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Estrous synchronization in ewes: The use of

progestogens and prostaglandins

X. J. Yu, J. Wang & Y. Y. Bai

To cite this article: X. J. Yu, J. Wang & Y. Y. Bai (2019): Estrous synchronization in ewes: The use
of progestogens and prostaglandins, Acta Agriculturae Scandinavica, Section A — Animal Science,
DOI: 10.1080/09064702.2019.1674373

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ACTA AGRICULTURAE SCANDINAVICA, SECTION A — ANIMAL SCIENCE
https://doi.org/10.1080/09064702.2019.1674373

REVIEW ARTICLE

Estrous synchronization in ewes: The use of progestogens and prostaglandins

X. J. Yu *, J. Wang * and Y. Y. Bai
College of Animal Science and Technology, Hebei North University, Zhangjiakou, People’s Republic of China

ABSTRACT ARTICLE HISTORY

This review summarizes the first principles, hormone types, and applied research related to the Received 17 April 2019
synchronization of estrus in ewes. The hormones used to induce synchronization include Accepted 21 September 2019
progestogens, such as medroxyprogesterone acetate (MAP), fluorogestone acetate (FGA), a
KEYWORDS
controlled internal drug-releasing (CIDR) device, progesterone (P4), norgestomet, and Ewe; estrous synchronization;
prostaglandins. These are usually combined with gonadotropins. Research on these hormones progestogen; prostaglandin;
indicates that CIDR, MAP, and FGA can be regarded as equally effective for induction of estrus in gonadotropins
ewes. Prostaglandins are a suitable alternative for estrous synchronization especially considering
that progestogens may have negative effects on the functionality of ovulatory follicles. Artificial
insemination, embryo transfer, and the acceleration of herd genetic trait improvement are
among the potential practical applications of synchronous estrus in ewes.

Introduction
Synchronization of estrus has been used for numerous
animal species worldwide, including swine (Kraeling
et al., 1981; Davis, 2004), dogs (Mogheiseh et al., 2017),
cows (Ribeiro et al., 2012; Ferreira et al., 2018), and
goats (Stelletta et al., 2017), and has been performed in
ewes for several decades (Scales, 1967; Allison & Kelly,
1978; Akusu & Egbunike, 1984; Kesler & Favero, 1997;
Zarkawi, 2001; Martinez-Ros et al., 2018). Although
different specific schemes for synchronization of estrus
have been applied in different regions, different
seasons, and different sheep breeds, the optimal syn-
chronization procedure has yet to be determined. It is
thus instructive to summarize the findings of previous
studies and draw reasonable conclusions.
Estrous synchronization technology refers to the artifi-
cial use of hormone preparations to induce estrus in
female animals to achieve the synchronization of
estrus, pregnancy, and birth (Wildeus, 2000). Successful
application of estrous synchronization has been reported
in several sheep breeds in a number of countries [Paki-
stan: Beetal × Dwarf ewes (Kausar et al., 2009); Mexico:
Suffolk ewes (Ruiz de Chávez et al., 2015); Brazil: Santa
Inês ewes (Oliveira et al., 2016); China: Lanzhou fat-
tailed ewes (Wei et al., 2016); Turkey: Kivircik ewes
(Dogan et al., 2018); and Uruguay: Santa Ines × Dorper
ewes (Cosentino et al., 2019)]. Its application in sheep
herds can promote the synchronized birth of lambs to
improve reproductive efficiency and enable better
farming care for ewes and lambs, thereby enhancing sur-
vival rates (Whitley & Jackson, 2004). Of course, hormone
treatment is not the only way to achieve synchronization
of lambing. Seasonal ewes are usually achieved easily by
delimiting the breeding season.
In ewes, estrus is a cyclic process that can be defined
both physiologically and behaviorally. The estrous cycle
of ewes includes four stages, namely, proestrus, estrus,
metestrus, and diestrus. The physiological characteristics
of estrus are corpus luteum degeneration, decreased pro-
gesterone levels, follicular development, and ovulation.
Diestrus is characterized by luteal formation, secretion
of progesterone, elevated progesterone levels, and inhi-
bition of ovulation and estrus. In ewes, synchronization
of estrus can be achieved using various approaches,
including artificial prolongation of the luteal phase or
shortening of the luteal phase to delay or advance the
arrival of estrus, respectively (Wildeus, 2000). The delay
of estrus in ewes involves the use of exogenous proges-
togen to maintain progesterone levels and inhibit ovu-
lation (Letelier et al., 2010). Induction of estrus entails
the use of prostaglandins and leads to lysing of the
corpus luteum and a reduction in the levels of progester-
one, thereby promoting follicular development and ovu-
lation (Fierro et al., 2013).
In this review, we summarize the types, application,
and influencing factors of protocols used for estrous

