RESEARCH PAPER
https://doi.org/10.1071/AN22074
Conception rate according to antral follicle count and oestrus
expression in Nelore cows submitted to timed artificial
insemination
Fábio Lucas Zito de Moraes A, Denis Vinicius Bonato A, Ana Clara Canto Souza A
Marcelo Marcondes Seneda A,*
For full list of author affiliations and
declarations see end of paper
*Correspondence to:
Marcelo Marcondes Seneda
State University of Londrina, Laboratory of
Animal Reproduction (REPROA), DCV,
CCA, UEL, Cep: 86057-970, Londrina,
PR, Brazil
Email: marcelo.seneda@uel.br
Handling Editor:
Rodolfo Ungerfeld
Received: 14 May 2021
Accepted: 30 April 2022
Published: 10 June 2022
Cite this:
Moraes FLZ et al. (2022)
Animal Production Science, 62(13),
1229–1235.
doi:10.1071/AN22074
© 2022 The Author(s) (or their
employer(s)). Published by
CSIRO Publishing.
, Fábio Morotti A and
ABSTRACT
Context. Antral follicle count (AFC) has been related to bovine fertility and response to
reproductive biotechnologies, but its relationship to oestrus expression during timed artificial
insemination (TAI) is not known. Aims. This study evaluated the relationship between the AFC
and oestrus expression in regard to the conception rate in beef cows that received an ovulation
synchronisation protocol for TAI. Methods. Multiparous Nelore cows up to 40 days
postpartum were evaluated by an ultrasound to determine the AFC, and were classified into
low (≤15 follicles; n = 150), intermediate (≥19 and ≤26 follicles; n = 150) and high count
(≥30 follicles; n = 150). AFC evaluation was performed on a random day of the oestrous cycle
(Day 0). The animals received a conventional TAI protocol based on progesterone and oestradiol
benzoate. On Day 8, the animals received cloprostenol, equine chorionic gonadotrophin and
oestradiol cypionate, in addition to a painting in the sacrococcygeal region. On Day 10, TAI was
performed, and oestrus expression was evaluated and classified as high, low or absent, according
to the removal of paint. Pregnancy diagnosis was performed by ultrasound 30 days after TAI.
The data were analysed using a generalised linear model and a binary logistic regression model
(P < 0.05). Key results. Regardless of AFC, a high proportion of cows showed high oestrus
expression (low 61.3%; intermediate 67.3% and high 58.7%) at the end of the TAI protocol. The
conception rate was influenced by AFC (low 69.3% vs high 56.7%; P = 0.04) and oestrus
expression (high oestrus: 67.3% vs no oestrus: 45.4%; P = 0.003). In addition, an interaction
(P = 0.008) between AFC and oestrus expression showed that low AFC cows and high oestrus
expression had the highest conception rate to TAI. Conclusion. AFC and oestrus expression
classification influenced the conception rate of Nelore cows in the TAI programme. In addition,
the interaction of AFC with oestrus expression revealed that the best conception rate was
observed in females with low AFC that expressed high-intensity oestrus. Implications. The
associated assessment of AFC and oestrus expression represents a strategy to increase the
conception rate in TAI programmes in cattle.
Keywords: antral follicle count, Bos indicus, conception rate, fertility, oestrus expression,
synchronisation, timed artificial insemination, ultrasound.
Introduction
The antral follicle count (AFC) has shown considerable importance in studies on fertility in
cattle. Many physiological and productive measures are associated with AFC, with
influence on biotechnique efficiency, such as artificial insemination (Mossa et al. 2012;
Martinez et al. 2016), timed artificial insemination (TAI; Morotti et al. 2018; de Moraes
et al. 2019; de Lima et al. 2020), and embryo in vivo and in vitro production (Ireland
et al. 2007; Pontes et al. 2009; Silva-Santos et al. 2014; Santos et al. 2016; Garcia et al.
2020). In this context, a full understanding of the relationship between AFC and fertility
F. L. Z. Moraes et al.
has become a matter of high relevance (Silva-Santos et al.
2014; Morotti et al. 2017a, 2017b; Zangirolamo et al. 2018;
Garcia et al. 2020).
The AFC is a highly variable reproductive trait among
females. However, it has high individual repeatability,
even if evaluated on a random day of the oestrous cycle
(Burns et al. 2005; Ireland et al. 2008; Cushman et al.
2009). Thus, AFC evaluation has become an interesting and
strategic tool for classifying cows in a herd into groups with
low, intermediate and high counts, with only an ultrasound
examination (Ireland et al. 2007; Silva-Santos et al. 2014;
Morotti et al. 2017b, 2018; Zangirolamo et al. 2018; de
Moraes et al. 2019; Bomtempo et al. 2020; de Lima et al.
2020). This classification is useful for assessing reproductive
performance or as a tool to guide the strategic use of
reproductive biotechnology (Morotti et al. 2017a, 2017b;
Zangirolamo et al. 2018; Seneda et al. 2019).
