C H A P T E R

4
Reproductive management of beef cattle
Pedro L.P. Fontes, Nicola Oosthuizen, G. Cliff Lamb
Department of Animal Science, Texas A&M University, College Station, TX, United States

O U T L I N E

Introduction 57 Estrus synchronization and fixed-time

artificial insemination 60
Challenges 58
Multiple ovulation embryo transfer 66
Replacement heifers 58
In vitro fertilization 68
Postpartum cows 58
Sex-sorted semen 68
Genotype 59
Conclusion 69
Available strategies 59
Breeding season 59 References 70
Artificial insemination 60

Introduction However, additional advancements in both

With the expected increase in demand for beef
over the next few decades,1 reproductive perfor-
mance of beef cattle will not only determine the
overall efficiency of cow-calf operations and
the United States (US) beef industry, but will
significantly impact the world’s food supply.
The production of beef cattle has become a
more efficient process during the past few de-
cades, which is largely due to the development
and adoption of new technologies, as well as
an overall improvement in herd genetics.
management and technologies are required to
reach a level of animal production that can pro-
vide for the global population by 2050.2
Cow-calf operations rely on their females to
produce a healthy calf once per year to generate
revenue and remain profitable. Females that do
not produce a calf annually are utilizing re-
sources that could be used to support more
productive cattle. Therefore, reproductive
management strategies aimed at improving the
overall prolificacy and quality of calves play a
large role in increasing overall output and