CONTACT J. Wang wangjing197410@163.com College of Animal Science and Technology, Hebei North University, Zhangjiakou, Hebei 075000, People’s
Republic of China
*These authors contributed equally to this work
© 2019 Informa UK Limited, trading as Taylor & Francis Group
2 X. J. YU ET AL.
synchronization in ewes, with the aim of distilling current
knowledge, such that it may be harnessed to enhance
the efficacy of this procedure in breeding.
Hormone types used for estrous
synchronization
Hormones used to delay estrus
The progestogens used for estrous synchronization
include medroxyprogesterone acetate (MAP) (Simonetti
et al., 2002; Barrett et al., 2004; Kausar et al., 2009), fluoro-
gestone acetate (FGA) (Amarantidis et al., 2004), pro-
gesterone (P4) (Knights et al., 2001; De et al., 2015),
melengestrol acetate (MGA) (Powell et al., 1996), chlorma-
dinone acetate (CAP), norgesterone, norgestomet, and
norethindrone. The use of progestogen maintains pro-
gesterone levels and inhibits ovulation, thereby inhibiting
estrus. After a period of time, the use of progestogen is
discontinued and the ovaries of ewes are no longer con-
trolled by exogenously administered artificial progesto-
gen. At this point, most ewes will show synchronized
follicular development and will go on to achieve simul-
taneous estrus.
Hormones used to advance estrus
Prostaglandins used to advance estrus include prosta-
glandin F2α (PGF2α) (Ott et al., 1980; Olivera-Muzante
et al., 2011), cloprostenol (an analogue of PGF2α)
(Romano, 1998; Kusina et al., 2001; Contreras-Solis
et al., 2009), methyl prostaglandin, oxyprostenol, and
fluoroprostenol. Prostaglandins can inhibit progesterone
production and accelerate corpus luteum degeneration,
causing the ovaries to lose progesterone control and
thereby promote the development of follicles. This can
shorten the estrous cycle and shorten the time to
estrus in ewes. In this regard, it is worth noting that pros-
taglandins should be used in non-pregnant ewes during
the luteal phase.
Auxiliary hormones
Gonadotropins are generally used in combination with
progestogens and prostaglandins for estrous synchroni-
zation. Numerous types of gonadotropins can be used
for this purpose, including pregnant mare serum gonado-
tropin / equine chorionic gonadotropin (PMSG / eCG)
(Aköz et al., 2006; Chao et al., 2008), gonadotropin-releas-
ing hormone (GnRH) (Titi et al., 2010), follicle-stimulating
hormone (FSH) (Forcada et al., 2011), and human chorio-
nic gonadotropin (hCG) (Abdullah et al., 2002). Although
gonadotropins are often used in conjunction with
progestogens and prostaglandins for estrous synchroni-
zation, these auxiliary hormones are used only to
promote follicle maturation and ovulation, thereby
improving accuracy and synchronicity.
Applied research on estrous synchronization
in ewes
Progestogen use: MAP, FGA, and CIDR
Numerous studies have demonstrated that estrus can be
synchronized in ewes by using progestogen. Common
means of delivery include intravaginal progestogen
sponges and controlled internal drug-releasing (CIDR)
devices. The progestogens most commonly used for
estrous synchronization are MAP and FGA, combined
with or without gonadotropins during the breeding
season or seasonal anestrus phase.
MAP (Table 1) is typically applied for 14 days as an
intravaginal sponge, often combined with an auxiliary
hormone. Merino ewes (n = 130) have been used to
examine the efficacy of different doses of MAP to syn-
chronize estrus outside the breeding season, and it has
been found that different doses (30, 40, and 60 mg) com-
bined with an intramuscular injection of 300 IU PMSG at
sponge withdrawal can induce estrous synchronization,
with a duration similar to that of the natural estrous
period (29.4 ± 7.6 vs. 25.9 ± 6.8 h, respectively) (Greyling
et al., 1994). In order to induce estrous synchronization
outside the normal breeding season, a similar treatment
(MAP 60 mg/14 d + 600 IU PMSG) has been used in indi-
genous Awassi ewes (Zarkawi et al., 1999), with results
indicating that estrus was induced within 36–48 h of
sponge withdrawal in 82% of treated ewes. There were
significant differences between the MAP-treated group
and the control ewes in total estrus response (96% vs.
32.6%), lambing rate (80.0% vs. 32.6%), and fecundity
(137.5% vs. 106.7%). The dosage of MAP has also been
concerned by researchers. In order to determine the
residual and absorbed levels of MAP, different doses of
MAP (40, 50, and 60 mg) have been used in Merino
ewes, with the results indicating that absorbed levels
of MAP did not differ among groups (21.62 ± 1.97 mg,
19.78 ± 2.03 mg, and 24.85 ± 3.23 mg, respectively)
(Simonetti et al., 2000). In addition, there were no differ-
ences in estrus incidence, interval to estrus onset, or rates
of pregnancy among groups, thereby illustrating that
doses lower than 40 mg MAP could be effective for
estrous synchronization in Merino ewes.
FGA (Table 2) is also applied using an intravaginal
sponge combined with an auxiliary hormone. The dur-
ation of FGA administration is variable, with reported
application times ranging from 7 to 14 days. PMSG is
 ACTA AGRICULTURAE SCANDINAVICA, SECTION A — ANIMAL SCIENCE 3

Table 1. Estrous synchronization in ewes with MAP intravaginal sponge.

