Animal Reproduction Science 102 (2007) 208216

Induction of cyclicity in postpartum anestrous beef

cows using progesterone, GnRH and
estradiol cypionate (ECP)
J.E. Wheaton a, , G.C. Lamb b
a Department of Animal Science, University of Minnesota, Twin Cities Campus, 495 Animal Science/Veterinary
Medicine Building, 1988 Fitch Avenue, St. Paul, MN 55108, United States
b North Central Research and Outreach Center, 1861 Highway 169 E, Grand Rapids, MN 55744, United States
Received 6 July 2006; accepted 16 November 2006
Available online 19 November 2006

Abstract
The objective of the present study was to determine whether treatment of postpartum multiparous and
primiparous anestrous beef cows with an intravaginal progesterone-releasing insert (CIDR) and PGF2 , with
and without the addition of GnRH or estradiol cypionate (ECP) at the time of CIDR insertion, is effective
in stimulating onset of estrous cycles. Postpartum lactating Angus primiparous (n = 47, 2 years of age,
495 6 kg) and multiparous (n = 76, 3 years of age, 553 9 kg) cows were assigned by calving date to
four blocks spaced 21-day apart. Cows were assigned sequentially by calving date to four treatment groups:
(1) PGF2 (n = 30), (2) CIDRPGF2 (n = 30), (3) GnRHCIDRPGF2 (n = 33), and (4) ECPCIDRPGF2
(n = 27). Intravaginal CIDR inserts were in place from days 7 to 0. A single 100 g injection of GnRH
or 2 mg ECP were administered on day 7, and 25 mg PGF2 was administered on day 0. Day 0 averaged
38 1 day postpartum. Blood samples were collected on days 19, 9, 0, 5, 9, 12, 16, 19, 23, 26, and 30 for
determination of plasma progesterone concentrations. Pre-treatment luteal activity (progesterone 1 ng/ml)
was detected in 19% of primiparous and 8% of multiparous cows. Progesterone concentrations on day 0
were greater (P < 0.001) in primiparous (3.2 0.3 ng/ml) than multiparous (2.0 0.2 ng/ml) cows. Following
CIDR withdrawal, progesterone concentrations from days 5 to 30 were used to categorize response profiles
as either: (1) treatment-induced onset of estrous cycles, (2) continued anestrus, or (3) spontaneous ovulation
and subsequent formation of a CL. Incidence of treatment-induced onset of estrous cycles, which was
defined as progesterone concentrations 1 ng/ml in three or more consecutive samples from days 9 to 19,
was influenced by treatment and parity. Percentages of cows initiating estrous cycles were greater (P < 0.001)
in the three CIDR-treated groups than in the PGF2 group (55 and 8%, respectively). Percentages of cows
initiating estrous cycles in the CIDRPGF2 , GnRHCIDRPGF2 , and ECPCIDRPGF2 groups were 55,

Corresponding author. Tel.: +1 612 624 1225; fax: +1 612 625 2743.
E-mail address: wheat001@umn.edu (J.E. Wheaton).

0378-4320/$ see front matter 2006 Elsevier B.V. All rights reserved.
doi:10.1016/j.anireprosci.2006.11.006
 J.E. Wheaton, G.C. Lamb / Animal Reproduction Science 102 (2007) 208216 209

58, and 52%, respectively. Incidence of treatment-induced estrous cycles in the three CIDR-treated groups of
cows was greater (P = 0.008) in primiparous (76%) than multiparous (43%) cows. Treatment of postpartum
anestrous primiparous and multiparous beef cows with CIDRPGF2 approximately 40-day postpartum
provides an approach to increase the percentage of cows that have reinitiated estrous cycles by the start of
the breeding season.
2006 Elsevier B.V. All rights reserved.