Animal Agriculture
https://doi.org/10.1016/B978-0-12-817052-6.00004-5 57 Copyright © 2020 Elsevier Inc. All rights reserved.
58 4. Reproductive management of beef cattle
profitability of a beef enterprise. There are tech-
nologies available to cattle producers that can
be utilized to introduce superior genetics into
their herds, reduce the transfer of diseases,
improve both male and female fertility, and ulti-
mately increase the value of their calves. These
technologies include, but are not limited to, the
use of a defined breeding season, estrus synchro-
nization, fixed-time artificial insemination (TAI),
breeding soundness evaluations of bulls, multi-
ple ovulation embryo transfer (MOET), in vitro
fertilization, and the use of sex-sorted semen.
However, many beef producers fail to incorpo-
rate them into their production system and opt
for more traditional approaches.3 Further im-
provements to fertility, ease of application, and
reductions in overall cost will persuade more
cattle producers to adopt these reproductive
management strategies in the future. This chap-
ter will focus on available reproductive manage-
ment strategies and their application to address
current challenges in beef production systems.
Challenges
There are a number of challenges associated
with reproductive management in both beef
heifers and cows, challenges which need to be
overcome to optimize reproductive performance
in the beef industry.
Replacement heifers
Replacement heifers are the future of a beef
cattle operation and producers need to focus on
heifer development strategies that maximize
their productive potential and keep them per-
forming in the herd for years to come. To maxi-
mize lifetime productivity, heifers need to be
managed to calve for the first time at approxi-
mately 24 months of age. When a lifetime pro-
ductivity comparison between heifers calving
at 2 versus 3 years old was performed, heifers
that calved as 2 year olds produced 138 more
I. Beef cattle production
kg of weaned calves and had 6%e8% greater
economic efficiency.4,5 In addition, heifers that
calve early in the calving season not only have
a greater probability of becoming pregnant as
first calf heifers,6 but also have increased
longevity in the herd and produce more kg of
weaned calves during their overall productive
life.7 Therefore, the timing of conception within
the first breeding season is key for long-term
productivity of beef females. To become preg-
nant early in the breeding season, it is para-
mount that heifers attain sexual maturity prior
to the initiation of the breeding season. In recent
decades, significant progress has been made in
understanding physiological events associated
with the attainment of puberty in heifers,8,9
which has allowed for the development of
several strategies that increase the percentage
of pubertal heifers in the herd prior their first
breeding season.
Postpartum cows
One of the main factors known to influence
reproductive performance of beef herds is the
proportion of mature cows in anestrus at the initi-
ation of the breeding season.10 After parturition,
beef cows undergo a transitional period of anes-
trus characterized by a wave-like pattern of follic-
ular growth where dominant follicles undergo
atresia prior to ovulation due to a lack of luteiniz-
ing hormone (LH) pulses.11 Results of multi-
location studies evaluating postpartum cyclicity
of Bos taurus beef cows in the US indicate that
an average of 50% of cows are anestrus prior to
the breeding season. Those studies also demon-
strated great variation among different locations,
with the proportion of cyclic cows ranging from
17% to 67%.12 Because beef cows in anestrus at
the beginning of the breeding season have lower
fertility when compared to cyclic cows,13 strate-
gies that increase fertility of non-cyclic cows or
increase the proportion of cows cycling prior to
the time of breeding have the potential to
improve reproductive efficiency of beef herds.
 Available strategies
The effects of days postpartum on pregnancy
rate to TAI in suckled beef cows is well docu-
mented. Cows that are less than 50 days post-
partum at the initiation of the breeding season
have significantly poorer pregnancy rates
compared to cows that are greater than 50 days
postpartum.14e16 A study including more than
8,500 postpartum suckled beef cows, revealed
that both multiparous and primiparous cows
have decreased pregnancy rates if the interval
between calving and TAI is less than 72 days.
Furthermore, results of the same study indicated
that early calving multiparous cows in adequate
body condition have the greatest pregnancy
rates, whereas late calving primiparous cows in
poor body condition have the poorest pregnancy
outcomes,17 indicating that parity, timing of
calving, and postpartum nutritional status are
crucial to subsequent reproductive performance.
Primiparous cows represent an even greater
challenge. Although heifers are commonly
mated to calve earlier and have greater days
postpartum at initiation of the breeding season
than primiparous cows, a greater proportion of
primiparous cows are in anestrus at the begin-
ning of the breeding season when compared to
multiparous cows.13 Since the percentage of cy-
clic cows at the beginning of the breeding season
increases curvilinearly from 9% at
30 days, to a
peak of 70% at 81e90 days postpartum, strate-
gies that increase the proportion of cows calving
early in the calving season can have major bene-
fits to improve fertility in beef cows, particularly
primiparous cows.
Genotype
It is estimated that approximately 70% of the
increase in beef production required to meet the
growing global demand will come from subtrop-
ical/tropical regions of the planet,1 including
the southern US, Mexico, Central/South
America, Africa, Asia, and Oceania. These re-
gions contain approximately 70% of the world’s
cattle population, which are predominantly Bos
I. Beef cattle production
59
indicus-influenced or Zebu breeds fed diets based
on warm-season forages and agricultural by-
products. In the US alone, approximately 30%
of cattle contain B. indicus genetics, and approxi-
mately 40% of beef cows and 50% of the country’s
cow-calf producers are located in the southern
US, where B. indicus cattle and their crosses are
located. Because these cattle have undergone nat-
ural selection in hot and humid environments,
they have acquired a greater tolerance to heat
and parasites when compared to B. taurus cattle,
which underwent environmental selection in the
arid climates of Europe and Africa. Therefore,
B. indicus cattle are better able to regulate body
temperature in response to heat stress, and conse-
quently experience a less severe decrease in feed
intake, growth, milk yield and reproductive per-
formance when exposed to heat stress condi-
tions.18 Zebu cattle have been subjected to less
selection pressure for production traits, such as
meat and milk production, compared to
B. taurus breeds. Currently, challenges include
enhancing beef production efficiency in tropical
and subtropical regions, and maximizing the ben-
efits of using B. indicus genetics while minimizing
their limitations in intensive beef production
systems. It is important to highlight that
several physiological differences exist between
B. indicus and B. taurus cattle, including differ-
ences related to reproductive physiology.19,20
Available strategies
Breeding season
One of the most important reproductive man-
agement practices for a cow-calf operation is the
establishment of a breeding season. By defining
a breeding season, a producer is able to develop
a defined calving season to match environmental
conditions and available resources to gestation
and calving periods of beef females, and to
ensure that they receive adequate nutrients dur-
ing those times. In addition, a breeding season
allows for the concentration of labor resources,
60 4. Reproductive management of beef cattle
and more attention devoted to cows and heifers
when calving to minimize the consequences of
dystocia events, and maximize calf survival. A
survey conducted in the United States showed
that only 34% of beef cattle operations had a
defined breeding season, 11.5% of operations
had two defined breeding seasons, and as
many as 54.5% of operations had no set breeding
season.21
Many traditional commercial breeding sea-
sons have the intent to place a young, growing
calf on forages that are at their peak of quality
and availability. Providing growing calves and
lactating dams high quality forages allows for
maximal calf weight gain through both
increased milk and forage intake.22 Without a
defined breeding season, producers have diffi-
culty implementing certain reproductive bio-
technologies, and must monitor cows for
calving throughout the year. By establishing a
defined breeding season, calving activity is
reduced from being year-round to a specific
period of time. More calves will be available
for sale at a given time, and these calves will
have greater uniformity in terms of size and
weight, resulting in an increase in market
value.23 Market value of calves increases as a
result of an increase in the number of calves in
an auction lot, resulting in greater financial
income.24
Artificial insemination
Artificial insemination (AI) was one of the
first biotechnologies used in farm animal species
to improve reproduction. The first reported AI
success in a domestic animal was performed by
Lazzaro Spallanzani in 1784 with successful AI
in a dog. However, it was only in the early
1900s that a Russian scientist named E.I. Ivan-
how accomplished the first successful AI in cat-
tle.25 Current techniques for AI have improved
drastically with the development of semen eval-
uation techniques, semen extenders, and proper
methods for freezing semen. Between 1990 and
I. Beef cattle production
2017 the number of units of dairy semen sold
increased 84%, whereas the number of units of
beef semen increased 145%.26 This increase in
units of semen sold per year indicates that there
has been a greater adoption of AI in the beef
cattle industry over time.
By utilizing AI, superior genetics can be intro-
duced into a herd in a shorter period of time than
through the use of a natural service sires alone.27
Semen from proven bulls with the best genetics
and most desirable expected progeny differences
(EPDs) can be purchased easily and utilized in
any beef cattle operation. By selecting for bulls
with low birth weights and calving ease EPDs,
calving difficulty can be minimized and calf los-
ses associated with dystocia can be reduced. In
addition, AI can reduce the number of natural
service sires required by a producer, as well as
their associated maintenance expenses.23 The
use of EPDs support rapid genetic advance-
ments, which subsequently lead to an increase
in overall profitability.28 Sires producing semen
for AI have EPDs and accuracies of EPDs that
are superior to the majority of those from natural
service sires. Even when EPDs between AI sires
and natural service sires are similar, the accu-
racies of the EPDs from AI sires are greater;
therefore, more confidence can be placed in the
performance characteristics of the AI-sired
offspring.29
Estrus synchronization and fixed-time
artificial insemination
Significant improvements have been made in
our understanding of the physiology controlling
the bovine estrous cycle. Those advances laid the
foundation for the development of estrous syn-
chronization (ES) protocols that utilize exoge-
nous hormones to synchronize estrus and
ovulation in cattle. The primary objective of
these hormonal treatments is to manipulate the
estrous cycle in order to facilitate the adoption
of biotechnologies such as AI and embryo trans-
fer by cattle producers. Prior to the establishment
 Available strategies
of effective protocols for synchronization of
estrus, the labor associated with visual detection
of females in estrus was a major factor limiting
adoption these biotechnologies. With the devel-
opment of ES it is now possible to AI a herd of
cows or heifers at a pre-determined fixed time
and achieve pregnancy rates greater than 50%
without the need for estrus detection.15,30 Conse-
quently, the establishment of effective ES proto-
cols has significantly impacted reproductive
management in beef cattle.31 Over the past two
decades, there has been a remarkable global in-
crease in the use of AI by beef cattle producers.
Although significant progress has been made,
and the rate of adoption of biotechnologies has
increased, natural service by itself is still by far
the main reproductive management strategy uti-
lized by beef producers. It has been estimated
that only 7.6% of the beef operations in the US
utilize AI, and only 1.6% make use of embryo
transfer.21 Conversely, it has been estimated
that 89.3% of dairy operations utilize AI, and
8.9% utilize embryo transfer.32 The low rate of
adoption for these biotechnologies in the beef in-
dustry is likely related to the extensive nature of
beef cow-calf operations, indicating that man-
agement strategies that minimize the amount
of times producers are required to handle
their cattle are more likely to be adopted and
incorporated into management strategies of pro-
ducers with large scale operations in the beef
industry.
As our understanding of the wave-like
pattern of follicular growth increased with the
use of ultrasonography, research on the exoge-
nous control of the lifespan of the corpus luteum
(CL), as well as the use of GnRH agonists to
induce ovulation,33 set the stage for develop-
ment of OvSynch. OvSynch was the first ES pro-
tocol that achieved sufficient synchrony of
ovulation in the herd to allow the use of TAI.34
The OvSynch protocol was developed and thor-
oughly validated for use in dairy cows.34e37
Together with its variations, the OvSynch proto-
col remains the most commonly utilized ES
I. Beef cattle production
61
protocol in the dairy industry.38 The OvSynch
protocol consists of an injection of a GnRH
analogue to induce ovulation of potentially
responsive follicles, and induce the start of a
new follicular wave. Seven days after the first
GnRH injection, an injection of prostaglandin
F2a (PGF) is administered to induce luteolysis
of a potentially present CL, thereby decreasing
circulating concentrations of progesterone (P4)
and facilitating final growth of the dominant fol-
licle. A second GnRH injection administered 48 h
after the PGF induces ovulation of the pre-
ovulatory dominant follicle, and TAI is per-
formed 16 h after the second GnRH injection.34
As previously mentioned, the more extensive
nature of beef cow-calf operations requires that
cattle handling is minimized in order to facilitate
the adoption of ES and TAI. Hence, efforts were
devoted to developing protocols that reduce the
number of times animals are handled. This led to
the development of the CO-Synch protocol in
which TAI is performed concurrently with the
second GnRH injection and yields similar preg-
nancy rates as those achieved using OvSynch.39
However, a disadvantage of both the OvSynch
and the CO-Synch protocols was that 10e20%
of the cows exhibited estrus prior to or directly
after the PGF injection. Unless those cows were
AI following visual detection of estrus, they
failed to become pregnant. Hence, a controlled
internal drug release (CIDR) insert impregnated
with P4 was developed, and incorporated into
the CO-Synch protocol from day 0e7 of the reg-
ular CO-Synch protocol. The supplemental P4
from the CIDR resulted in significant increases
in pregnancy rates to TAI,13e15 and has, there-
fore, been adopted extensively in the beef indus-
try. Today, the 7-d CO-Synch þ CIDR protocol is
the most commonly utilized protocol in the US to
synchronize both beef cows and heifers for TAI.
By handling cows or heifers three times, beef cat-
tle producers can now synchronize ovulation
and AI hundreds of beef females in a single
day and attain more than 50% pregnancy rates
on the first day of the breeding season.15,30
62 4. Reproductive management of beef cattle