Initially, the low AFC in Bos taurus females was related
to low fertility characteristics. This includes reduced
capacity for in vitro embryo production, reduced response
to superovulatory treatment for in vivo embryo production
and less viable embryo production (Singh et al. 2004;
Ireland et al. 2007). In addition, a lesser probability of
pregnancy at the end of the reproductive season (Mossa
et al. 2012), low concentrations of circulating progesterone
(P4) and anti-Müllerian hormone (Evans et al. 2012;
Jimenez-Krassel et al. 2015), lesser endometrial thickness
(Jimenez-Krassel et al. 2009), and small ovaries (Ireland
et al. 2008) have also been reported in cows with low AFC.
In contrast, such findings do not seem to be a consensus,
and several studies in Bos indicus cattle have shown that
low AFC resulted in a greater conception rate at TAI
(Morotti et al. 2015, 2017b, 2018; de Moraes et al. 2019;
Seneda et al. 2019; de Lima et al. 2020). This included a
larger diameter in the dominant follicle (Morotti et al.
2018), larger corpus luteum and greater P4 concentration
(de Lima et al. 2020), and greater productive longevity/in
Bos indicus cattle (Jimenez-Krassel et al. 2017). In addition,
low AFC also showed favourable expression of genes
considered important for follicular development, oocyte
competence and female fertility (de Lima et al. 2020).
In the fertility context, it has been shown that even in TAI
protocols, which aim at insemination at a prefixed time
without the need for oestrus observation, females that
express oestrus during TAI have a greater pregnancy rate
than those that do not (Bó and Cedeno ˜ 2018; Nogueira
et al. 2019; Rodrigues et al. 2019). In addition, the largest
diameter of the dominant follicle in TAI is positively related
to oestrus expression (Sá Filho et al. 2010), ovulatory
potential (Gimenes et al. 2008) and greater pregnancy rates
(Meneghetti et al. 2009; Sá Filho et al. 2010; Pfeifer et al.
2012, 2015). Therefore, to strategically take advantage of
this fertility, oestrus evaluation with the aid of painting at
the base of the tail has become a routine during TAI
programmes (Nogueira et al. 2019; Rodrigues et al. 2019).
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Animal Production Science
The association between AFC and oestrus expression in TAI
has not yet been studied, and it is possible that these factors
can interact and have some impact on fertility. Therefore,
our study hypothesised that there is a positive relation-
ship between low AFC and oestrus, resulting in better
pregnancy rates.
Materials and methods
Location and animals
The study was conducted during the typical time of breeding
on a commercial farm of Bos taurus indicus cattle. This farm is
located in the south-western region of Brazil, latitude
22°11 0 43.9″S and longitude 48°30 0 25.7″W. The climate in
this region is tropical, with wet and dry seasons, an average
temperature of 22.1°C, an average annual rainfall of
1200–1400 mm and an altitude of 325 m.
A total of 470 Nelore females ranging in age from 36 to
96 months, newly calving (<40 days postpartum), bred on
extensive grazing system, eating pasture of Brachiaria brizantha
and supplied with mineral salt ad libitum were used. Animals
had an average body condition score (BCS) of 3.04 ± 0.03,
ranging from 2.50 to 4.25 (scale 1–5; Lowman et al. 1976).
All animals received a standard ovulation synchronisation
protocol for TAI, which was initiated on a random day of the
oestrous cycle (Day 0). On the same day, the BCS and AFC
were evaluated by a single experienced veterinarian.
Experimental design and antral follicle count
The AFC of each animal was determined at the beginning of
the TAI protocol (Day 0) throughout the breeding season. This
was performed through ultrasound evaluation using a 5-MHz
transrectal linear transducer (Aloka SSD-500, Aloka Co. Ltd.,
Tokyo, Japan). Both ovaries were evaluated, and the total
number of antral follicles (≥3 mm in diameter) was estimated
per female, as described by Burns et al. (2005) and Ireland
et al. (2008). To standardise the antral follicle count, each
face of the ovary was analysed from the end to the pedicle.
This aimed to identify the number of antral follicles present
in each gonad with greater precision, as described by
Morotti et al. (2018). According to their ovarian follicular
population, the cows were divided for analysis into groups
of high, intermediate and low AFC (Fig. 1).
Hormonal protocol and oestrus expression
evaluation
The beginning of the protocol (Fig. 2) was established on
random days of the oestrous cycle (Day 0) with the inser-
tion of an intravaginal device containing 1.2 g progesterone
(P4; FertilCare®1200; Vallée, Brazil). This was associated
with the application of 2.0 mg estradiol benzoate (RIQ-BE®,
www.publish.csiro.au/an Animal Production Science