Keywords: Beef cattle; Postpartum anestrus; CIDR; GnRH; Estradiol cypionate

1. Introduction

Research into the physiology of the postpartum cow has led to the development of treatment
protocols and products that can be applied during the postpartum period to increase fertility of
beef cows by the start of the breeding season (Day, 1998; Yavas and Walton, 2000a,b). Conception
during the early part of the breeding season concentrates the calving season, yields a more uniform
and heavier calf crop and lengthens intervals between calving and the onset of the subsequent
breeding season. Longer intervals between calving and the onset of the subsequent breeding
season increases the proportion of cows initiating estrous cycles before the start of the breeding
season (Lamb and Dahlan, 2002).
In the mid-western states approximately 50% of postpartum cows remain anestrus by the start
of the breeding season (Lamb and Dahlan, 2002). Short-term (<10 days) progestogen treatment
stimulates onset of estrous cycles in some postpartum anestrous beef cows (Day, 1998). More-
over, progestogen treatment prior to ovulation leads to the normal duration of the CL formed
following ovulation (Cooper et al., 1991). More recent progestogen-based protocols have used
6- or 7-day progesterone treatment delivered via an intravaginal CIDR insert (Fike et al., 1997;
Perry et al., 2004; Lucy et al., 2001). In multiparous postpartum anestrous beef cows, the inci-
dence of estrus and onset of estrous cyclicity (ovulation and subsequent formation of a CL of
normal life span) is increased following CIDR removal. Lamb et al. (2001) and other investi-
gators (Day, 1998; Stevenson et al., 2003; Rivera et al., 1998; Colazo et al., 2003) incorporated
CIDR treatment into protocols designed to synchronize ovarian follicular waves and ovulation for
fixed-timed AI. The protocols generally include: (1) the administration of GnRH or an estrogen
at or near the time of CIDR insertion to initiate the emergence of a follicular wave, (2) proges-
terone delivered via the CIDR to prevent estrus and ovulation, (3) an injection of PGF2 at the
time of CIDR removal to induce luteolysis in cows that bear a CL, and (4) a second adminis-
tration of GnRH or an estrogen following CIDR withdrawal to induce ovulation for fixed-timed
AI. Although Lamb et al. (2001) and Stevenson et al. (2003) reported that one such integrated
protocol was effective in increasing pregnancy rates, it could not be ascertained from these
previous results whether treatment influenced pregnancy rates in anestrous primiparous cows.
Furthermore, assessment of incidence of estrous cycle induction was precluded by the inclu-
sion of the hormonal induction of ovulation, AI, and pregnancy rate as the measure of treatment
effectiveness.
The objective of the present study was to determine whether CIDRPGF2 treatment, with
and without the addition of GnRH or estradiol cypionate (ECP) at the time of CIDR insertion,
is effective in stimulating estrous cyclicity in postpartum multiparous and primiparous anestrous
cows. Information gained may prove useful in the planning of reproductive management strategies
for mature and first-calf cows.
210 J.E. Wheaton, G.C. Lamb / Animal Reproduction Science 102 (2007) 208216

2. Material and methods

Suckled Angus primiparous (n = 47, 2 years of age, BW at calving = 495 6 kg) and mul-
tiparous (n = 76, 3 years of age, mean = 4.7 years, range = 311, BW at calving = 553 9 kg)
were assigned by calving date (range February 27, 2003May 19, 2003) to four blocks spaced
21-day apart. Data from three mature cows were not used because two of the animals had not
been suckled and another had expelled the CIDR. Numbers and ages of cows assigned to each
block are shown in Table 1. Dams were assigned sequentially by calving date to four treatment
groups: (1) PGF2 (n = 30), (2) CIDRPGF2 (n = 30), (3) ECPCIDRPGF2 (n = 33), and (4)
GnRHCIDRPGF2 (n = 27). All cows were administered a single im injection of PGF2 (25 mg,
Lutalyse, Pfizer Animal Health, New York, NY) on day 0. Corresponding dates for day 0 for each
block are given in Table 1. Exogenous progesterone was delivered via a CIDR (EAZI-BREED,
1.38 g progesterone, Pfizer Animal Health) from days 7 to 0. A single im injection of GnRH
(100 g, Gonadorelin Hydrochloride, Fort Dodge Animal Health, Fort Dodge, IA) or ECP (2 mg,
Pfizer Animal Health) was administered on day 7. Day 0 averaged 40 days (range 2848 days) in
2-year-old cows and 37 days (range 2848 days) in postpartum 3-year-old cows. Body weights
and body condition scores (BCS, one to nine with three being thin, five moderate and seven fleshy
to fat (Whitman, 1975) were recorded at calving and on day 7. Tail vein blood samples were
collected into evacuated, heparinized tubes on days 19, 9, 0, 5, 9, 12, 16, 19, 23, 26, and 30.
Cows were not exposed to bulls until after the last day of blood collection to avoid confounding
effects of treatment and bull exposure.
Plasma samples (0.1 ml) were assayed in duplicate for progesterone concentration using a solid
phase RIA (Coat-A-Count, Diagnostic Products Co., Los Angeles, CA) as described previously
(Hegstad et al., 1992). Six assays were conducted. The intra-assay CV was 9.6% and the interassay
CV was 12.4%. Assay sensitivity was 0.1 ng/ml. Plasma samples collected from ovariectomized
cows had non-detectable progesterone concentrations.
Data were analyzed using the Statistical Analysis System Release 9.1, licensed to the University
of Minnesota (SAS Inst. Inc., Cary, NC, USA). Data from cows with evidence of pre-treatment
luteal activity were removed from data sets that were analyzed for treatment effects. Evidence
of pre-treatment luteal activity was defined as a progesterone concentration 1 ng/ml in the day