Reference Breed Dose1 mg Dur.2 d Aux. treat3 n Interval4 h Estrus% Mating5 CR6% LR7% FR8 Time9
Greyling et al. (1994) Merino T1: 60 14 T: 300 IU 35 43.8 ± 2.8 100 Fixed time AI 75 79.2 1.06* out
T2: 40 14 PMSG 35 44.4 ± 10.4 100 62.5 70.8 1.13*
T3: 30 14 35 42.3 ± 1.3 91.3 65.2 95.7 1.47*
C: 0 — C: 0 25 88 72.7 90.9 1.25*
Zarkawi et al. (1999) Awassi T: 60 14 T: 600 IU 50 96* Hand-mating 80.0* 137.5% out
PMSG
C: 0 — C: 0 46 32.6* 32.6* 106.7%
Simonetti et al. (2000) Merino T1: 40 14 — 82 55.94 ± 1.87 79.27 Cervical AI 43.75 — — in
T2: 50 14 187 56.74 ± 1.13 77.42 52.94
T3: 60 14 280 57.70 ± 1.02 80.87 45.45
Kuru et al. (2017) Pirlak BS
T1: 60 11 T: 500 IU 50 44.52 ± 1.95# 100 Natural mating 44 100 — out
T2: 60 11 eCG 25 45.08 ± 2.93# 92 43.48 100
BS
T3: 60 14 49 34.42 ± 1.71# 97.96 41.76 95
T4: 60 14 24 33.74 ± 2.37# 100 37.5 100
Atalla (2018) Assaf C: 0 — C: 0 20 70. 4 ± 30. 4 10 Natural mating — — — out
T1: 60 14 300 eCG 30 60. 7 ± 20. 3 80
T2: 60 14 600 eCG 30 51. 3 ± 25. 2 80
Note: Detailed data on the application of MAP can be found in Table 1, which shows that compared with natural estrus, a single use of MAP or combination of
MAP with PMSG can effectively induce estrus and increase the percentage of estrus in ewes. In addition, the dosage of MAP may be a potential factor influen-
cing lambing rate and fecundity rate.
1
‘Dose’ means the dose of medroxyprogesterone acetate (MAP); 2‘Dur.’ means duration of the MAP treatment; 3‘Aux. treat’ represents ‘auxiliary treatment’, which
means ewes were injected intramuscularly at the sponge withdrawal with different doses of PMSG (eCG); 4‘Interval’ means the interval time from the sponge
withdrawal to the onset of estrus; 5‘Mating’ represents the method of mating and ‘AI’ represents artificial insemination; 6‘CR’ represents conception rate; 7‘LR’
represents lambing rate (percentage of ewes lambing/total ewes mated); 8‘FR’ represents fecundity rate; 9‘Time’ means the research time including ‘out’ rep-
resents outside the breeding season and ‘in’ represents in the breeding season; 10‘BS’ means the ewes of T1 and T3 were injected subcutaneously with barium
selenate. ‘*’ means there is significant difference among groups (P < 0.05). ‘#’ means there is significant difference among groups (P < 0.001).

the most commonly used in combination with FGA and
the combination of FGA and PMSG has been used in
many sheep breeds. A study conducted by Ainsworth &
Shrestha (1985) demonstrated the effect of PMSG dose
on the reproductive performance of ewes using intrava-
ginal sponges containing 40 mg FGA for 14 days to
achieve estrous synchronization. Their results showed
that a higher dose of PMSG had no additional advantage
in terms of ewe fecundity. To determine the best sponge
type and optimum PMSG dose, Abdullah et al. (2002)
conducted three trials in Awassi ewes, in which all
ewes from two trials were administered intravaginal
sponges containing 40 mg FGA for 12 days prior to
receiving different doses of PMSG at the time of

Table 2. Estrous synchronization in ewes with FGA intravaginal sponge.

Dose1 Estrus Litter
Reference Breed mg Dur.2 d A.T.3 n Interval4 h % CR5% LR6% size FR7 Time8
Timurkan & Yildiz (2005) Hamdani PMSG in
T1: 40 14 500 IU 32 100 90.62b 1.06b
T2: 40 14 600 IU 32 100 93.75b 1.25b
T3: 40 14 750 IU 32 100 100b 1.40b
C: 0 — 0 34 97.05 79.41b 0.88b
Kridli & Al-Khetib (2006) Awassi T1:40 12 C: none 7 59.6 ± 7b 100 43a 29b 0.29 TD
RJ 100 75a 50b 0.63
b
T2: 40 12 250 mg/d 8 59.4 ± 7
T3: 40 12 500 mg/d 8 49.6 ± 7b 100 100a 100b 1.13
T4: 40 12 750 mg/d 7 49 ± 8b 85.7 71a 57b 0.71
T5: 40 12 eCG 600IU 7 34 ± 8b 100 86a 71b 0.86
Quintero-Elisea et al. Pelibuey, PMSG
(2011) Blackbelly T1:40 10 C: 0 27 36.3 ± 1.5 89 62.5 66.7b 0.79b NB
T2:40 10 100 IU 24 31.2 ± 1.5 92.7 77.3 58.8b 0.68b
T3:40 10 200 IU 24 28.4 ± 1.5 95.8 69.6 93.7b 1.13b
T4:40 10 400 IU 24 26.1 ± 1.5 91.7 81.8 100b 1.77b
Altinçekiç & Koyuncu Merino T1:30 7 500 IU 26 65.4b — 61.5b 1.25b 0.77c out
(2019) T2:30 10 PMSG 26 80.8b 88.5b 1.48b 1.31c
T3:30 14 26 96.2b 92.3b 1.63b 1.50c
Note: Data relating to results of the application of FGA are summarized in Table 2, which shows that compared with natural estrus, the application of FGA
increases the percentage of estrus in ewes of different sheep breeds. Furthermore, the types and doses of auxiliary hormones are factors that can potentially
influence litter size and the rates of conception, lambing, and fecundity.
1
‘Dose’ means the dose of fluorogestone acetate (FGA); 2‘Dur.’ means duration of the FGA treatment; 3‘A.T.’ represents ‘auxiliary treatment’, which means ewes
were treated with different doses of PMSG (eCG) or royal jelly; 4‘Interval’ means the interval time from the sponge withdrawal to the onset of estrus; 5‘CR’
represents conception rate; 6‘LR’ represents lambing rate (percentage of ewes lambing/total ewes mated); 7‘FR’ represents fecundity rate; 8‘Time’ means
the research time including ‘in’ represents in the breeding season, ‘TD’ represents transition period (from anestrus to the breeding season), ‘NB’ represents
nonseasonal breeds; ‘out’ represents outside the breeding season; Superscript ‘a’ means there is significant difference among groups (P < 0.1); Superscript
‘b’ means there is significant difference among groups (P < 0.05); Superscript ‘c’ means there is significant difference among groups (P < 0.01).
4 X. J. YU ET AL.

Table 3. Comparison research between MAP and FGA.