90 n = 470 maintained stable (defroster semen and embryos, WTA®;

Mean = 25.8 WTA, Craven, SP, Brazil) set at 36°C for 30 s.
80 s.d. = 16.5
On Day 8, the sacrococcygeal region was painted with
70 1st quartile = 15 (3−15)
Median = 20 (16−28) a red stick (All-Weather®; Paint Stick, EUA) to assess
Frequency

60
3rd quartile = 30 (30−100)
50
40
30
20
10
0
0 15 30 45 60 75 90
AFC on Day 0
Fig. 1. Distribution frequency of the antral follicle count (AFC),
mean, standard deviation (s.d.), first quartile, median and third
quartile from animals monitored in this study.
Agener União, Brazil) intramuscularly (i.m.). After 8 days
(Day 8), the P4 devices were removed. This was followed
by intramuscular administration of 450 μg of sodium
cloprostenol (Estron®; Agener União, Brazil), 300 IU of
equine chorionic gonadotropin (Folligon®; MSD Animal
Health, São Paulo, Brazil) and 1.0 mg estradiol cypionate
(Fertilcare® Ovulation; Vallée). Artificial insemination was
performed 48 h after the removal of the intravaginal P4
device (Day 10) using cryopreserved semen from three
bulls with previously known fertility, and equally distributed
among the three treatments (low, intermediate and high
AFC). Semen doses were acquired at commercial centres
that follow the quality criteria established by the Brazilian
College of Animal Reproduction (CBRA 2013). For thawing
of the semen, an automatic defroster temperature that
oestrus expression, following the methodology described
by Nogueira et al. (2019), as shown in Fig. 2. On Day 10,
during TAI, the cows were evaluated and classified according
to the expression of oestrus in: I (no oestrus), paint was not
removed; II (low oestrus), paint was partially removed; and
III (high oestrus), paint was completely removed.
Pregnancy diagnosis
For the diagnosis of pregnancy, the cows were subjected to a
single transrectal ultrasound with a 5-MHz transrectal linear
transducer (Aloka SSD-500; Aloka Co. Ltd., Tokyo, Japan)
30 days after artificial insemination. The pregnancy was
confirmed by visualisation of a viable conceptus, with the
presence of a heartbeat. To calculate the conception rate,
the total number of pregnant cows was divided by the
number of inseminated cows.
Statistical analysis
For data analysis, the first quartile (Q1, 15 follicles), the
second quartile/median (Q2, 20 follicles) and the third
quartile (Q3, 30 follicles) were extracted from the antral
follicle population of 470 cows, as proposed by Droher
et al. (2021). Subsequently, the AFC groups were defined as
low (Q1 ≤15 follicles), intermediate (cows with AFC ≥19
and ≤26 follicles) or high count (Q3 ≥30 follicles). Animals
with AFC at the interface of the referred groups were
excluded from the statistical analysis to better categorise

AFC evaluation Estrus expression

I II III

Low Intermediate High

No estrus Low estrus High estrus
(≤15 follicles) (≥19 ≤ 26 follicles) (≥30 follicles)
(n = 150) (n = 150) (n = 150)

300 IU eCG
450 μg PGF2α
1 mg EC
+ Pregnancy
2 mg EB Tail painting TAI diagnosis

P4 device

Day 0 Day 8 Day 10 Day 40

Fig. 2. Experimental design and hormonal protocol used in Nelore cows to evaluate the relationship of antral
follicle count (AFC) and oestrus expression with the conception rate in a timed artificial insemination (TAI)
programme. At the time of artificial insemination, cows with low, intermediate and high AFC were classified
as follows: I (no oestrus expression), paint was not removed; II (low oestrus expression), paint partially
removed; and III (high oestrus expression), paint completely removed.