Table 1
Number, age, and days postpartum of cows assigned to blocks 14
Block (day 0) Age (years) n Calving to day 0 (d kg)

1 (April 16) 2 21 41 1.4

3 5 28 0.2
2 (May 7) 2 14 41 1.8
3 31 39 1.4
3 (May 28) 2 8 36 1.9
3 18 37 1.6
4 (June 18) 2 4 40 1.9
3 19 38 1.3
Total 2 47 40 0.9
3 73 37 0.8

Day 0 refers to the day of CIDR removal and (or) PGF2 administration.
 J.E. Wheaton, G.C. Lamb / Animal Reproduction Science 102 (2007) 208216 211

19 or 9 plasma sample. In cows without evidence of pre-treatment luteal activity, progesterone

profiles determined from concentrations in the days 530 plasma samples were grouped into
three categories: treatment-induced estrous cyclicity, anestrus, and spontaneous CL. Treatment-
induced estrous cyclicity was defined by progesterone concentrations 1 ng/ml in three or more
consecutive samples from days 9 to 19. Anestrous cows had 0 or 1 sample with a progesterone
concentration 1 ng/ml from days 5 to 30. Spontaneous CL included cows with two consecutive
samples having 1 ng/ml from days 9 to 19 (presumed short-lived CL) or two or more consecutive
samples 1 ng/ml from days 19 to 30 (spontaneous ovulation and CL formation).
Categorical data were tested using the CATMOD Procedure. Initial analyses tested for effects
of parity, block and block parity interaction on the incidence of pre-treatment luteal activity, and
for effects of treatment, parity, treatment parity, block and block treatment, and block parity
interactions on the incidence of posttreatment estrous cyclicity. Probabilities were > 0.1 for block
and block interactions and these factors were not included in final analyses. A contrast state-
ment within CATMOD was used to test for differences between treatment groups PGF2 versus
CIDRPGF2 + ECPCIDRPGF2 + GnRHCIDRPGF2 . Continuous, serial data (BW and
BCS recorded at calving and again at the time of treatment and plasma progesterone concentra-
tions following CIDR removal (days 530) were analyzed using the MIXED Procedure with a
repeated measures statement using the AR (1) autoregressive covariance structure. Means were
compared using the TUKEY method.
Primiparous and multiparous cows were similar genetically and managed at locations 11.2 km
apart. Prepartum diets were designed to meet or exceed NRC (1996) requirements for non-
lactating, pregnant cows (last third of gestation). Following parturition, primiparous cows received
a diet that included ad libitum access to a legume and grass silage (16.0% crude protein [CP]
and 61.3% total digestible nutrients [TDN]) plus 2.3 kg of a supplement containing dry cracked
corn and a complete inorganic mineral mix (9.8% CP and 90.0% TDN). On June 4, 2003, all
primiparous cows were turned out to pasture containing alfalfa (Medicago sativa) and orchard-
grass (Dactylis glomerata). They also received 2.3 kg supplement. Multiparous cows received
a diet that included corn silage (9.2% CP and 66.0% TDN) and ad libitum access to grass hay
(11.4% CP and 57.2% TDN) and free choice inorganic mineral to achieve an individual daily
consumption of 0.11 kg mineral/day. On May 28, 2003, all multiparous cows were placed into
pastures containing either alfalfa (M. sativa) and reed canarygrass (Phalaris arundinacea) or red
clover (Trifolium pratense) and orchardgrass (D. glomerata). The cows had free choice access
to inorganic mineral. Animal care and use protocols were approved by the Institutional Animal
Care and Use Committee (IACUC; #0305A48281).