Dose1 Aux.
Reference Breed mg Dur.2 d treat3 n Interval4 h Estrus CR5% LR6% Litter size FR7 Time8
Ainsworth & Shrestha Synthetic FGA: 40 14 500 IU 314 n = 278 2.3 ± 0.08 52.8 ± 2.3 Out
(1983) strains MAP: 60 14 PMSG 304 n = 267 2.1 ± 0.08 57 ± 2.4
Abdullah et al. (2002) Awassi FGA: 30 12 600 IU 20 49 ± 3.1* n = 18 35 Out
FGA: 40 12 PMSG 20 49 ± 3.1* n = 17 35
MAP: 60 12 20 42 ± 3.1* n = 16 50
Zeleke et al. (2005) Dorper FGA: 40 14 300 IU 100 41.7 ± 0.9 96% 74.0 97.0 1.4 ± 0.1 131.1 TD
MAP: 60 14 PMSG 102 41.9 ± 0.9 98% 70.6 85.3 1.5 ± 0.1 120.8
Note: The data presented in Table 3 highlight the different effects of MAP and FGA on the estrous synchronization in the ewes of different sheep breeds. When the
duration of FGA and MAP treatment was same, it was found that a 60 mg MAP sponge could advance the estrus of Awassi ewes (P < 0.05) to a greater extent
than the use of a 30 or 40 mg FGA sponge.
1
‘Dose’ means the dose of medroxyprogesterone acetate (MAP) and fluorogestone acetate (FGA); 2‘Dur.’ means duration of the MAP or FGA treatment; 3‘Aux.
treat.’ represents ‘auxiliary treatment’, which means ewes were injected intramuscularly at the sponge withdrawal with different doses of PMSG (eCG); 4‘Interval’
means the interval time from the sponge withdrawal to the onset of estrus; 5‘CR’ represents conception rate; 6‘LR’ represents lambing rate (percentage of ewes
lambing/total ewes mated); 7‘FR’ represents fecundity rate; 8‘Time’ means the research time including ‘Out’ represents outside the breeding season and ‘TD’
represents transition period (from the natural breeding to the anestrus season); ‘*’ means there is significant difference among groups (P < 0.05).

sponge removal to induce estrous synchronization. Simi-
larly, Timurkan & Yildiz (2005) used sponges containing
40 mg FGA applied for 12 days, in conjunction with
different doses of PMSG (500, 600, and 750 IU, respect-
ively), to induce the synchronization of estrus in
Hamdani ewes, and accordingly showed that FGA and
PMSG administration may be effective in synchronizing
estrus and parturition and in enhancing pregnancy rate
during the breeding season. To examine the efficacy of
using intravaginal sponges containing 30 mg FGA for
14 days to induce synchronized estrus in Kivircik ewes,
Koyuncu & Ozis Altıcekic (2010) administered PMSG at
three different time points (24 h before, at, or 24 h
after FGA sponge withdrawal) and via two different
routes (intramuscular and subcutaneous). The results
showed that all treated Kivircik ewes achieved synchro-
nized estrus, with no significant differences in estrous
response related to the timing or route of PMSG admin-
istration. Furthermore, to investigate whether royal jelly
could be used as a substitute for gonadotropin, Kridli
and colleagues applied intravaginal sponges containing
40 mg FGA for 12 days in Awassi ewes and compared
GnRH and natural royal jelly paste for reproductive
responses (Kridli et al., 2003), and examined the effects
of eCG or different doses of royal jelly after the
sponges were removed (Kridli & Al-Khetib, 2006).
However, the results revealed that royal jelly did not
affect estrus synchronization in manner comparable to
GnRH and eCG. To investigate the effects of different
doses (0, 100, 200, or 400 IU) and different application
times (48 h before, 24 h before, and at FGA sponge with-
drawal) of PMSG on reproductive performance, Quin-
tero-Elisea et al. (2011) synchronized estrus in hair
sheep ewes (Blackbelly and Pelibuey breeds) using intra-
vaginal sponges for 10 days. They accordingly demon-
strated that administration of FGA intravaginal sponges
for 10 days was effective in inducing synchronized
estrus, with the time of estrus advanced and the
response enhanced in a PMSG dose-dependent
manner. Aköz et al. (2006) sought to determine an
optimal PMSG dose for enhancing prolificacy in Akkara-
man cross-bred ewes synchronized with different doses
of FGA in intravaginal sponges (containing 30 or 40 mg
FGA) for 7 days in the non-breeding season. The results
indicated that treatment for 7 days was sufficient to
achieve estrous synchronization. To determine the
efficacy of estrous synchronization using short-term
and long-term progestogen treatments, sponges con-
taining 30 mg FGA were used for 7 or 12 days (Karaca
et al., 2009), with the results showing that the longer-
term regimen could be replaced with the short-term
regimen at the onset of the breeding season.
Comparative studies on MAP and FGA intravaginal
sponges (Table 3) have been conducted to assess their
respective effectiveness in inducing the synchronization
of estrus. In one such study, Ainsworth & Shrestha
(1983) applied an intravaginal FGA sponge containing
40 mg FGA or an intravaginal MAP sponge containing
60 mg MAP for 14 days, with 500 IU PMSG administrated
at sponge removal to induce the synchronization of
estrus. No significant differences between the FGA and
MAP groups were found with regards to fertility or litter
size. A similar experiment was conducted by Zeleke
et al. (2005) using FGA at 40 mg or MAP at 60 mg for 14
days to compare their efficacy in inducing estrous syn-
chronization in Dorper ewes, with the results indicating
that there were no significant differences in pregnancy,
lambing, or fecundity rates between the two treatments.
Furthermore, in a comparative trial examining the use of
FGA at 30 or 40 mg and MAP at 60 mg for 12 days, Abdul-
lah et al. (2002) showed that estrus was more advanced
(P < 0.05) in ewes receiving MAP compared with FGA.
Accordingly, 60 mg MAP sponges may be more appropri-
ate for estrous synchronization.
Controlled internal drug-releasing (CIDR) devices are
intravaginal devices comprising a progesterone-
 ACTA AGRICULTURAE SCANDINAVICA, SECTION A — ANIMAL SCIENCE 5

Table 4. Comparison research among MAP, FGA and CIDR.