1231
F. L. Z. Moraes et al.
the groups, as proposed by Morotti et al. (2018). From a total
of 470 cows, we excluded 20, and we considered the final
number of 450 cows. The excluded animals (n = 20) were
not included, because they presented a number of follicles
close to the cutoff of each category. For the cut-off between
the low and intermediate AFC groups, those animals
(n = 11) with 16, 17 or 18 antral follicles were not
considered. For the interface of intermediate and high AFC
groups, we excluded the cows (n = 9) with 27, 28 or
29 antral follicles. This strategy was adopted to improve
the accuracy of the establishment of AFC groups. Because
we worked with many animals under field conditions, an
error of one or two follicles at the counting moment was
possible. Considering the exclusion of those 20 animals, we
are confident that no errors occurred at the classification of
the animals as low, intermediate or high AFC.
The number of antral follicles was analysed by ANOVA
using a generalised linear model. The main effect of the
AFC group (low, intermediate and high) was considered as
the fixed effect, and BCS was included as the covariate in
the model. For significance in ANOVA, Tukey’s test was
used as a post-hoc mean test. The TAI conception rate was
analysed using a binary logistic regression model, including
the main effect of the AFC groups (low, intermediate and
high), oestrus expression (no, low and high), interactions
(AFC × oestrus) and bulls as fixed variables. The BCS of
cows was included as a continuous covariate in the model.
The bull had no significant effect (P > 0.1) and was
removed from the final model. In the presence of a
significant effect, the oestrus and conception rates were
compared using the Chi-squared test to rank the categories.
For descriptive analysis, quantitative data are presented as
the mean (M), standard error (s.e.) and qualitative data as a
percentage. All statistical analyses were performed using
Minitab® statistical programme, version 18.1. A significance
level of 5% was used to indicate the effect of categorical
variables and their interactions.
Results
In general, the highest proportion of cows in this study
showed a high oestrus expression (62.4%) at the end of the
Table 1. Proportion of Nelore cows with low, intermediate and high antral follicle count that presented no, low or high oestrus expression during
the timed artificial insemination protocol.
Group Animals (n)
Low AFC 150
Intermediate AFC 150
High AFC 150
Total/mean 450
a and b in the same column indicates a statistical difference (P ≤ 0.05) among the AFC groups.
A–C on the same line indicates a statistical difference (P ≤ 0.05) among oestrus expression classifications.
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Animal Production Science
TAI protocol, regardless of the AFC group evaluated
(Table 1). Females with low, intermediate and high AFC
showed similar (P > 0.1) proportions of oestrus expression,
classified as low or high intensity.
The conception rate was influenced by AFC (P = 0.04)
and oestrus expression (P = 0.003; Table 2), with low AFC
cows having greater conception rates compared with cows
with high AFC (69.3% vs 56.7%, respectively). Cows with
intermediate AFC had intermediate conception rates. Cows
with high oestrus expression also had greater conception
rates in relation to cows with no oestrus expression (67.3%
vs 45.4%, respectively). The BCS was similar (P > 0.1)
among groups with low (3.0 ± 0.1), intermediate (3.0 ± 0.1)
and high AFC (3.1 ± 0.1). Neither BCS nor the bull affected
(P > 0.05) the conception rate.
In addition to the isolated effects, an interaction
(P = 0.008) between AFC and oestrus expression on the
conception rate to TAI was observed. This interaction showed
that low AFC cows with high oestrus expression had a greater
conception rate than intermediate and high AFC cows also
classified with high oestrus expression (Table 3).
Discussion
The present study presents practical and applicable
information regarding the relationship between AFC, estrus
expression and conception rates in Bos indicus cattle
submitted to the ovulation synchronisation programme for
TAI. Our results indicate that both AFC and oestrus
expression can affect pregnancy by TAI. We demonstrated
that cows with low AFC had a greater conception rate at
TAI than high AFC cows, and cows that demonstrated greater
oestrus expression had greater conception rates than cows
that did not exhibit oestrus. We emphasise that this is the
first study that relates oestrus expression with AFC and
conception to TAI. Therefore, our findings contribute to a
better understanding of the relationship between AFC and
fertility. Also, we believe that our data may provide
adjustments of the management to TAI, considering the
best result in low AFC Bos indicus females according to
oestrus expression.
No oestrus % (n) Low oestrus % (n) High oestrus % (n)
16.7aB (25) 22.0aB (33) 61.3aA (92)
8.7bC (13) 24.0aB (36) 67.3aA (101)
18.7aB (28) 22.7aB (34) 58.7aA (88)
14.7 (66) 22.9 (103) 62.4 (281)
www.publish.csiro.au/an Animal Production Science