The adoption of ES and TAI can have a major
impact on the profitability of a cow-calf opera-
tion. Modeling exercises indicate a potential in-
crease in net return of $25e40 per calf born
from AI.40 Additionally, 72% of respondents to
a survey estimated that the additional value of
calves from AI breeding compared with natural
service breeding was more than $20, whereas
48% of respondents estimated the additional
value to be more than $50.41 Data generated
from the sales of the Show-Me Replacement
Heifer Inc. revealed a premium of $18.69 per
pregnant heifer with a calf from AI, and a pre-
mium of $24.30 per pregnant heifer that was
due to calve during the first 30 days of the
calving season.42 However, it is important to
mention that some of these economic models
may underestimate the magnitude of the eco-
nomic impact ES and TAI. Some of the benefits
associated with the adoption of these technolo-
gies go beyond the incorporation of superior
genetics through the use of AI. When a multi-
location study compared ES and TAI with
natural service, an increase in the proportion of
cows calving early in the calving season was
observed for cows exposed to TAI. Since calves
that are born earlier in the calving season are
older at the time of weaning and have more
time to gain weight between birth and weaning,
shifting the time of calving through the use of ES
and TAI increased the weaning weights of
calves. Furthermore, a greater proportion of
cows exposed to TAI weaned a calf compared
to cows only exposed to natural service23
(Fig. 4.1).
The impacts of calving date on productivity of
offspring and reproductive efficiency of cows are
well documented. Data collected over 13 years at
the University of Nebraska’s Gudmundsen
Sandhills Laboratory indicated that weaning
weights are greater for steers born in the first
21 days of the calving season, when compared
to steers born in the second and third 21 day in-
tervals.43 In the same study, steers were tracked
until slaughter, and final body weight, hot
carcass weight, marbling score, and carcass

FIG. 4.1 Percentage of cows calved in 10-day increments of the calving season for cows exposed to the 7-d CO-
Synch þ CIDR protocol followed by fixed-time artificial insemination (TAI) or cows mated by natural service with no estrous
synchronization (Control; adapted from Rodgers et al., 2012). *Within 10-day interval treatments differ (P < 0.01). **Within
10-day interval treatments differ (P < 0.05).

I. Beef cattle production

 Available strategies
value were greater for steers born earlier in the
calving season when compared to steers from
the same herd that were born later in the calving
season. The influence of early calving on produc-
tivity is not restricted to male calves. Heifers
born in the first 21 days of the calving season
were heavier at weaning and at the time of
breeding than those born later in the calving sea-
son. Consequently, a greater proportion of those
heifers were pubertal prior to the first breeding
season, more became pregnant, and a greater
percentage calved earlier as first calf-heifers
when compared to heifers born in the following
21 days or later.43 Calving distribution is also
associated with longevity and lifetime produc-
tivity of beef females. By culling females strictly
based on reproductive failure, heifers that calve
in the first 21 days of their first calving season
have increased longevity compared to heifers
that calve later,44 indicating that calving date in-
fluences the chances of primiparous cows
becoming pregnant in the subsequent breeding
season. Additionally, heifers that calved in the
first 21 days of their first calving season weaned
heavier calves during their first six breeding
seasons.
The main challenge limiting the ability of
cows to become pregnant early in the breeding
season is postpartum anestrus. The end of gesta-
tion in cattle is characterized by reduced ovarian
activity. High circulating concentrations of
placental-derived steroids suppress the release
of gonadotropins and result in the accumulation
of follicle stimulating hormone (FSH) and deple-
tion of LH stores in gonadotrophs of the anterior
pituitary.45 After calving, circulating concentra-
tions of FSH increase, and the wave-like pattern
of follicular growth is re-established.46 However,
dominant follicles fail to ovulate due to a lack
of LH stored in the anterior pituitary, which
limits the pre-ovulatory LH surge required for
final follicular maturation and ovulation, and
results in atresia of the dominant follicle.47
Luteinizing hormone stores are re-established
approximately 2e3 weeks postpartum in cows.
I. Beef cattle production
63
Ovarian cyclicity and ovulation are rapidly
resumed in cows when calves are weaned at
birth.47 However, cows that continue to nurse
their calves experience a transient suppression
of the GnRH surge, which is required for an
LH surge; therefore, follicles fail to ovulate and
undergo atresia during the early postpartum
period in suckled beef cows. As the postpartum
period progresses, the negative effects of the
presence of the calf decrease, allowing for devel-
opment and ovulation of the dominant follicle.48
Having a large number of cows cycling prior to
the breeding season is key to increasing the num-
ber of cows pregnant at the beginning of the
breeding season. The proportion of cyclic cows
at the beginning of the breeding season increases
as the number of days postpartum increase.13
Consequently, cows that calve earlier in the
calving season are more likely to have resume
estrous cycles prior to the subsequent breeding
season which increases their chances of
becoming pregnant early in the season. This
concept is well illustrated in Figs. 4.2 and 4.3.
When multi-herd fertility studies are conducted,
it is common to observe a wide variation in preg-
nancy rates to TAI among herds, and several fac-
tors may contribute to these differences, such as
genetics, nutrition, herd management, and
others. Fig. 4.2 summarizes data collected from
1541 cows in eight different herds. When
comparing herds that had pregnancy rates to
TAI greater than 50% with herds that had preg-
nancy rates
50%, the better performing herds
had, on average, 88% of the cows calving in the
first 30 days of the calving season, whereas the
less fertile herds had only 44% of cows calving
in the first 30 days of the breeding season
(Fig. 4.3A, B). Consequently, the average number
of days postpartum at the beginning of the
breeding season for cows in the top performing
herds was 79 days versus 64 days in the remain-
ing herds. Furthermore, the top performing
herds only had 7% of cows that were less than
50 days postpartum at the beginning of the
breeding season, whereas the other herds had
64 4. Reproductive management of beef cattle

FIG. 4.2 Differences in pregnancy rates among eight different herds. All cows were exposed to a 7-d CO-Synch þ CIDR
estrus synchronization protocol followed by fixed-time artificial insemination 60e66 h after CIDR removal. Marron
(black in print version) bars represent the herds with pregnancy rates greater than 50%, whereas gray bars represent the herds
with pregnancy rates of less than 50%.