Hormone Dur.2 Litter
Reference Breed type1 d A.T.3 n Interval4 h Estrus CR5% size FR6% Time7
Ainsworth & Downey crosses of Dorset, Leicester CIDR: 0.55 g P 14 500 IU 192 — 92% — 2.3 69 in
(1986) and Suffolk FGA: 40 mg 14 PMSG 194 91% 2.6 64
Rhodes & Nathanielsz crossbred Western range CIDR: 0.3 g P 14 — 97 — n = 85 57.7% — in
(1988) MAP: 60 mg 14 99 n = 88 56.6%
Ungerfeld & Rubianes Polwarth and Polwarth x Ile MAP: 60 mg 6 380 IU 51 44.6 ± 1.7* 94.1% 62.5% — — non
(2002) de France FGA: 30 mg 6 eCG 47 38.8 ± 1.6* 91.5% 67.4%
CIDR: 0.3 g P 6 49 39.9 ± 2.1* 95.9% 59.6%
Hashemi et al. (2006) Karakul CIDR: 0.3 g P 12 500 IU 30 30.1 ± 7.6 93.3% — — — non
MAP: 60 mg 12 eCG 30 29.6 ± 5.6 100%
Swelum et al. (2015) Najdi CIDR: 0.3 g P 14 600 IU 150 96.95% 77.86* — 75.57* in
FGA: 30 mg 14 eCG 150 95.75% 62.41* 60.99*
Note: On the basis of the aforementioned studies, CIDR, MAP, and FGA can be regarded as equally effective for estrous induction (Table 4). Although compared with
MAP and FGA, CIDR may advance estrus, there are no differences among these treatments with regard to the percentage of estrus and conception rate in ewes.
1
‘Hormone type’ include medroxyprogesterone acetate (MAP), fluorogestone acetate (FGA) or the controlled internal drug-releasing (CIDR) device and ‘P’ represents
progesterone; 2‘Dur.’ means duration of the MAP, FGA or CIDR treatment; 3‘A.T.’ represents ‘auxiliary treatment’, which means ewes were injected intramuscularly
at the sponge withdrawal with different doses of PMSG (eCG); 4‘Interval’ means the interval time from the sponge or device withdrawal to the onset of estrus; 5‘CR’
represents conception rate; 6‘FR’ represents fertility rate; 7‘Time’ means the research time including ‘non’ represents non-breeding season and ‘in’ represents in
breeding season; ‘*’ means there is significant difference among groups (P < 0.05).