Table 2. Relationship of antral follicle count (low, intermediate and (Jimenez-Krassel et al. 2009, 2015), reduced response to
high) and oestrus expression (no, low and high) with the conception superovulatory treatment, fewer viable embryos produced
rate in Bos taurus indicus cows (Nelore) submitted to the timed in vivo and in vitro (Singh et al. 2004; Ireland et al.
artificial insemination protocol. 2007, 2011), small ovaries (Ireland et al. 2008), reduced
Groups Animals AFC Conception endometrial thickness during estrus (Jimenez-Krassel et al.
(n) (m ± s.e.) rate % (n) 2009), and a lesser probability of pregnancy until the end
AFC of the reproductive period (Mossa et al. 2012). However,
Jimenez-Krassel et al. (2017) found that the low AFC in a
Low 150 12.2 ± 0.2c 69.3a (104)
dairy herd (Holstein) led to greater fertility and greater
Intermediate 150 21.9 ± 0.2b 60.0ab (90)
productive longevity compared with females with high
High 150 44.7 ± 1.3a 56.7b (85) AFC. Therefore, our findings in Zebu animals are similar to
P-value – <0.0001 0.04 the data found in Holstein cows reported by Jimenez-Krassel
Oestrus et al. (2017).
High 281 26.3 ± 1.0a 67.3a (189) Another relevant finding in our study was the greater
Low 103 24.5 ± 1.3a 58.2ab (60)
conception rate in cows that had a high intensity of oestrus
expression compared with animals that had absent or low
No 66 29.1 ± 2.4a 45.4b (30)
expression oestrus. Similarly, several studies reported a
P-value – 0.22 0.003 greater conception rate in bovine females that demonstrated
a and b in the same column indicates a statistical difference (P ≤ 0.05) among the oestrus in response to the TAI protocol (Sá Filho et al. 2010;
AFC groups. ˜ 2018; Nogueira et al.
Baruselli et al. 2017; Bó and Cedeno
2019). In a study of 445 newly calving Nelore cows, the
pregnancy rate was greater in cows that had oestrus detected
In the present study, cows with low AFC (≤15 follicles) (67%; 174/257) than in cows that did not express oestrus
(36.2%; 68/188; Sá Filho et al. 2010).
had a greater conception rate at 30 days post-artificial
Our data agree with the findings of Morotti et al. (2018),
insemination than those with intermediate and high counts.
who reported larger diameters of dominant and preovu-
Similar results were found by Morotti et al. (2018), who
latory follicles to TAI. Similarly, de Lima et al. (2020)
evaluated Bos indicus females classified as low (≤15 follicles;
showed not only the diameter of the dominant follicle, but
n = 243), intermediate (≥20 ≤ 40 follicles; n = 495) and high
also the size of the CL and the P4 concentration were
AFC (≥45 follicles; n = 194), with a greater conception rate in
greater in females with low AFC than in females with a
cows with low AFC (61.7%) than intermediate (52.9%) or high concentration. de Lima et al. (2020) also evaluated the
high AFC (49.5%). Similarly, in Bos indicus cows (n = 736), expression pattern of genes related to oocyte competence,
de Moraes et al. (2019) found a greater conception rate in and proved that oocytes and cumulus cells of females with
low AFC cows (≤10 follicles; 57.7%), in comparison with low AFC were upregulated for intercellular communication,
intermediate (≥11 and ≤29 follicles; 49.7%) and high AFC meiotic control, epigenetic modulation, cell division,
(≥30 follicles; 47.9%). Therefore, in the present study, the follicular growth, cell maintenance, steroidogenesis and cell
data indicate that the lowest AFC in Bos taurus indicus stress. Therefore, based on the present study, we suggest
cattle is related to better conception rates for TAI. that in the Bos indicus beef herd subjected to TAI, greater
In contrast, the first studies in Bos taurus related pregnancy in low AFC cows may be positively related to
low AFC with characteristics of lesser fertility (Ireland et al. follicular diameters (Morotti et al. 2018) and also to the
2008, 2011; Mossa et al. 2012; Martinez et al. 2016). In high intensity of oestrus expression.
this context, low AFC was associated with low circulat- In contrast, in dairy cows (Bos taurus), Martinez et al.
ing concentrations of P4 and anti-Müllerian hormone (2016) reported a relationship between high AFC and

Table 3. Conception rate from the interaction between antral follicle count (low, intermediate and high) with estrus expression (no, low or high)
in Nelore cows subjected to the timed artificial insemination protocol.
Groups No oestrus % (n) Low oestrus % (n) High oestrus % (n) Total/mean % (n)
Low AFC 48.0aB (12/25) 66.7aAB (22/33) 76.1aA (70/92) 69.3 (104/150)
Intermediate AFC 53.8aAB (7/13) 47.2bB(17/36) 65.3bA (66/101) 60.0 (90/150)
High AFC 32.3aB (11/28) 61.8abA (21/34) 60.2bA (53/88) 56.7 (85/150)
Total/mean 45.4 (30/66) 58.2 (60/103) 67.2 (189/281) 62.0 (279/450)

a and b in the same column indicates a statistical difference (P ≤ 0.05) among the AFC groups.
A and B in the same line indicates a statistical difference (P ≤ 0.05) among oestrus expression classifications.