43% of cows that were less than 50 days post-
partum. These results highlight the importance
of having cows calve early in the breeding sea-
son, and indicates that management strategies
that increase the proportion of cows calving
early in the calving season also influence fertility
during subsequent breeding seasons.
Since exogenous P4 can induce cyclicity in
anestrous cows and prepubertal heifers, ES pro-
tocols that include supplemental P4 can be used
strategically by cattle producers to increase the
number of cows and heifers that are pregnant
early in the breeding season. The benefits of P4
supplementation are particularly important for
producers with B. indicus cattle. Zebu females
generally reach puberty at 22e36 months of
age when managed in an extensive production
system.49 Accordingly, age at first calving in
these heifers can be as high as 44e48 months.
Attention has been focused on the development
of strategies to reduce the age at puberty in
Zebu heifers through genetic selection, nutri-
tional management,50 and pharmacological
treatments.51e53 It is important to mention that
the regulation of use of exogenous hormones
for ES, such as estradiol and equine chorionic
gonadotropin (eCG), differ among countries.
Although the use of estradiol benzoate, estradiol
cypionate, and eCG for ES and puberty induc-
tion is not legal in the US, their use has been le-
gally approved in countries such as Brazil and
Australia. Therefore, protocols with acceptable
pregnancy rates have been established for TAI
in both B. indicus cows54,55 and heifers in those
countries.53 Approved protocols can effectively
induce puberty in approximately 80% of prepu-
bertal B. indicus heifers,52 which can then be
exposed to ES and TAI, yielding conception rates
of approximately 50% for those heifers that
respond to induction of puberty.53 The effective-
ness of these strategies, however, is highly
dependent on the nutritional and metabolic sta-
tus of the heifers. Increased nutrient intake and
accelerated rates of body weight gain during
specific periods of heifer growth facilitate puber-
tal development by programming hypothalamic

I. Beef cattle production

 Available strategies 65

FIG. 4.3 Calving distribution of cows exposed to a 7-d CO-Synch þ CIDR estrus synchronization protocol followed by
fixed-time artificial insemination 60e66 h after CIDR removal. Panel A: These results are from herds that had pregnancy rates
greater than 50%. Eighty-eight percent of the cows calved within the first 30 days of the previous calving season. Panel B: These
results are from herds that had pregnancy rates of less than 50%. Only 44% of the cows calved within the first 30 days of the
previous calving season.

centers that regulate the onset of puberty.50 heifers that have reached puberty prior to their
Therefore, feeding high energy diets during spe- first breeding season.56 Combining an intensive
cific periods of development can be useful to nutritional management program with a phar-
increase the proportion of B. indicus-influenced macological protocol for induction of puberty