impregnated medical silicone elastomer molded over a
nylon core (Wheaton et al., 1993). Numerous
studies (Table 4) have been conducted to evaluate the
effectiveness of these devices in controlling the estrous
cycle of ewes, including comparisons between the
device and MAP (Rhodes & Nathanielsz, 1988; Iida et al.,
2004; Hashemi et al., 2006) or FGA (Ainsworth &
Downey, 1986), as well as among CIDR, MAP, and FGA
combined (Fukui et al., 1999; Ungerfeld & Rubianes,
2002; Romano, 2004).
The most commonly used dose of MAP is 60 mg,
whereas the dose of FGA is 40 mg. Rhodes & Nathanielsz
(1988) used MAP sponges at 60 mg and a CIDR device
containing 366 mg of natural progesterone in crossbred
Western range ewes without PMSG after sponge
removal, and found no significant difference in the
rates of marking by rams or pregnancy between the
two groups. However, earlier estrus and closer synchrony
were achieved using the CIDR device, thereby indicating
that CIDR is comparable to the MAP sponge in terms of
synchronizing estrus during the breeding season. Simi-
larly, Iida et al. (2004) used CIDR containing 300 mg pro-
gesterone, an MAP sponge at 60 mg, and a custom-made
sponge containing 500 mg progesterone to treat Suffolk,
South Down, and Dorset crossed ewes for 12 days. Estrus
was induced using both CIDR and MAP, with no consider-
able difference in the mean onset time of estrus (23 h vs.
21 h, respectively). These results illustrate that both the
MAP sponge and CIDR device are effective in inducing
estrous synchronization in ewes. Additionally, Hashemi
et al. (2006) used a CIDR device containing 300 mg pro-
gesterone and an MAP 60 mg sponge for 12 days, fol-
lowed by 500 IU eCG in Karakul ewes, and found no
significant difference between the two groups in terms
of the mean interval times between treatment cessation,
onset of estrus, and estrus duration. These findings
accordingly indicate that both CIDR devices and MAP
sponges can be used to induce estrous synchronization
in Karakul ewes outside the natural breeding season.
To facilitate estrus synchronization in three-way
crosses of Dorset, Leicester, and Suffolk breeds, Ains-
worth & Downey (1986) used a CIDR device and FGA
sponges containing 40 mg for 14 days, followed by 500
IU PMSG, and found that mating response, fertility, and
litter size were similar between groups. These results
indicate that CIDR devices can be as effective as FGA-
impregnated intravaginal sponges, and thus constitute
a promising alternative regimen.
Further comparisons have been made to investigate
the best regimen among the CIDR device, MAP sponge,
and/or FGA sponge. These three types of progestogen
intravaginal devices were used by Ungerfeld & Rubianes
(2002) to examine estrus induction in Polwarth and Pol-
warth × Ile de France ewes during the non-breeding
season. MAP sponge, FGA sponge, or CIDR device were
used for 6 days, followed by 380 IU eCG; however, no sig-
nificant differences were detected in either estrus or con-
ception rates among recipients of the three treatments,
indicating that the CIDR device and FGA and MAP
sponges may be equally effective over 6 days for estrus
induction in anestrous ewes.
The use of other progestogens: P4, MGA, and
norgestomet
In studies conducted to examine the efficacy of different
doses of progesterone (P4 at 350 and 300 mg) used for
12 days in crossbred Bharat Merino ewes during the
autumn breeding season, Das et al. (2000) found that
ewes treated with the higher dose showed better
estrus (75% vs. 42%) and synchronization (93% vs. 56%)
responses than those treated with a lower dose at 72 h
6 X. J. YU ET AL.
after sponge removal. Although lower doses of P4 signifi-
cantly delayed the onset of estrus, the dose of P4 had no
marked influence on estrus length, conception rate, or
the proportion of ewes lambing at term. The findings
of this study thus indicate that the dose of progesterone
may affect the occurrence of estrus and the response
time in crossbred Bharat Merino ewes. Biehl et al.
(2019) evaluated the effects of new and reused pro-
gesterone (P4) devices in long (11 days) and short (7
days) protocols on time to estrus, estrus synchronization,
and pregnancy rate in Santa Inês ewes. The results
showed that estrous synchronization was efficiently pro-
moted by re-utilization of P4 devices, which did not affect
pregnancy rates at synchronization protocols. In
addition, long hormonal protocols (11 days) tended to
increase the pregnancy rate at estrus synchronization.
Melengestrol acetate (MGA), a commercialized pro-
gesterone, is also effective for estrous synchronization in
ewes. A study by Quispe et al. (1994) demonstrated that
dietary MGA can be an effective alternative protocol for
estrous synchronization in Suffolk, Dorset, and Pelibuey
ewes. Ewes provided with an MGA-containing diet
(0.25 mg MGA·ewe−1·d−1) showed a more pronounced
estrous response and higher lambing rates than ewes in
the control group (P < 0.05). In addition, a protocol of
MGA + estradiol (0.25 mg MGA·ewe−1·d−1 for 8 days,
20 μg estradiol-17β i.m. injection at 54 h after the final
feeding of MGA) was found effective in inducing estrous
synchronization in ewes (Powell et al., 1996). Reproductive
response of 415 fat tailed ewes synchronized with different
lengths of MGA treatments (9 days vs. 12 days) were eval-
uated (Emsen et al., 2011). The results showed that estrus
rates were significantly lower in ewes treated with MGA for
9 days than in the ewes treated with MGA for 12 days (62%
vs. 89%, P < 0.0001), and that the pregnancy rates were
similar in short term and long term MGA-treated ewes
(41% vs. 44%, P > 0.05). Different doses of MGA were
used to estimate the efficiency in inhibition of ovulation
in synchronized Dorper and Dorper-Pelibuey ewes (Salas-
Razo et al., 2014). The results showed that an intermediate
dose of 0.22 mg of MGA per ewe was effective to inhibit
ovulation without affecting the normal development of
follicles; therefore, an intermediate dose of MGA can facili-
tate better estrous synchronization and fertility rates.
Norgestomet, another commercialized progesterone,
has been used in cattle for synchronized estrus. Recent
studies have shown that it is also effective in ewes for
estrous synchronization. In a previous study, Luther
et al. (2007) used norgestomet implants for 14 days to
induce estrous synchronization in Hampshire and Mon-
tadale ewes, followed by various treatments after
implant removal, including 400 IU eCG, 25 μg GnRH, or
400 IU eCG + 25 μg GnRH. The results indicated that
the use of norgestomet combined with eCG was
effective for estrous synchronization and increasing
pregnancy rates. Conception rate and lambing rate
were estimated in Iranian Kourdish breed fat-tailed
ewes treated using implant norgestomet with or
without PMSG (Garoussi et al., 2012). The results demon-
strated that combination of implant norgestomet with
PMSG could increase and improve the fertilization rate
of ewes in the out-of-breeding season. Fifty-five Texel
ewes were used to assess the efficacy of re-utilization
of norgestomet by ear implant insertion in estrous syn-
chronization response and pregnancy rates (Bragança
et al., 2019). The results showed that estrus behavior
(93.3% vs. 90%, P > 0.05) and pregnancy rates (73.3%
vs. 68%, P > 0.05) did not differ statistically between the
new norgestomet treatment group and re-utilization
ear implant group. These results indicate that norgesto-
met ear implant treatment was an effective protocol
for estrous synchronization in ewes.
Prostaglandin use
Prostaglandin F2α and its synthetic analogues have been
used extensively for luteolysis to facilitate estrous syn-
chronization. However, it is important to note that prosta-
glandins cannot be used to dissolve the corpus luteum at
all points in the estrous cycle, as ovine corpus luteum is
refractory to prostaglandin treatment for up to 2 days
after ovulation (Fierro et al., 2013). Follicular response to
prostaglandin treatment occurs according to the
specific phase of estrus, and therefore, the timing of pros-
taglandin treatment is critical. However, for a flock of
sheep, the specific stage of the estrous cycle in individual
ewes is uncertain, and thus, when prostaglandin is used to
promote estrous synchronization, researchers typically
apply a second treatment at 7–12 days after the first use
of prostaglandin to ensure that all ewes can reach estrus
and conceive. In this regard, Hackett et al. (1981a)
assessed the fertility and prolificacy among ewes of
three breeds of sheep after synchronization of estrus
using prostaglandin F2α via two intramuscular injections
of 15 mg PGF2α administered 11 days apart, followed by
artificial insemination at 60, 72 h, or at 60 and 72 h after
the second PGF2α injection. Although differences were
detected in fertility among the different artificial insemi-
nation times, there were no differences in prolificacy
among the different breeds inseminated at different
times. To determine the efficacy of GnRH-PGF2α treatment
for synchronizing estrus in sheep during the breeding
season, Ataman & Aköz (2006), administered a control
group with two injections of PGF2α at a 9-day interval,
with the results showing that GnRH-PGF2α treatment
was an effective measure for inducing estrous
 ACTA AGRICULTURAE SCANDINAVICA, SECTION A — ANIMAL SCIENCE 7