1233
F. L. Z. Moraes et al.
greater oestrus expression. However, in Brahman (Bos indicus)
females, Bomtempo et al. (2020), reported variation in the
oestrus expression intensity (94.7%a, 70.6%b and 84.9%ab),
but not in the conception rate (52.6%, 51.0% and 45.5%)
among low, intermediate and high AFC groups, respectively.
There is still no consensus on the relationship between AFC
and fertility across Bos taurus and Bos indicus females.
However, the present study makes it possible to suggest
management strategies that can contribute to a more
efficient use of reproductive biotechniques that may be
influenced by AFC number in Bos indicus cattle. AFC is a
characteristic of high repeatability in the same female
(Burns et al. 2005; Ireland et al. 2008; Silva-Santos et al.
2014; Morotti et al. 2018). The combination of AFC and
oestrus expression may be a strategy for adjustments in the
arificial insemination moment.
The relationship between AFC and oestrus expression was
clear in the present study. However, there are still many
questions that need to be clarified regarding the use of
AFC as a tool to adjust the performance of reproductive
programmes. For example, Morotti et al. (2018) suggested
that the dominant follicle size and the response to TAI
synchronisation programmes may be related to hormonal
signalling that may vary depending on the female’s AFC.
There is evidence that follicle-stimulating hormone concen-
trations are inversely associated with the number of
follicles >3 mm in diameter during follicular waves (Burns
et al. 2005).
Conclusion
In conclusion, both AFC and oestrus expression classification
influenced the conception rate of Nelore cows submitted to
the TAI programme. The low AFC cows with high intensity
of oestrus expression resulted in greater conception rates.
References
Baruselli PS, Ferreira RM, Colli MHA, Elliff FM, Sá Filho MF, Vieira L,
de Freitas BG (2017) Timed artificial insemination: current challenges
and recent advances in reproductive efficiency in beef and dairy herds
in Brazil. Animal Reproduction 14, 558–571. doi:10.21451/1984-3143-
AR999
Bó GA, Cedeño A (2018) Expression of estrus as a relevant factor in fixed-
time embryo transfer programs using estradiol/progesterone-based
protocols in cattle. Animal Reproduction 15, 224–230. doi:10.21451/
1984-3143-AR2018-0060
Bomtempo MES, Cerezetti MB, de Moraes FLZ, de Campos Silva RCV,
Lorenzetti E, Morotti F (2020) Relationship of conception rate with
antral follicle count, nutritional parameters and animal behavior in
Brahman cows submitted to TAI. Ensaios e Ciencias 24, 455–461.
doi:10.17921/1415-6938.2020v24n5-esp.p455-461
Burns DS, Jimenez-Krassel F, Ireland JLH, Knight PG, Ireland JJ
(2005) Numbers of antral follicles during follicular waves in cattle:
evidence for high variation among animals, very high repeatability
in individuals, and an inverse association with serum follicle-
stimulating hormone concentrations. Biology of Reproduction 73,
54–62. doi:10.1095/biolreprod.104.036277
1234
Animal Production Science
CBRA (2013) ‘Manual para exame andrológico e avaliação de sêmen
animal.’ (Colégio Brasileiro de Reprodução Animal)
Cushman RA, Allan MF, Kuehn LA, Snelling WM, Cupp AS, Freetly HC
(2009) Evaluation of antral follicle count and ovarian morphology
in crossbred beef cows: investigation of influence of stage of the
estrous cycle, age, and birth weight. Journal of Animal Science 87,
1971–1980. doi:10.2527/jas.2008-1728
de Lima MA, Morotti F, Bayeux BM, de Rezende RG, Botigelli RC, De Bem
THC, Fontes PK, Nogueira MFG, Meirelles FV, Baruselli PS, da Silveira
JC, Perecin F, Seneda MM (2020) Ovarian follicular dynamics,
progesterone concentrations, pregnancy rates and transcriptional
patterns in Bos indicus females with a high or low antral follicle
count. Scientific Reports 10, 19557. doi:10.1038/s41598-020-76601-5
de Moraes FLZ, Morotti F, Costa CB, Lunardelli PA, Seneda MM (2019)
Relationships between antral follicle count, body condition, and
pregnancy rates after timed-AI in Bos indicus cattle. Theriogenology
136, 10–14. doi:10.1016/j.theriogenology.2019.06.024
Droher RG, Cavalieri FLB, Morotti F, Zangirolamo AF, Andreazzi MA,
Seneda MM (2021) Increased antral follicle population and in vitro
embryo production in pregnant Holstein. Semina: Ciências Agrárias
42, 2851–2866. doi:10.5433/1679-0359.2021v42n5p2851