I. Beef cattle production

66 4. Reproductive management of beef cattle
can yield acceptable pregnancy rates in Zebu
heifers between 12 and 15 months of age.
When B. indicus and B. indicus  B. taurus heifers
were fed to reach approximately 300 kg prior to
breeding between 12 and 15 months of age and
were exposed to a P4 and estradiol-based pu-
berty induction protocol, acceptable pregnancy
rates to TAI were observed.57 However, there
are currently no effective GnRH-based protocols
for ES and TAI in B. indicus beef heifers. The use
of short-term protocols, such as the PG 5-d CO-
Synch þ CIDR protocol have been evaluated in
B. indicus-influenced replacement heifers.58
Exposing heifers to ES and TAI resulted in a
greater proportion of heifers becoming pregnant
within the first 21 days of the breeding season;
however, others have reported inconsistent
results when utilizing the same protocol for
B. indicus heifers.59 Therefore, there is a need
for the development of ES strategies tailored spe-
cifically for B. indicus-influenced heifers that do
not rely on estradiol-based products.
There are several ES and TAI protocols for use
in mature B. indicus beef cows; however, these
protocols rely on the use of estradiol and often
require the use of eCG or temporary calf
removal. Because estradiol and eCG products
are not commercially available for cattle pro-
ducers in the US, there are limited alternatives
available for ES and TAI in cows with a large
B. indicus influence. The PG 5-d CO-
Synch þ CIDR protocol is the only GnRH-
based protocol currently recommended for TAI
in mature beef cows that does not rely on
estradiol-based products. This protocol is similar
to the 5-d CO-Synch þ CIDR widely utilized in
B. taurus animals; however, a PGF injection is
given at the beginning of the protocol in conjunc-
tion with the first GnRH, and a second injection
of PGF is administered 8 h after the first at CIDR
removal. Additionally, TAI is performed 66 h af-
ter CIDR removal rather than at 72 h. The ratio-
nale behind the PGF injection at CIDR insertion
is based on data indicating that B. indicus females
appear to be more sensitive to the effects of P4 on
I. Beef cattle production
gonadotropin release.60,61 Hence, decreasing
concentrations of P4 may facilitate follicular
development, and consequently improve
pregnancy rates in these females. The value of
elimination of the initial GnRH in the PG 5-
d CO-Synch þ CIDR protocol is under investiga-
tion.62 Removing the initial GnRH injection may
allow for the elimination of the second PGF injec-
tion 8 h after CIDR removal; however, limited
data are available for the recommendation of
the later strategy. In summary, the combination
of pharmacological hormonal supplementation
and adequate nutritional management, together
with continuous genetic selection for early
maturing heifers, have the potential to impact
reproductive efficiency of beef females raised in
tropical and subtropical regions.
Multiple ovulation embryo transfer
The utilization of embryo transfer is an addi-
tional opportunity for genetic improvement in
a cattle operation. Through embryo transfer, a
single, genetically superior female is able to
generate a greater number of offspring than
through a conventional system, and when
coupled with spermatozoa from a genetically
outstanding sire, embryos of exceptional genetic
quality can be produced. In addition, recipient
females of poor or average genetic merit have
the opportunity to serve as surrogates and
receive an embryo with high genetic value and
give birth to calves with greater genetic merit.
Through embryo transfer, genetic progress can
be hastened, which is particularly useful in cattle
due to their relatively long generation interval
when compared to other livestock species.
Another great advantage of this technology is
the ability to transport embryos to areas where
biotechnologies for the production of beef need
to be advanced, instead of having to transport
live animals themselves.
Since 1951, when the first calf was produced
by embryo transfer, biotechnologies have
evolved to allow embryo transfer to take place
 Available strategies
in a commercial setting.63 Initially, embryos
were transferred using a surgical procedure in
which the uterine horn was exteriorized and
the embryo was transferred into the lumen.
However, improvements now allow for success-
ful transcervical transfer of bovine embryos.64 In
current MOET protocols, donor females are
superovulated using FSH, subjected to AI, and
a number of embryos are recovered through a
uterine flushing technique. Following collection,
viable embryos are either transferred fresh to
recipient females or frozen for future use. On
average 6.9 viable embryos are recovered per
uterine flushing in beef females65; however,
this number fluctuates depending on breed and
age of cow, as well as within breed variation.
The transfer of fresh embryos typically yields
10%e15% greater pregnancy rates than those us-
ing frozen-thawed embryos.66,67
According to the annual statistical survey of
the Data Retrieval Committee for the Interna-
tional Embryo Transfer Society (IETS), the num-
ber of fresh and frozen bovine embryos
transferred has increased exponentially from
200,000 in 2008, to more than 400,000 in 2017.65
Of those embryos, 52% were transferred in dairy
cattle, and 48% in beef cattle. Furthermore, the
number of embryo transfers performed globally
increased from 361,000 in 1997 to 506,000 in 2012
(Fig. 4.4).22 Overall, the success of embryo trans-
fer depends on a variety of factors associated
with the embryo, the recipient, the embryo trans-
fer technician, or an interaction among those
factors.68 Suitability of recipients is dependent
on several management, nutritional, and estrous
cycle control factors to ensure the presence of a
functional CL at the time of embryo transfer.68
It is necessary to control the stage of the estrous
cycle in order to achieve acceptable synchrony
between donors and recipients of the embryos.
Development of ES has allowed this synchrony
between uterus and embryo to be established,
reduced the amount of recipients required, and
increased the ease of incorporation of embryo
transfer into cattle operations. In addition, ES
I. Beef cattle production
67
FIG. 4.4 The number of embryos transferred that were
produced per year by in vivo and in vitro techniques (IETS
Data Retrieval Committee Reports; https://www.iets.org/
accessed December 5, 2018).
has enabled the use of fixed-time embryo trans-
fer (FTET), which eliminates the need for estrus
detection in recipient cows. Pregnancy establish-
ment is most successful when embryos are trans-
ferred into estrus synchronized cows 6e8 days
after being detected estrus or GnRH injection.69
Superovulation protocols increase the num-
ber of oocytes ovulated and fertilized to produce
multiple embryos per estrous cycle. Currently,
the recommended superstimulatory protocol
for B. taurus donors involves inserting a CIDR
on day 0, followed by injection of 100 mg GnRH
2 days later. Beginning on day 4, donors receive
injections of FSH every 12 h. The amount of FSH
per injection decreases each day until day 7,
resulting in a total of 8 injections and a total of
400 mg of FSH. On day 7, donors receive 2 injec-
tions of PGF2a 12 h apart (AM/PM). At the time
of the second PGF2a injection, the CIDR insert is
removed, and heat detection begins 24 h after
CIDR removal and continues until day 11. Do-
nors that are detected in estrus during this
period are AI at both 12 and 24 h after onset of
estrus. Embryos are flushed 7 days after AI.70
A major limitation to the production of embryos
from MOET has been the lack of reliability
in successfully inducing superovulation in
donor females.71 However, research into the
68 4. Reproductive management of beef cattle
development of superovulation protocols and
techniques to predict which donor females may
respond well to superovulation is ongoing.
In vitro fertilization
An alternative to embryos derived from
MOET is the production of embryos in a labora-
tory via in vitro maturation (IVM) and in vitro
fertilization (IVF) of oocytes followed by
in vitro embryo culture (IVC), which are collec-
tively referred to as IVM/IVF. The first success-
ful generation of live offspring by IVM/IVF
was achieved in rabbits in 1959.72 From then on-
wards, IVM/IVF technology has improved
drastically to include development of embryo
cryopreservation, which was first successful in
1972 with mouse embryos, and a year later
with bovine embryos.73 Following cryopreserva-
tion, the need for in vitro sperm capacitation was
demonstrated and led to the birth of the first live
calf from IVF using fresh semen in 1981.74 Two
years later IVM/IVF embryos were successfully
generated using frozen semen.75
During the IVF process, oocyte maturation is
required such that oocytes complete their first
meiotic division.76 Similarly, spermatozoa used
for IVF need to undergo capacitation before
they are able to fertilize the oocyte.77 Oocytes
that mature spontaneously in vitro or in vivo
are highly receptive to fertilization. However,
oocytes matured in vitro have reduced develop-
mental capacities in comparison to those
matured in vivo.76 In addition, the viability of
IVF-derived embryos decreases with cryopreser-
vation to a greater extent than in vivo-derived
embryos; therefore, these embryos are more
likely to be transferred fresh.73
The predominant oocyte collection technique
is known as aspiration or ovum pick up (OPU).
Through OPU, unfertilized oocytes can be har-
vested directly from the ovarian follicles of a
donor cow or heifer using an ultrasound probe
and an aspiration needle. This technique may
be performed two to three times during a
I. Beef cattle production
cow’s estrous cycle for as long as 6 months,78
which is more frequent than what MOET can
be performed. Therefore, a greater number of
transferrable embryos per donor can be gener-
ated through IVM/IVF than through MOET.
As improvements to IVM/IVF techniques are
made, costs to generate embryos will likely be
reduced, leading to a greater increase in the
adoption of IVM/IVF produced embryos.
Over the past 15 years the number of in vitro-
produced embryos has increased by more than
300% (Fig. 4.4),22 and according to the IETS
(2017), 58% of the embryo transfers performed
in 2016 were with embryos generated through
IVM/IVF. In vitro fertilization may generate
pregnancies from a donor female that is already
pregnant, and requires fewer units of semen.
Furthermore, oocytes can be collected from the
antral follicles of ovaries obtained from slaughter
facilities, which greatly increases the number of
embryos that can be produced, and can eliminate
the need for donor females. Finally, the potential
disadvantage of a poor response to a superovu-
lation protocol can be avoided by utilizing
IVM/IVF.
Sex-sorted semen
One of the more recent biotechnologies used
in beef cattle operations is that of sex-sorted or
sexed semen. Through flow cytometry, sperm
cells carrying either an X (X-sperm) or Y chromo-
some (Y-sperm) are separated based on DNA
content, where X-sperm contain approximately
4% more DNA than Y-sperm.79 Flow cytometry
was first developed in the early 1980s; however,
it produced de-membraned, unviable sperm. By
1989 the procedure had been refined and was
able to sort sperm cells without killing or