synchronization in ewes and that two injections of prosta-
glandin can effectively induce synchronization. In order to
compare the effects of PGF2α and D-cloprostenol for
estrous synchronization in hair sheep during the breeding
season, 61 hair sheep were treated with two doses of
50 μg PGF2α (n = 30) and D-cloprostenol (n = 31) intra-
muscular injection with a 12-day interval (Ramírez et al.,
2018). Ewes in estrus of 48 h after the last injection
showed significant difference (37.1% vs. 65.7%, P < 0.05),
whereas the estrus duration (42 + 6.1 h vs. 41.1 + 11.2 h,
P > 0.05) and pregnancy rate (38.4% vs. 52.1%, P > 0.05)
were similar between the two groups. These results
accordingly indicate that prostaglandins and their ana-
logues are effective in synchronizing estrus in the ewes
of different breeds.
Comparative studies between progestogens and
prostaglandins
To date, there have been only a few studies studies
that have examined the induction of estrus in ewes
using prostaglandins and their analogues alone, and
these have generally compared prostaglandins with
progestogens to determine the most effective protocol
for inducing synchronous estrus in the ewes of a
specific breed.
Based on the available information, it appears that pro-
gestogens can be used earlier than prostaglandins.
However, the hormone residues and spent delivery
devices associated with intravaginal application of proges-
togen can constitute sources of environmental pollution,
making prostaglandins a more sustainable option.
Although various studies on estrous synchronization that
have compared progestogens with prostaglandins have
demonstrated that prostaglandins can replace
progestogens, their effectiveness may vary depending
on the sheep breed (Table 5).
A comparison of PGF2α (two injections with an 11-day
interval) and FGA sponges (40 mg for 12 days) con-
ducted by Hackett et al. (1981b) in Corriedale ewes
during the breeding season revealed that the pro-
portions of estrous response and fertility of ewes did
not differ, thereby demonstrating that PGF2α may be
effective as an alternative to FGA for estrous synchroniza-
tion. To compare the reproductive performance of ewes
treated with PGF2α or P4, Olivera-Muzante et al. (2011)
examined Australian Merino ewes during the breeding
season using timed artificial insemination with fresh or
chilled semen, and found that ewes treated with PGF2α
had lower fertility and fecundity than those treated
with P4. Furthermore, in a comparison of PGF2α adminis-
tered as two injections 12 days apart and progestogen
sponges for the induction of synchronized estrus in
parous Ethiopian Menze ewes, Mutiga & Mukasa-
Mugerwa (1992) found that ewes treated with PGF2α
showed significantly earlier estrus than those treated
with progestogen sponges, although lamb births did
not differ significantly between the two treatments.
In a comparative study conducted to examine estrous
synchronization, Godfrey et al. (1997) used two PGF2α
injections of 15 mg (total 30 mg) administered 10 days
apart and CIDR for 12 days to induce synchronization in
St. Croix White ewes, and showed that ewes treated
with the CIDR device exhibited estrus significantly
earlier than those treated with PGF2α, with a non-signifi-
cant difference in the conception rate. In a further com-
parative study, Naderipour et al. (2012) examined
estrous synchronization in fat-tailed Iranian Kalkuhi
ewes during the breeding season in response to treat-
ment with sponges containing 60 mg medroxyprogester-
one applied for 14 days, a CIDR device applied for 12

Table 5. Comparison research between progestogens and prostaglandins.