Evans ACO, Mossa F, Walsh SW, Scheetz D, Jimenez-Krassel F, Ireland
JLH, Smith GW, Ireland JJ (2012) Effects of maternal environment
during gestation on ovarian folliculogenesis and consequences for
fertility in bovine offspring. Reproduction in Domestic Animals
47(s4), 31–37. doi:10.1111/j.1439-0531.2012.02052.x
Garcia SM, Morotti F, Cavalieri FLB, Lunardelli PA, Santos AdO,
Membrive CMB, Castilho C, Puelker RZ, Silva JOF, Zangirolamo AF,
Seneda MM (2020) Synchronization of stage of follicle development
before OPU improves embryo production in cows with large
antral follicle counts. Animal Reproduction Science 221, 106601.
doi:10.1016/j.anireprosci.2020.106601
Gimenes LU, Sá Filho MF, Carvalho NAT, Torres-Júnior JRS, Souza AH,
Madureira EH, Trinca LA, Sartorelli ES, Barros CM, Carvalho JBP,
Mapletoft RJ, Baruselli PS (2008) Follicle deviation and ovulatory
capacity in Bos indicus heifers. Theriogenology 69, 852–858.
doi:10.1016/j.theriogenology.2008.01.001
Ireland JJ, Ward F, Jimenez-Krassel F, Ireland JLH, Smith GW, Lonergan
P, Evans ACO (2007) Follicle numbers are highly repeatable within
individual animals but are inversely correlated with FSH
concentrations and the proportion of good-quality embryos after
ovarian stimulation in cattle. Human Reproduction 22, 1687–1695.
doi:10.1093/humrep/dem071
Ireland JLH, Scheetz D, Jimenez-Krassel F, Themmen APN, Ward F,
Lonergan P, Smith GW, Perez GI, Evans ACO, Ireland JJ (2008)
Antral follicle count reliably predicts number of morphologically
healthy oocytes and follicles in ovaries of young adult cattle.
Biology of Reproduction 79, 1219–1225. doi:10.1095/biolreprod.
108.071670
Ireland JJ, Smith GW, Scheetz D, Jimenez-Krassel F, Folger JK, Ireland
JLH, Mossa F, Lonergan P, Evans ACO (2011) Does size matter in
females? An overview of the impact of the high variation in the
ovarian reserve on ovarian function and fertility, utility of anti-
Müllerian hormone as a diagnostic marker for fertility and causes of
variation in the ovarian reserve in cattle. Reproduction, Fertility and
Development 23, 1–14. doi:10.1071/RD10226
Jimenez-Krassel F, Folger JK, Ireland JLH, Smith GW, Hou X, Davis JS,
Lonergan P, Evans ACO, Ireland JJ (2009) Evidence that high
variation in ovarian reserves of healthy young adults has a negative
impact on the corpus luteum and endometrium during estrous
cycles in cattle. Biology of Reproduction 80, 1272–1281. doi:10.1095/
biolreprod.108.075093
Jimenez-Krassel F, Scheetz DM, Neuder LM, Ireland JLH, Pursley JR,
Smith GW, Tempelman RJ, Ferris T, Roudebush WE, Mossa F,
Lonergan P, Evans ACO, Ireland JJ (2015) Concentration of anti-
Müllerian hormone in dairy heifers is positively associated with
productive herd life. Journal of Dairy Science 98, 3036–3045.
doi:10.3168/jds.2014-8130
Jimenez-Krassel F, Scheetz DM, Neuder LM, Pursley JR, Ireland JJ (2017)
A single ultrasound determination of ≥25 follicles ≥3 mm in diameter
in dairy heifers is predictive of a reduced productive herd life. Journal
of Dairy Science 100, 5019–5027. doi:10.3168/jds.2016-12277
www.publish.csiro.au/an
Lowman BG, Scott N, Somerville S (1976) ‘Condition scoring of cattle.’
Revised edn. Bulletin No. 6. (East of Scotland College of Agriculture)
Martinez MF, Sanderson N, Quirke LD, Lawrence SB, Juengel JL (2016)
Association between antral follicle count and reproductive measures
in New Zealand lactating dairy cows maintained in a pasture-based
production system. Theriogenology 85, 466–475. doi:10.1016/
j.theriogenology.2015.09.026
Meneghetti M, Sá Filho OG, Peres RFG, Lamb GC, Vasconcelos JLM (2009)
Fixed-time artificial insemination with estradiol and progesterone for
Bos indicus cows I: basis for development of protocols. Theriogenology
72, 179–189. doi:10.1016/j.theriogenology.2009.02.010
Morotti F, Barreiros TRR, Machado FZ, González SM, Marinho LSR,
Seneda MM (2015) Is the number of antral follicles an interesting
selection criterium for fertility in cattle. Animal Reproduction 12,
479–486.
Morotti F, Santos GMG, Júnior CK, Silva-Santos KC, Roso VM, Seneda MM
(2017a) Correlation between phenotype, genotype and antral follicle
population in beef heifers. Theriogenology 91, 21–26. doi:10.1016/
j.theriogenology.2016.12.025
Morotti F, Zangirolamo AF, da Silva NC, da Silva CB, Rosa CO, Seneda MM