3. Results

Pre-treatment luteal activity was evident in 19% (9/47) of primiparous cows and 8% (6/73) of
multiparous cows (P = 0.08). Mean BW and BCS were similar (P > 0.1) for primiparous cows in
which pre-treatment luteal activity was and was not detected (Table 2). Mean BW and BCS were
likewise similar in multiparous cows that did and did not express pre-treatment luteal activity.
At calving BCS were greater (P < 0.001) in primiparous than multiparous cows, but were sim-
ilar at the time of treatment initiation (Table 2). Primiparous cows lost a greater percentage of
their BW during the calving-to-treatment interval than multiparous cows (0.16 0.03% versus
0.01 0.02% of their BW at calving per day, P < 0.001). In those cows that had been treated with
a CIDR, progesterone concentrations on the day of CIDR removal averaged 3.2 0.3 ng/ml in
primiparous cows and 2.0 0.2 ng/ml in multiparous cows (P < 0.001). The greater progesterone
212 J.E. Wheaton, G.C. Lamb / Animal Reproduction Science 102 (2007) 208216

Table 2
Body weight and body condition scores (19 scale) for primiparous and multiparous cows recorded at calving and on day
7
Parity BW (kg) BCS
Calving Day 7 Calving Day 7

Primiparous
No pre-treatment luteal activitya 503 6 478 6 5.4 0.1 d 5.0 0.1
Pre-treatment luteal activityb 469 14 439 9 5.3 0.1 d 5.0 0.0
Multiparous
No pre-treatment luteal activity 555 9 555 9 4.8 0.1 e 4.9 0.1
Pre-treatment luteal activity 544 9 526 11 4.9 0.2 e 4.8 0.1

Day 7 averaged 33 and 30 days postpartum for primiparous and multiparous cows, respectively. Values within a column
with different letters differ (d and e; P < 0.001). Progesterone-releasing devices were inserted on day 7 and removed on
day 0.
a Progesterone concentrations < 1 ng/mL on days 19 and 9.
b Progesterone concentrations 1 ng/mL on days 19 or 9.

concentration in younger cows remained significant when adjusted for differences in BW. Plasma
progesterone concentrations on day 0 also were analyzed for differences between cows that initi-
ated onset of estrous cycles in response to treatment and those that remained anestrus. Progesterone
concentrations were greater (P = 0.03) in the cows that initiated estrous cycles (2.9 0.3 ng/ml)
compared to those that did not (1.9 0.1 ng/ml). The parity by estrous cyclicity interaction was
non-significant.
Progesterone profiles following day 0 in cows without pre-treatment luteal activity are summa-
rized in Table 3 and illustrated in Fig. 1. The percentage of cows that initiated estrous cycles (i.e.,

Table 3
Treatment responses based upon plasma progesterone concentrations from days 5 to 30 following CIDR removal
Treatment Age (years) n Response, n (%)
Cycling Anestrus Spontaneous CL

PGF2 2 9 1 (11) 4 (44) 4 (44)

3 16 1 (6) 10 (63) 5 (31)
25 2 (8) a 14 (56) 9 (36)
CIDR + PGF2 2 11 9 (82) 2 (18) 0
3 18 7 (39) 10 (56) 1 (6)
29 16 (55) b 12 (41) 1 (3)
GnRH + CIDR + PGF2 2 9 6 (67) 1 (11) 2 (22)
3 15 8 (53) 5 (33) 2 (13)
24 14 (58) b 6 (25) 4 (17)
ECP + CIDR + PGF2 2 9 7 (78) 1 (11) 1 (11)
3 18 7 (39) 10 (56) 1 (6)
27 14 (52) b 11 (41) 2 (7)