severely damaging them, and in 1991 the sorting
procedure was patented by the United States
Department of Agriculture.79 The first live birth
from sexed semen was in 1989 when rabbits
were surgically inseminated,80 and the first
calves were produced using sexed semen by
 Conclusion
nonsurgical AI in 1997.81 In 2003, commercial
use of sexed semen accelerated when Sexing
Technologies Inc. (Navasota, TX) was granted a
sorting license.82
Since 2007, the commercialization of sex-
sorted semen has drastically increased due to
enhanced equipment and improvements in pro-
cessing procedures. During the sorting proced-
ure, sperm cells are stained using a fluorescent
dye, Hoechst 33342, which penetrates the sperm
membranes and binds to DNA. A laser provides
a wavelength of light that causes sperm cells to
fluoresce, and a computer will detect and
analyze the amount of fluorescence given off.
X-chromosome bearing sperm give off approxi-
mately 4% more fluorescence than Y-sperm,
because they have an additional 4% of DNA.
X- and Y-sperm with varying levels of fluores-
cence are given different electrical charges that
allows them to be sorted into different containers
when passing between electrical fields.83
There are a number of benefits associated
with the utilization of sex-sorted semen, such
as selecting calf gender with greater than 90% ac-
curacy, faster genetic progress, and the removal
of defective sperm through the sorting process.79
In addition, it is easy to incorporate the use of
sexed semen into a management system if AI is
already being performed, as it will not change
the workflow. However, there are disadvantages
that can hamper the adoption of this technology.
First, pregnancy rates are generally lower when
using sexed semen compared to conventional
semen, and are usually in the range of 80%e90%
of those from conventional semen.79,82 This
reduction in fertility is one of the largest hin-
drances to its use in beef cattle, and is largely
due to a lower post-thaw motility, a reduced
number of sperm cells with intact membranes,
and acrosomal alterations that can occur during
the sorting process.84 Another disadvantage is
that sexed semen is more expensive than conven-
tional semen, which limits its economic feasi-
bility. Last, there are currently no official TAI
protocols established specifically for the use of
I. Beef cattle production
69
sex-sorted semen, which limits its adoption in
the beef industry. Therefore, opportunities exist
to optimize pregnancy rates to TAI by developing
protocols specifically for the use of sex-sorted
semen.
The primary utilization of sexed semen is in
the dairy industry to generate heifer calves that
are able to replace cows for milk production. In
the beef industry, sexed semen is used to pro-
duce replacement females, and to produce males
for the production of beef, since bulls and steers
are more efficient at converting feed to muscle.
Purebred operations use sexed semen to
generate progeny of the desired sex, such as
bulls from superior sires or daughters from elite
cows. Sex-sorted semen can be used in conjunc-
tion with IVF to produce embryos of a desired
sex. Combining sexed semen with IVM/IVF
has achieved pregnancy rates greater than those
of sexed semen used in combination with TAI.85
Although sexed semen is currently utilized in the
beef industry, large-scale adoption will require
TAI protocols that result in acceptable preg-
nancy rates. In the past, TAI with sexed semen
has resulted in pregnancy rates between 47%
and 70% of those from conventional semen.86
However recent improvements in the semen
sorting procedure has yielded pregnancy rates
of approximately 87% of those from conven-
tional semen.87 Advancements to sperm sexing
technologies are continually being made.
Together with genetic selection, the use of sexed
semen is a good strategy to produce genetically
superior animals of the desired sex. As costs
decline, and as greater pregnancy rates are
achieved, sexed sperm may be increasingly
adopted in the beef cattle industry.
Conclusion
Through the incorporation of reproductive
management strategies and biotechnologies,
there is potential to improve reproductive effi-
ciency, genetic quality, and animal performance
70 4. Reproductive management of beef cattle
in the beef cattle industry. The level of incorpora-
tion of these strategies will depend on future
environmental conditions, production systems,
ease of use, infrastructure, and cattle markets;
however, these technologies will allow pro-
ducers to maximize the potential of available
resources, and aid in the production of sufficient
animal protein to provide for the expanding
global population.
References
1. FAO. Global agriculture towards 2050. In: High Lev
Expert Forum-How to Feed World 2050. 2009:1e4.
2. FAO. How to feed the world in 2050. Insights from an
Expert Meet FAO. 2009;2050(1):1e35. https://doi.org/
10.1111/j.1728-4457.2009.00312.x.
3. NAHMS. Reference of Beef Cow-Calf Management Practices
in the United States, 2007-08. 2009. Fort Collins, CO.
4. N
~ ez-Dominguez R, Cundiff LV, Dickerson GE,
un
Gregory KE, Koch RM. Lifetime production of beef
heifers calving first at two vs three years of age.
J Anim Sci. 1991;69(9):3467e3479. https://doi.org/
10.2527/1991.6993467x.
5. Nunez-Dominquez R, Cundiff LV, Dickerson GE,
Gregory KE, Koch RM. Effects of managing heifers to
calve first at two vs three years of age on longevity and
lifetime production of beef cows. In: Beef Research Program
Progress Report 2. Ames: USDA-ARS; 1985:33e35.
6. Burris MJ, Priode BM. Effect of calving date on subse-
quent calving performance. J Anim Sci. 1958;17(3):
527e533. https://doi.org/10.2527/jas1958.173527x.
7. Lesmeister JL, Burfening PJ, Blackwell RL. Date of first
calving in beef cows and subsequent calf production.
J Anim Sci. 1973;36(1):1e6. https://doi.org/10.2134/
jas1973.3611.
8. Amstalden M, Cardoso RC, Alves BRC, Williams GL.
Reproduction Symposium: hypothalamic neuropep-
tides and the nutritional programming of puberty in
heifers. J Anim Sci. 2014;92(8):3211e3222. https://
doi.org/10.2527/jas2014-7808.
9. Perry GA. Factors affecting puberty in replacement beef
heifers. Theriogenology. 2016;86(1):373e378. https://
doi.org/10.1016/j.theriogenology.2016.04.051.
10. Short RE, Bellows RA, Staigmiller RB, Bernardinelli JG,
Custer EE. Physiological mechanisms controlling anestrus
and infertiliy in postpartum beff cattle. J Anim Sci. 1990;68:
799e816. https://doi.org/10.2527/1990.683799x.
11. Yavas Y, Walton JS. Postpartum acyclicity in suckled
beef cows: a review. Theriogenology. 2000;54(1):25e55.
https://doi.org/10.1016/S0093-691X(00)00323-X.
12. Day ML. Hormonal induction of estrous cycles in
anestrous Bos taurus beef cows. Anim Reprod Sci. 2004;
I. Beef cattle production
82e83:487e494. https://doi.org/10.1016/j.anireprosci.
2004.05.002.
13. Stevenson JS, Lamb GC, Johnson SK, et al. Supplemental
norgestomet, progesterone, or melengestrol acetate in-
creases pregnancy rates in suckled beef cows after timed
inseminations. J Anim Sci. 2003;81(3):571e586.
14. Lamb GC, Stevenson JS, Kesler DJ, Garverick HA,
Brown DR, Salfen BE. Inclusion of an intravaginal pro-
gesterone insert plus GnRH and prostaglandin F2a for
ovulation control in postpartum suckled beef cows.
J Anim Sci. 2001;79(9):2253e2259. https://doi.org/
10.2527/2001.7992253x.
15. Larson JE, Lamb GC, Stevenson JS, et al. Synchronization
of estrus and artificial insemination in replacement beef
heifers using gonadotropin-releasing hormone, prosta-
glandin F2a, and progesterone. J Anim Sci. 2006;84(11):
3000e3009. https://doi.org/10.2527/jas.2006-220.
16. Hill SL, Perry GA, Mercadante VRG, et al. Altered pro-
gesterone concentrations by hormonal manipulations
before a fixed-time artificial insemination CO-Synch þ
CIDR program in suckled beef cows. Theriogenology.
2014;82(1):104e113. https://doi.org/10.1016/j.therio
genology.2014.03.008.
17. Stevenson JS, Hill SL, Bridges GA, Larson JE, Lamb GC.
Progesterone status, parity, body condition, and days
postpartum before estrus or ovulation synchronization
in suckled beef cattle influence artificial insemination
pregnancy outcomes. J Anim Sci. 2015;93(5):
2111e2123. https://doi.org/10.2527/jas.2014-8391.
18. Hansen PJ. Physiological and cellular adaptations of zebu
cattle to thermal stress. Anim Reprod Sci. 2004;82e83:
349e360. https://doi.org/10.1016/j.anireprosci.2004.
04.011.
19. Randel RD. Unique Reproductive Traits of Brahman
and Brahman Based Cows. In: 39th Annual Beef Cattle
Short Course. 1990:60e82.
20. Sartori R, Gimenes LU, Monteiro PLJ, Melo LF,
Baruselli PS, Bastos MR. Metabolic and endocrine differ-
ences between Bos taurus and Bos indicus females that
impact the interaction of nutrition with reproduction.
Theriogenology. 2016;86(1):32e40. https://doi.org/
10.1016/j.theriogenology.2016.04.016.
21. Nahms. Usda. Beef 2007-08. Management. 2009
(February):Part IV.
22. Lamb GC, Mercadante VRG, Henry DD, et al. Invited
Review: advantages of current and future reproduc-
tive technologies for beef cattle production. Prof Anim
Sci. 2016;32(2):162e171. https://doi.org/10.15232/
pas.2015-01455.
23. Rodgers JC, Bird SL, Larson JE, et al. An economic
evaluation of estrous synchronization and timed artifi-
cial insemination in suckled beef cows. J Anim Sci.
2012;90(11):4055e4062. https://doi.org/10.2527/jas.
2011-4836.
 References
24. Leupp JL, Lardy GP, Daly R, Wright CL, Paterson JA.
Factors influencing price of North Dakota, South
Dakota and Montana feeder calves. NDSU Beef Cattle
Range Res Rep. 2008:46e49.
25. Foote RH. The history of artificial insemination: selected
notes and notables. J Anim Sci. 2002;80(E-suppl_2):1e10.
26. NAAB. reportAnnual Reports of Semen Sales and
Custom Freezing. Semen Sales.
27. Lamb GC, Dahlen CR, Larson JE, Marquezini G,
Stevenson JS. Control of the estrous cycle to improve
fertility for fixed-time artificial insemination in beef cat-
tle: a review. J Anim Sci. 2010;88(13 suppl l). https://
doi.org/10.2527/jas.2009-2349.
28. Harris DL, Newman S. Breeding for profit: synergism
between genetic improvement and livestock production
(A review). J Anim Sci. 1994;72:2178e2200.
29. Pruzzo L, Cantet RJC, Fioretti CC. Risk-adjusted ex-
pected return for selection decisions. J Anim Sci. 2003;
81(12):2984e2988.
30. Lamb GC, Larson JE, Geary TW, et al. Synchronization
of estrus and artificial insemination in replacement
beef heifers using gonadotropin-releasing hormone,
prostaglandin F2a, and progesterone. J Anim Sci.
2006;84(11):3000e3009. https://doi.org/10.2527/jas.
2006-220.
31. Lamb GC, Mercadante VRG. Synchronization and arti-
ficial insemination strategies in beef cattle. Vet Clin
North Am - Food Anim Pract. 2016;32(2):335e347.
https://doi.org/10.1016/j.cvfa.2016.01.006.
32. National Animal Health Monitoring System - United
States Department of Agriculture. Dairy Herd and Man-
agement Practices Ob U. S. Dairy Operations. 2014.
33. Macmillan KL, Thatcher WW. Effects of an agonist of
gonadotropin-releasing hormone on ovarian follicles
in cattle. Biol Reprod. 1991;45(6):883e889. https://
doi.org/10.1095/biolreprod45.6.883.
34. Pursley JR, Mee MO, Wiltbank MC. Synchronization of
ovulation in dairy cows using PGF2a and GnRH.
Theriogenology. 1995;44(7):915e923. https://doi.org/
10.1016/0093-691X(95)00279-H.
35. Burke JM, De La Sota RL, Risco CA, Staples CR,