(2017b) Antral follicle count in cattle: advantages, challenges, and
controversy. Animal Reproduction 14, 514–520. doi:10.21451/1984-
3143-AR994
Morotti F, Moretti R, dos Santos GMG, Silva-Santos KC, Ramos Cerqueira
PH, Seneda MM (2018) Ovarian follicular dynamics and conception
rate in Bos indicus cows with different antral follicle counts
subjected to timed artificial insemination. Animal Reproduction
Science 188, 170–177. doi:10.1016/j.anireprosci.2017.12.001
Mossa F, Walsh SW, Butler ST, Berry DP, Carter F, Lonergan P, Smith GW,
Ireland JJ, Evans ACO (2012) Low numbers of ovarian follicles ≥3mm
in diameter are associated with low fertility in dairy cows. Journal of
Dairy Science 95, 2355–2361. doi:10.3168/jds.2011-4325
Nogueira E, Silva MR, Silva JCB, Abreu UPG, Anache NA, Silva KC,
Cardoso CJT, Sutovsky P, Rodrigues WB (2019) Timed artificial
insemination plus heat I: effect of estrus expression scores on
pregnancy of cows subjected to progesterone–estradiol-based
protocols. Animal 13, 2305–2312. doi:10.1017/S1751731119000442
Pfeifer LFM, Leal SdCBdS, Schneider A, Schmitt E, Corrêa MN (2012)
Effect of the ovulatory follicle diameter and progesterone
concentration on the pregnancy rate of fixed-time inseminated
lactating beef cows. Revista Brasileira de Zootecnia 41, 1004–1008.
doi:10.1590/S1516-35982012000400024
Pfeifer LFM, Castro NA, Melo VTO, Neves PMA, Cestaro JP, Schneider A
(2015) Timed artificial insemination in blocks: a new alternative to
Data availability. The data that support this study will be shared upon reasonable request to the corresponding author.
Conflicts of interest. The authors declare no conflicts of interest.
Declaration of funding. This research did not receive any specific funding.
Author affiliation
A
State University of Londrina (UEL), Laboratory of Animal Reproduction, Londrina, Brazil.
Animal Production Science
improve fertility in lactating beef cows. Animal Reproduction Science
163, 89–96. doi:10.1016/j.anireprosci.2015.10.002
Pontes JHF, Nonato-Junior I, Sanches BV, Ereno-Junior JC, Uvo S,
Barreiros TRR, Oliveira JA, Hasler JF, Seneda MM (2009)
Comparison of embryo yield and pregnancy rate between in vivo
and in vitro methods in the same Nelore (Bos indicus) donor cows.
Theriogenology 71, 690–697. doi:10.1016/j.theriogenology.2008.
09.031
Rodrigues WB, Silva AS, Silva JCB, Anache NA, Silva KC, Cardoso CJT,
Garcia WR, Sutovsky P, Nogueira E (2019) Timed artificial
insemination plus heat II: gonadorelin injection in cows with low
estrus expression scores increased pregnancy in progesterone/
estradiol-based protocol. Animal 13, 2313–2318. doi:10.1017/
S1751731119000454
Sá Filho MF, Crespilho AM, Santos JEP, Perry GA, Baruselli PS (2010)
Ovarian follicle diameter at timed insemination and estrous
response influence likelihood of ovulation and pregnancy after
estrous synchronization with progesterone or progestin-based
protocols in suckled Bos indicus cows. Animal Reproduction Science
120, 23–30. doi:10.1016/j.anireprosci.2010.03.007
Santos GMGd, Silva-Santos KC, Barreiros TRR, Morotti F, Sanches BV,
de Moraes FLZ, Blaschi W, Seneda MM (2016) High numbers of
antral follicles are positively associated with in vitro embryo
production but not the conception rate for FTAI in Nelore cattle.
Animal Reproduction Science 165, 17–21. doi:10.1016/j.anireprosci.
2015.11.024
Seneda MM, Morotti F, Zangirolamo AF, da Silva NC, Sanches TK,
Blaschi W, Barreiros TRR (2019) Antral follicle population in
prepubertal and pubertal heifers. Reproduction, Fertility and
Development 31, 10–16. doi:10.1071/RD18344
Silva-Santos KC, Santos GMG, Koetz Júnior C, Morotti F, Siloto LS,
Marcantonio TN, Urbano MR, Oliveira RL, Lima DCM, Seneda MM
(2014) Antral follicle populations and embryo production – in vitro
and in vivo – of Bos indicus–taurus donors from weaning to yearling
ages. Reproduction in Domestic Animals 49, 228–232. doi:10.1111/
rda.12255
Singh J, Domínguez M, Jaiswal R, Adams GP (2004) A simple ultrasound
test to predict the superstimulatory response in cattle. Theriogenology
62, 227–243. doi:10.1016/j.theriogenology.2003.09.020
Zangirolamo AF, Morotti F, da Silva NC, Sanches TK, Seneda MM (2018)
Ovarian antral follicle populations and embryo production in cattle.
Animal Reproduction 15, 310–315. doi:10.21451/1984-3143-
AR2018-0072
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