Values within a column with different letters differ (a and b; P < 0.001). Response categories are defined in Section 2.
 J.E. Wheaton, G.C. Lamb / Animal Reproduction Science 102 (2007) 208216 213

formed a CL of normal duration within 12 days following PGF2 injection) was 44% (46/105) and
was influenced by treatment and parity. The treatment effect was due to a difference (P < 0.001) in
the percentages of cows that initiated estrous cycles within the PGF2 (8%, 2/25) group and those
in the CIDRPGF2 (55%, 16/29), GnRHCIDRPGF2 (58%, 14/24), and ECPCIDRPGF2
(52%, 14/27) groups. The parity effect (P = 0.025) was attributable to a greater percentage of
primiparous (61%, 23/38) than multiparous (34%, 23/67) cows that initiated estrous cycles. Luteal
phase plasma progesterone concentrations were similar in primiparous and multiparous cows from
days 5 to 30 (Fig. 1). Forty-one percent (43/105) of the cows remained anestrus to day 30. The
16 of 105 cows that neither initiated estrous cycles due to treatments nor remained anestrus had
spontaneous luteal activity. Five had short-lived CL and 11 ovulated spontaneously from dom-
inant ovarian follicles and formed a CL. These cows were spread across treatment groups and
parities.

Fig. 1. Plasma progesterone profiles in the 105 cows, regardless of age, that were anestrus on day 9. Day 0 corresponds
to the day that CIDR were removed and PGF2 was administered. Day 0 averaged 38-day postpartum. Forty-three cows
were categorized as remaining anestrus, 16 had spontaneous ovulations and formed a short-lived (n = 5) or late (n = 11)
CL, and 46 cows had ovulations in response to treatment and formed a CL of normal duration (estrous cycling).
214 J.E. Wheaton, G.C. Lamb / Animal Reproduction Science 102 (2007) 208216

4. Discussion

Results of the present study are similar to those of Fike et al. (1997) in that 7-day progesterone
administration via an intravaginal CIDR device increased the proportion of multiparous anestrous
suckled beef cows that ovulated and formed a CL of normal life-span following CIDR removal.
Increases were 44% in the study of Fike et al. (1997) and 43% for multiparous cows in the present
study. Present results further show that CIDR treatment is capable of inducing estrous cyclicity
in primiparous postpartum anestrous cows. The CIDR-induced increase was 76% in the younger
animals.
Two postpartum anestrous cows initiated estrous cycles subsequent to the single injection
of PGF2 (n = 25). Treatment with CIDRPGF2 (n = 29), GnRHCIDRPGF2 (n = 24), or
ECPCIDRPGF2 (n = 27)-induced estrous cyclicity in 55, 58, and 52% of the cows, respec-
tively. The similarity of response rates indicates that the addition of GnRH or ECP to CIDRPGF2
treatment may not have augmented the CIDRPGF2 response rate. Nonetheless, the conclusion
of little, if any, GnRH or ECP enhancement cannot be made with confidence due to the limited
number of animals. Gonadotropin-releasing hormone and ECP were tested because GnRH and
estrogen have been used in conjunction with CIDRPGF2 treatment to induce estrus and ovu-
lation in both cows with estrous cyclicity, and anestrous cows (Day, 1998; Lamb et al., 2001;
Stevenson et al., 2003; Rivera et al., 1998; Colazo et al., 2003). Estradiol cypionate is a long-
acting, oil-soluble ester of estradiol-17 that is a licensed product for the correction of anestrus
in cattle (Pfizer Animal Health).
Exogenous progesterone has been shown to elicit an increase in LH pulse frequency in post-
partum anestrous beef cows (Rhodes et al., 2002). Progesterone may decrease the number of
hypothalamic estradiol-17 receptors and thereby diminish the potency of estradiol-17 negative
feedback (Day and Anderson, 1998). Progesterone treatment applied to anestrous cows nearing
the onset of spontaneous estrous cyclicity may be most effective as theses cows have already
undergone a partial reduction in the potency of estradiol-17 negative feedback (Acosta et al.,
1983). Treatment of such cows with progesterone may further diminish the suppressive effect of
estradiol-17, enabling sufficient LH secretion to stimulate pre-ovulatory follicular development.
Day (1998) postulated from his review of literature that the primary variable that controls the
degree of response to progestogen treatment is the nearness in time to the spontaneous onset of
estrous cycles.
The greater estrous cyclicity response rate in primiparous than multiparous cows was incon-
sistent with the more problematic reproductive management of the first-calf cow. Postpartum
intervals were similar in primiparous and multiparous cows so this was not a contributing fac-
tor. Parity differences did exist in BCS at calving and in BW change from calving to the onset
of treatment. At calving, primiparous cows were in moderate to good body condition, whereas
multiparous animals were moderate to thin. Houghton et al. (1990) reported that cows that calve
with BCS of 4, 5, and 6 have anestrous intervals of 70, 60, and 52 days, respectively. Although
the difference between the mean BCS of multiparous (4.8) and primiparous (5.4) cows at calv-
ing was small, it was highly significant. Based upon the percentages of multiparous (52%) and
primiparous (21%) cows that remained anestrus until the end of the present study (about 70 days
postpartum), multiparous cows would have had longer postpartum intervals than primiparous
cows. Longer postpartum intervals in multiparous than primiparous cows is consistent with the
tendency (P = 0.08) towards a lesser rate of pre-treatment luteal activity in multiparous than prim-
iparous cows. Primiparous cows were evidently nearer in time to the onset of spontaneous luteal
activity than multiparous cows.
 J.E. Wheaton, G.C. Lamb / Animal Reproduction Science 102 (2007) 208216 215