Thatcher WW. Evaluation of timed insemi-
Schmitt EJP,
nation using a gonadotropin-releasing hormone agonist
in lactating dairy cows. J Dairy Sci. 1996;79:1385e1393.
https://doi.org/10.3168/jds.S0022-0302(96)76496-2.
36. Pursley JR, Kosorok MR, Wiltbank MC. Reproductive
management of lactating dairy cows using synchr-
onization of ovulation. J Dairy Sci. 1997;80(2):301e306.
https://doi.org/10.3168/jds.S0022-0302(97)75938-1.
37. Pursley JR, Wiltbank MC, Stevenson JS, Ottobre JS,
Garverick HA, Anderson LL. Pregnancy rates per artifi-
cial insemination for cows and heifers inseminated at a
synchronized ovulation or synchronized estrus. J Dairy
Sci. 1997;80(2):295e300. https://doi.org/10.3168/jds.
S0022-0302(97)75937-X.
I. Beef cattle production
71
38. Bisinotto RS, Ribeiro ES, Santos JEP. Synchronisation of
ovulation for management of reproduction in dairy
cows. Animal. 2014;8(SUPPL. 1):151e159. https://
doi.org/10.1017/S1751731114000858.
39. Geary TW, Whittier JC, Hallford DM, MacNeil MD. Calf
removal improves conception rates to the Ovsynch and
CO-Synch protocols. J Anim Sci. 2001;79(1):1e4.
https://doi.org/10.2527/2001.7911.
40. Johnson SK. Possibilities with today’s reproductive
technologies. Theriogenology. 2005;64(3):639e656. https://
doi.org/10.1016/j.theriogenology.2005.05.033.
41. Johnson SK, Funston RN, Hall JB, et al. Multi-state beef
reproduction Task Force provides science-based recom-
mendations for the application of reproductive
technologies. J Anim Sci. 2011;89(9):2950e2954.
https://doi.org/10.2527/jas.2010-3719.
42. Parcell JL, Dhuyvetter KC, Patterson DJ, Randle R. The
value of heifer and calf characteristics in bred heifer
price. Prof Anim Sci. 2006;22(3):217e224. https://
doi.org/10.15232/S1080-7446(15)31097-4.
43. Funston RN, Musgrave JA, Meyer TL, Larson DM. Ef-
fect of calving distribution on beef cattle progeny
performance. J Anim Sci. 2012;90:5118e5121. https://
doi.org/10.2527/jas2012-5263.
44. Cushman RA, Kill LK, Funston RN, Mousel EM,
Perry GA. Heifer calving date positively influences
calf weaning weights through six parturitions. J Anim
Sci. 2013;91(9):4486e4491. https://doi.org/10.2527/
jas2013-6465.
45. Moss GE, Crowder ME, Nett TM. GnRH-receptor inter-
action. VI. Effect of progesterone and estradiol on hypo-
physeal receptors for GnRH, and serum and
hypophyseal concentrations of gonadotropins in ovari-
ectomized ewes. Biol Reprod. 1981;25(5):938e944.
https://doi.org/10.1095/biolreprod25.5.938.
46. Wiltbank MC, G€ umen A, Sartori R. Physiological
classification of anovulatory conditions in cattle.
Theriogenology. 2002;57(1):21e52. https://doi.org/
10.1016/S0093-691X(01)00656-2.
47. Williams GL. Suckling as a regulator of postpartum
rebreeding in cattle: a review. J Anim Sci. 1990;68(3):
831e852.
48. Baruselli PS, Reis EL, Marques MO, Nasser LF, B
o GA.
The use of hormonal treatments to improve reproduc-
tive performance of anestrous beef cattle in tropical
climates. Anim Reprod Sci. 2004;82e83:479e486.
https://doi.org/10.1016/j.anireprosci.2004.04.025.
49. Nogueira GP. Puberty in south American Bos indicus
(zebu) cattle. 2004;83:361e372. https://doi.org/
10.1016/j.anireprosci.2004.04.007.
50. Cardoso RC, Alves BRC, Williams GL. Neuroendocrine
signaling pathways and the nutritional control of
puberty in heifers. Anim Reprod. 2018;15(Suppl. 1):
868e878. https://doi.org/10.21451/1984-3143-AR2018-
0013.
72 4. Reproductive management of beef cattle
51. J