Dur.2 Litter
Reference Breed Dose1 d A.T.3 n Interval4 d Estrus CR5% LR6% size Fertility
Hackett et al. Corricdale, Finn PGF2α: 15 mg 11 — 70 84 ± 4.3 1.3 63 ± 7.6
(1981b) FGA: 40 mg 12 — 71 89 ± 4.1 1.3 78 ± 6.7
Godfrey et al., St. Croix White PGF2α: 15 mg 10 — 14 2.9 ± 0.4** 71.4%** 86 1.9 ± 0.2
1997 CIDR 12 — 14 1.4 ± 0.4** 100%** 100 2.2 ± 0.2
Alnimer et al., Awassi PGF2α: 20 mg 10 — 20 85% 55% 40
2005 FGA: 40 mg 14 600 IU 15 80% 47% 40
PMSG
Wei et al., 2016 Lanzhou fat tailed CLO: 0.24 mg 11 — 117 45.35 ± 6.16* 70.91%* 88.89* 132.69*
ewes FGA: 50 mg 11 400 IU eCG 452 50.46 ± 7.15* 95.97%* 91.37* 150.00*
Note: Some of the comparative results relating to the reproductive performance of ewes treated with PGF2α, cloprostenol (the analogue of PGF2α), FGA, or CIDR
are presented in Table 5, which shows that progestogen and prostaglandin are effective for synchronizing estrus in ewes, although it is not possible to gen-
eralize the effects on different sheep breeds.
1
‘Dose’ means the dose of PGF2α, fluorogestone acetate (FGA), the controlled internal drug-releasing (CIDR) device and Cloprostenol (CLO, the analogue of PGF2α);
2
‘Dur.’ means duration of the FGA and CIDR treatment or the interval time of two injection of PGF2α and CLO; 3‘A.T.’ represents ‘auxiliary treatment’, which means
ewes were treated with different doses of PMSG (eCG); 4‘Interval’ means the interval time from the sponge withdrawal to the onset of estrus; 5‘CR’ represents
conception rate; 6‘LR’ represents lambing rate (percentage of ewes lambing/total ewes mated); ‘*’ means there is significant difference among groups (P < 0.05);
‘**’ means there is significant difference among groups (P < 0.01).
8 X. J. YU ET AL.
days, and PGF2α administered via two injections 11 days
apart. They accordingly found that the proportion of
ewes exhibiting estrous behavior was not significantly
different among the three treatment groups. Moreover,
ewes treated with CIDR exhibited clear signs of estrus
approximately 5 and 10 h earlier than the ewes treated
with PGF2α and medroxyprogesterone sponges, respect-
ively. Using St. Croix White and Barbados Blackbelly
ewes, Godfrey et al. (1999) assessed the efficacy of
three methods of estrous synchronization, namely,
PGF2α administered via two 15 mg injections 10 days
apart (total 30 mg), 300 mg progesterone via CIDR for
12 days, and sponges containing 500 mg progesterone
for 12 days. The results revealed that there were no sig-
nificant differences among the treated ewes with
regard to the time from ram introduction to ovulation,
the time to the preovulatory luteinizing hormone surge,
and progesterone levels during 16 days after synchroni-
zation, thereby indicating that progesterone sponges,
CIDR, and PGF2α can be considered comparably
effective alternatives.
During the early anestrous season in Kalkuhi ewes, Yadi
et al. (2011) investigated estrous synchronization via two
injections of prostaglandin 11 days apart, MAP sponge
for 14 days, or CIDR for 12 days, and comparisons of the
percentage of gestation, twinning rate, and lambing
numbers indicated that the use of prostaglandins was
more effective than either the MAP sponge or CIDR.
Cloprostenol, a PGF2α analogue, has been used in
West African ewes at 43.75 μg, administered via two
intramuscular injections 9 or 7 days apart (Contreras-
Solis et al., 2009), with results indicating that this is a suit-
able alternative method for estrous synchronization.
Given that a previous study has found that progestogens
may have negative effects on the functionality of ovula-
tory follicles (Gonzalez-Bulnes et al., 2005), prostaglandin
analogues may be useful for assisted reproduction. Thus,
prostaglandins can be considered to have extensive
potential applications.
Conclusions and implications
Synchronization of estrus in ewes can be achieved by
delaying or advancing estrus, using progesterone and/
or prostaglandin hormones. These hormones usually
combined with PMSG. The most commonly used dose
of MAP is 60 mg, though a dose less than 40 mg is also
effective. The most common use time of MAP is usually
14 days. The most commonly used dose of FGA is
40 mg, though the dose of 30 mg is equally effective.
The duration of FGA varies from 7 to 14 days. In fact,
FGA treatment for 7 days was sufficient to achieve
estrous synchronization. Compared with 30 mg FGA or
40 mg FGA, 60 mg MAP sponges may be more appropri-
ate for estrous synchronization. A CIDR device containing
300 mg progesterone is as effective as the 60 mg MAP
sponge. PGF2α or Cloprostenol, a PGF2α analogue, is a
suitable alternative method for estrous synchronization,
especially considering that progestogens may have
negative effects on the functionality of ovulatory follicles.
Thus, prostaglandins can be considered to have exten-
sive potential applications. However, although estrous
synchronization schemes can effectively improve the fer-
tility and lambing rates of ewes as well as the subsequent
rates of lamb survival, each method is associated with
certain shortcomings and is preferably used in conjunc-
tion with other treatments. The use of intravaginal pro-
gestogens can lead to environmental contamination
via dissemination of hormone residues in addition to
physical material waste, and may also contribute to vagi-
nitis in ewes. The use of prostaglandins has some limit-
ations, in that it is more time-sensitive, given that these
hormone-like compounds accelerate luteal dissolution
and cannot be used during seasonal anestrus (Wildeus,
2000). Furthermore, it is noteworthy that there is one
major potential drawback of hormonally synchronizing
ewes, namely, the potential breeding of animals with a
low natural fertility, leading to an unwanted and easily
overlooked long-term decrease in population fecundity.
Despite these drawbacks, synchronization of estrus has
broad potential applications in terms of artificial insemi-
nation (Palo et al., 2007; Blaschi et al., 2014; De et al.,
2015), embryo transfer in intensive breeding (Rowson &
Moor, 1966; Larsson et al., 1991), and accelerating herd
genetic improvement (Lupi et al., 2016). In addition, the
application of synchronous estrus technology can also
facilitate the management of large-scale farms, by estab-
lishing chronologically consistent reproductive events
and facilitating better seasonal division of labor tasks,
thereby improving efficiency in terms of lamb survival
rates and economic gains.
Acknowledgements
We would like to thank Editage [www.editage.cn] for English
language editing.
Disclosure statement
No potential conflict of interest was reported by the authors.
Funding
This work was supported financially by the [Key Research and
Development Plan Program of Hebei Province, China. #1]
under [grant number 18236609D]; [Funding for scientific
research projects from the training fund of the Talent Project
 ACTA AGRICULTURAE SCANDINAVICA, SECTION A — ANIMAL SCIENCE
of Hebei Province, China. #2] under [grant number
A2017002061]; and [Science and Technology Research and
Development Plan Program of Zhangjiakou City, Hebei, China.
#3] under [grant number 1711056I].
ORCID
X. J. Yu http://orcid.org/0000-0002-7568-1029
J. Wang http://orcid.org/0000-0003-4833-6024
Y. Y. Bai http://orcid.org/0000-0002-8306-3722
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