2 and 61.93 cm2 for
7
Animal 15 (2021) 100381
grain and grass-fed, respectively, contrasting with another result
that reported a similar amount of 61 cm2 for grass-fed carcasses
but different area of 11 cm2 for grain-fed carcasses (Bowling
et al., 1977). Speculation for the reported differences is that ani-
mals that are receiving grain use the abundant energy to put on
lean tissue to the limit of their genetic potential, with the excess
going to fat tissue formation. Therefore, the similarity of REA
observed in our study could be explained by the genetic resem-
blance of animals and that both groups received the energy
required to achieve optimum lean tissue deposition.
The ribeye area/carcass weight ratio diverged between both
feed regimens. After knowing the divergence of carcass weight
and the similarity of the ribeye area, this result is evident.
Nonetheless, it is essential to consider that lighter grass-fed car-
casses produce similar amounts (measured in the area) of highly
valuable cuts (steaks) compared to carcasses from grain-fed steers.
This is particularly relevant for high-quality grass-fed beef produc-
tion, as most of the revenue depends on the sale of steaks. The
average Yield Grade for grain-fed carcasses was 3.86 compared to
the 2.64 value assigned to grass-fed carcasses.
Interestingly, a previous study reported no significant differ-
ence between grain and grass-finished carcasses with 2.4 and 2.1
yield grades correspondingly (Bowling et al., 1977). Another study
found a better yield grade of 1.4 in Angus  Hereford grass-fed
steers that were employed as control and never received any sup-
plement (May et al., 1992). Yield grades assess the total amount of
boneless, carefully trimmed retail cuts from the carcass. However,
the evaluation mainly considers the high-valuable parts of the car-
cass, like the round, loin, rib, and chuck. A numerically low yield
grade is preferred because it indicates higher cutability than higher
yield grade. Therefore, grass-fed steers produce carcasses with a
better percentage of boneless, carefully trimmed retail cuts than
grain-fed finished individuals, demonstrating that grass-fed beef
production could increase revenue, and particularly the benefits
generated through the commercialization of high-value cuts.
However, a recent study using molecular breeding values sug-
gests that increasing genetic potential for yield grade was unlikely
to improve overall carcass quality, given its sizeable detrimental
effect on quality grade. For instance, concurrent genetic improve-
ments for yield grade and marbling constitute a more appropriate
strategy to achieve better overall carcass quality (Thompson et al.,
2015). We also observed that although grass-fed carcasses showed
better yield grades, their quality grades were generally lower than
its grain-fed counterpart; 75% of grain-fed carcasses were classified
as USDA Choice while only 27.5% of grass-fed carcasses received
the same grade. Furthermore, only 2.5% of grass-fed carcasses
reached the maximum score observed in this study (CH+) com-
pared to the 22.5% of grain-fed carcasses that were classified as
CH+. Contrarily, 42.5% of grass-fed carcasses received the lowest
grade attained (Sel), against only 5% of grain-fed carcasses with
the equivalent score. These observations concerning quality grade
suggest that high-quality grass-fed beef production is feasible but
not easy to achieve. Therefore, beef producers should evaluate the
economic consequences of transforming conventional to grass-fed
systems, especially considering whether the extra compensation
for ‘green’ products counterbalances the loss of ‘standard’ quality
with the corresponding economic risk.
Regardless of consumers’ tradition or habits, tenderness always
has been central for beef quality judgment and became a common
desirable characteristic across consumers from different cultures.
We found no difference in WBSF between different diet types.
However, the grass-fed shear force was slightly lower than the
grain-fed value, with their correspondent score of 2.11 and
2.27 kg. This finding conflicts with previous studies that reported
lower WBSF values for grain-fed compared to forage-fed carcasses,
however, the shear forces obtained in our research for both groups
J.A. Carrillo, Y. Bai, Y. He et al.
were small compared to others (Bowling et al., 1977; Bjorklund
et al., 2014). As mentioned before, subcutaneous fat could produce
less shortening of muscle fibers and increase postmortem enzy-
matic proteolysis, improving beef tenderness. Nevertheless, we
found different back fat values, but similar tenderness measures
in both groups, suggesting that other factors such as genetics or
epigenetics could be critical for the tenderness of beef. Genetically
identical animals used in both feeding systems were still very
young at harvest; therefore, we speculate that the homogeny of
tenderness was sustained by the genetic make-up and the lack of
enough time for the age factor to manifest.
Overall, grass-fed finished steers showed a distinct growth
curve compared to their grain-fed counterparts, requiring around
180 extra days to reach a similar harvest weight. The ADG of
grain-fed animals was approximately 70% greater than grass-fed
individuals. Grass-fed Angus steers produce lighter carcasses, with
reduced subcutaneous fat and a lower percent dressing than grain-
fed finished individuals. These results will help beef producers
make management decisions. Meanwhile, compared to conven-
tional beef, the data from the research suggest an urgent need
for a unique grading system for grass-fed beef quality evaluation.
Ethics approval
All animal experiments were conducted following NIH guideli-
nes for housing and care of laboratory animals and following the
University of Maryland and College Park (UMCP) regulations after
review and approval by the UMCP Institutional Animal Care and
Use Committee (permit number R-08–62).
Data and model availability statement
None of the data were deposited in an official repository.
Author ORCIDs
Jiuzhou Song: https://orcid.org/0000-0002-6973-5959.
Tad Sonstegard: https://orcid.org/0000-0002-6446-9276.
Wentao Cai: https://orcid.org/0000-0002-5639-1755.
Yanghua He: https://orcid.org/0000-0001-6849-6493.
Yaokun Li: https://orcid.org/0000-0002-9211-0244.
George Liu: https://orcid.org/0000-0003-0192-6705.
Scott M. Barao: https://orcid.org/0000-0001-6989-7547.
Jose A. Carrillo: https://orcid.org/0000-0002-5644-7448.
Derek M. Bichhart: https://orcid.org/0000-0003-2223-9285.
Bai Ying: https://orcid.org/0000-0002-95445-3929.
Author contributions
JZ: Conceptualization, Supervision, Funding acquisition, Writing
- Review & Editing. JAC and YB: Investigation, Data Curation, For-
mal analysis, Writing - Original Draft. YH, WC and YL: Methodol-
ogy, Formal analysis, Validation. DMB, GL SMB and TS: Sampling
and SNP data.
Declaration of interest
The authors have declared no conflict of interest in the funding,
planning, design, conduction, analyses, or interpretation of the pre-
sent study.
Acknowledgements
None.
8
Animal 15 (2021) 100381
Financial support statement
This work was supported by the Jorgensen Endowment Funds
(2014) and MAES (2014) Maryland Agricultural Experimental
Station.
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