Plasma progesterone concentrations were less in multiparous than primiparous cows on the
day of CIDR removal (day 0). The cause of the difference is uncertain. It may have resulted
from physical differences in body size and in the size and flaccidity of the reproductive tract, or
from differences in metabolic factors, such as feed intake and milk production. The latter may
have altered the rate of progesterone clearance (Butler, 2000). Progesterone concentrations on
day 0 were analyzed retrospectively to determine whether they differed in cows that subsequently
initiated estrous cycles in response to the three CIDRPGF2 treatments or remained anestrus.
Concentrations were greater in cows that initiated estrous cycles compared to those that remained
anestrus. Most (15/17, 88%) multiparous and primiparous cows that had a progesterone concentra-
tion >3 ng/ml on day 0 cycled, whereas in cows with 3 ng/ml there was no apparent relationship
between the progesterone concentration and the subsequent estrous cyclicity response.
Fifteen cows had plasma progesterone concentrations 1 ng/ml on day 19 or 9. These cows
were categorized as having pre-treatment luteal activity and their data were subsequently removed
from analyses of treatment effects. In the 25 anestrous cows not receiving a CIDR on day 7
and administered PGF2 on day 0, four had elevated progesterone concentrations on day 0. These
cows expressed spontaneous onset of luteal activity sometime between days 9 and 0. Following
PGF2 administration, two of the four animals had a typical estrous cycle, one formed a short-
lived CL and the other was anestrus. Presuming that the same percentage of cows (16%, 4/25)
would have expressed spontaneous luteal activity during the period of CIDR treatment in the 80
cows that received a CIDR, then 13 cows would have been expected to express spontaneous luteal
activity while CIDR were in place. A further presumption is that 6 or 7 of the 13 cows would have
initiated estrous cycles in response to PGF2 administration. Thus, the estrous cyclicity response
rate following CIDRPGF2 treatment of 55% (44/80) represents both CIDRPGF2 -induced
estrous cyclicity and synchronization of ovulation. An estimate of the latter is 15% (6.5/44).
Results of the present study show that CIDRPGF2 treatment initiated approximately 31-day
postpartum stimulated/synchronized estrous cyclicity in 76% of primiparous and 43% of multi-
parous beef cows that were anestrus near the time of CIDR insertion. Treatment with CIDRPGF2
provides an effective reproductive management tool to increase the proportion of cows initiating
estrous cycles by the start of the breeding season.

Acknowledgements

The authors acknowledge the assistance of Theresa Anttila and Dan Brown in planning and
conducting the experiment. Appreciation also is expressed to Dr. J.R. Chenault, Pharmacia Animal
Health, for providing CIDR, Lutalyse, and ECP.

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