unior IC, Sa Filho OG, Peres RFG, Aono FHS, Day ML.
Reproductive performance of prepubertal Bos indicus
heifers after progesterone-based treatments.
Theriogenology. 2010;74(6):903e911. https://doi.org/
10.1016/j.theriogenology.2010.04.015.
52. Rodrigues ADP, Peres RFG, Lemes AP, et al.
Progesterone-based strategies to induce ovulation in
prepubertal Nellore heifers. Theriogenology. 2013;79(1):
135e141. https://doi.org/10.1016/j.theriogenology.
2012.09.018.
53. Rodrigues ADP, Peres RFG, Lemes AP, et al. Effect of in-
terval from induction of puberty to initiation of a timed
AI protocol on pregnancy rate in Nellore heifers.
Theriogenology. 2014;82(5):760e766. https://doi.org/
10.1016/j.theriogenology.2014.06.008.
54. Meneghetti M, Filho OGS, Peres RFG, Lamb GC,
Vasconcelos JLM. Fixed-time artificial insemination
with estradiol and progesterone for Bos indicus cows I:
basis for development of protocols. Theriogenology.
2009;72(2):179e189. https://doi.org/10.1016/j.therio
genology.2009.02.010.
55. Peres RFG, J
unior IC, Filho OGS, Nogueira GP,
Vasconcelos JLM. Strategies to improve fertility in Bos
indicus postpubertal heifers and nonlactating cows sub-
mitted to fixed-time artificial insemination.
Theriogenology. 2009;72(5):681e689. https://doi.org/
10.1016/j.theriogenology.2009.04.026.
56. Cardoso RC, Alves BRC, Prezotto LD, et al. Use of a
stair-step compensatory gain nutritional regimen to
program the onset of puberty in beef heifers. J Anim
Sci. 2014;92(7):2942e2949. https://doi.org/10.2527/
jas.2014-7713.
57. Day ML, Nogueira GP. Management of age at puberty
in beef heifers to optimize efficiency of beef
production. Anim Front. 2013;3:6e11. https://doi.org/
10.2527/af.2013-0027.
58. Oosthuizen N, Fontes PLP, Sanford CD, et al.
Estrus synchronization and fixed-time artificial
insemination alter calving distribution in Bos indicus
influenced beef heifers. Theriogenology. 2018;106:
210e213. https://doi.org/10.1016/j.theriogenology.
2017.10.028.
59. Yelich JV, Bridges GA. Synchronization response: Bos
taurus vs. Bos indicus cattle. Spring. 2012;1:1e12.
60. Randel RD. Seasonal effects on female reproductive
functions in the bovine (Indian breeds). Theriogenology.
1984;21(1):170e185. https://doi.org/10.1016/0093-
691X(84)90315-7.
61. Carvalho JBP, Carvalho NAT, Reis EL, Nichi M,
Souza AH, Baruselli PS. Effect of early luteolysis in
progesterone-based timed AI protocols in Bos indicus,
Bos indicus  Bos taurus, and Bos taurus heifers.
Theriogenology. 2008;69(2):167e175. https://doi.org/
10.1016/j.theriogenology.2007.08.035.
I. Beef cattle production
62. Scarpa JO, O’Neil MM, Cardoso RC, Stanko RL,
Williams GL. Follicle dynamics and fertility at fixed-
time AI of Bos indicus-influenced beef cows synchronized
with the 5-Day Bee Synch þ Cidr protocol with or
without GnRH on day 0. J Anim Sci. 2017;95(suppl_4):
234. https://doi.org/10.2527/asasann.2017.479, 234.
63. Willett EL, Black WG, Casida LE, Stone WH,
Buckner PJ. Successful transplantation of a fertilized
bovine ovum. Science. 1951;113:247.
64. Betteridge KJ. A history of farm animal embryo transfer
and some associated techniques. Anim Reprod Sci. 2003;
(79):203e244. https://doi.org/10.1016/S0378-4320(03)
00166-0.
65. American Embryo Transfer Association. reportAnnual
Report of the AETA Statistics Committee for Calendar
Year 2017.
66. Leibo SP. Commercial production of pregnancies from
One-Step diluted frozen-thawed bovine embryos.
Theriogenology. 1986;25(1):166.
67. Sreenan JM, Diskin MG. Factors affecting pregnancy
rate following embryo transfer in the cow.
Theriogenology. 1987;27(1):99e113. https://doi.org/
10.1016/0093-691X(87)90073-2.
68. Lamb GC, Mercadante VRG. Selection & management
of the embryo recipient herd for embryo transfer. In:
Hopper RM, ed. Bovine Reproduction. 2014:723e732.
69. B
o GA, Baruselli PS, Moreno D, et al. The control of follic-
ular wave development for self-appointed embryo trans-
fer programs in cattle. Theriogenology. 2002;57:53e72.
70. Lamb GC, Mercadante VRG, Fontes PLP. Donor and
recipient management to optimize embryo technology suc-
cess. Applied Reprod Strategies Beef Cattle. 2016:197e209.
71. Betteridge KJ. Farm animal embryo technologies:
achievements and perspectives. Theriogenology. 2006;
65(5):905e913. https://doi.org/10.1016/j.therio
genology.2005.09.005.
72. Hasler JF. Forty years of embryo transfer in cattle: a re-
view focusing on the journal Theriogenology, the
growth of the industry in North America, and personal
reminisces. Theriogenology. 2014;81(1):152e169. https://
doi.org/10.1016/j.theriogenology.2013.09.010.
73. Palasz AT, Mapletoft RJ. Cryopreservation of mamma-
lian embryos and oocytes: recent advances. Biotechnol
Adv. 1996;14(2):127e149.
74. Brackett B, Bousquet D, Boice M, Donawick W, Evans J,
Dressel M. Normal development following in vitro
fertilization in the cow. Biol Reprod. 1982;1:147e158.
75. Bondioli KR. In vitro fertilization of bovine ooctytes by
spermatozoa capacitated in vitro. J Anim Sci. 1983;57(4):
1001e1005.
76. Leibfried-Rutledge ML, Critser ES, Eyestone WH,
Northey DL, First NL. Development potential of bovine
oocytes matured in vitro or in vivo. Biol Reprod. 1987;36:
376e383.
 References
77. Parrish JJ, Susko-Parrish JL, Leibfried-Rutledge ML,
Critser ES, Eyestone WH, First NL. Bovine in vitro fertil-
ization with frozen-thawed semen. Theriogenology. 1986;
25(4):591e600.
78. Greve T, Madison V. In vitro fertilization in cattle: a
review. Reprod Nutr Dev. 1991;31:147e157.
79. Seidel GE. Update on sexed semen technology in cattle.
Animal. 2014;8(s1):160e164. https://doi.org/10.1017/
S1751731114000202.
80. Johnson LA, Flook JP, Hawk HW. Sex preselection in
rabbits: live births from X and Y sperm separated by
DNA and cell sorting. Biol Reprod. 1989;41:199e203.
81. Seidel GE, Allen CH, Johnson LA, et al. Uterine horn
insemination of heifers with very low numbers of
nonfrozen and sexed spermatozoa. Theriogenology.
1997;48(8):1255e1264. https://doi.org/10.1016/S0093-
691X(97)00368-3.
82. DeJarnette JM, Nebel RL, Marshall CE. Evaluating the
success of sex-sorted semen in US dairy herds from
on farm records. Theriogenology. 2009;71(1):49e58.
https://doi.org/10.1016/j.theriogenology.2008.09.042.
83. Seidel GE. Overview of sexing sperm. Theriogenology.
2007;68(3):443e446. https://doi.org/10.1016/j.therio
genology.2007.04.005.
I. Beef cattle production
73
84. Carvalho JO, Sartori R, Machado GM, Mour~ao GB,
Dode MAN. Quality assessment of bovine cryopre-
served sperm after sexing by flow cytometry and their
use in in vitro embryo production. Theriogenology.
2010;74(9):1521e1530. https://doi.org/10.1016/
j.theriogenology.2010.06.030.
85. Pellegrino CAG, Morotti F, Untura RM, et al. Use
of sexed sorted semen for fixed-time artificial insem-
ination or fixed-time embryo transfer of in vitroe
produced embryos in cattle. Theriogenology. 2016;86(3):
888e893. https://doi.org/10.1016/j.theriogenology.
2016.03.010.
86. Thomas JM, Lock SL, Poock SE, Ellersieck MR,
Smith MF, Patterson DJ. Delayed insemination of nones-
trous cows improves pregnancy rates when using sex-
sorted semen in timed artificial insemination of suckled
beef cows. J Anim Sci. 2014;92(4):1747e1752. https://
doi.org/10.2527/jas2013-7131.
87. Thomas JM, Locke JWC, Vishwanath R, et al. Effective
use of SexedULTRATM sex-sorted semen for timed arti-
ficial insemination of beef heifers. Theriogenology. 2017;
98:88e93. https://doi.org/10.1016/j.theriogenology.
2017.03.018.