Prostaglandins and Reproduction Infemale Animals

Prostaglandins and reproduction in female farm
animals The Veterinary Journal172:206-228, 2006.

C.W. Weemsa, , Y.S. Weemsa and R.D. Randelb
a
Department of Human Nutrition, Food, and Animal Sciences, University of Hawaii,
1955 East West Road, Honolulu, HI 96822, USA
b
Agricultural Research and Extension Center, Texas A&M University, Overton, TX
75684, USA
Accepted 20 November 2004. Available online 26 January 2005.

172:206-228
Abstract
Prostaglandins impact on ovarian, uterine, placental, and pituitary function to regulate
reproduction in female livestock. They play important roles in ovulation, luteal function,
maternal recognition of pregnancy, implantation, maintenance of gestation, microbial-
induced abortion, parturition, postpartum uterine and ovarian infections, and resumption
of postpartum ovarian cyclicity. Prostaglandins have both positive and negative effects on
reproduction; they are used to synchronize oestrus, terminate pseudopregnancy in mares,
induce parturition, and treat retained placenta, luteinized cysts, pyometra, and chronic
endometritis. Improved therapeutic uses for prostaglandins will be developed when we
understand better their involvement in implantation, maintenance of luteal function, and
establishment and maintenance of pregnancy.
Keywords: Prostaglandins; Reproduction; Corpus luteum; Pregnancy; Placenta;

Ovulation; Implantation; Parturition; Postpartum; Oestrous synchronization
No se pude mostrar la imagen v inculada. Puede que se hay a mov ido, cambiado de nombre o eliminado el archiv o. Compruebe que el v ínculo señala al archiv o y ubicaciones correctos.
Article Outline
1. Introduction
2. Prostanoid metabolism
3. Follicular growth and ovulation
4. Luteolysis
5. Implantation
6. Maternal recognition and maintenance of pregnancy
7. Parturition
8. Postpartum period
9. Synchronization of oestrus
Acknowledgements
References
1. Introduction
Prostaglandins (PGs) affect ovulation, luteal regression, the implantation and
maintenance of pregnancy, parturition, postpartum physiology, and have been used for
synchronization of oestrus alone or with progestins, oestrogens, and gonadotropin
releasing hormone (GnRH). To understand PGs and reproduction, knowledge of their
metabolism is important.
2. Prostanoid metabolism
Prostanoids (comprising the PGs, leukotrienes [LT], and thromboxanes [TX]) are 20-
carbon molecules derived from arachidonic acid (AA) or di-homo-3b3;-linolenic acid.
Most PGs (PGF23b1;, PGI2, PGE2) and TXA2 have double bonds at carbons 52013;6 and
132013;14 and are derived from AA, while PGE1 has a double bond only at carbons
132013;14 and is derived from di-homo-3b3;-linolenic acid. Double bonds at carbons
132013;14 and an OH group at C-15 are essential for biological activity (Bergstrom et al.,
1968, Schneider, 1976 and Ramwell et al., 1977).
Leukotrienes are derived from AA, but unlike PGs and TXA do not contain a 5 or 6
member ring (Smith and DeWitt, 1996). Catabolism of PGs can occur within organs, but
most occurs in the lung (Piper et al., 1970). This involves reducing the double bond at
carbons 132013;14 and dehydrogenation of the OH group on C-15 to make the 13-14-
dihydro-15-keto-PG catabolite (Samuelsson et al., 1971). Cyclo-oxygenase (COX) occurs
as a constitutive (COX-1) and an inducible (COX-2) enzyme for both PG and TXA
synthesis. Arachidonic acid is converted by COX to PGG 2 and then to PGH2, which is
then converted by synthases or reductases to a specific PG or TXA (Smith and DeWitt,
1996).
Pharmaceutical agents influence PG metabolism. Aspirin or indomethacin inhibits COX-

1 and celecoxib inhibits COX-1 and COX-2, while cortisol blocks phospholipase-A 2
release of AA from phospholipids to decrease PG synthesis (Vane and Botting, 1995).
Inhibitors of PG synthesis are currently used for understanding reproductive processes,
but not for clinical therapy (Murdoch, 1988b and Bridges et al., 1999a). Little
information exists on downstream synthases or reductases after PGH 2 formation on
reproduction. Lipoxygenases synthesize LT from AA, which can be inhibited by 1-
benzo-[b]-thien-2-ylethyl-1-hydroxyurea (Vane and Botting, 1995). Various isoforms of
prostaglandin receptors (PG receptors) have been identified in tissues: receptors with low
and high affinity binding sites for PGF23b1; (FP); four isoforms for PGE1 and PGE2 (EP1,
EP2, EP3, EP4), and only one isoform each for TXA2, PGI2, and PGD2 (Narumiya, 1995).
Effects of PGE1, PGE2, and PGF23b1; on reproduction have been studied, but other PGs
may also have roles (Weems et al., 1995).
3. Follicular growth and ovulation

Whether PGs have direct effects on follicular growth prior to the luteinizing hormone
(LH) surge to initiate ovulation in livestock is unknown, but PGF 23b1; increases pituitary
responsiveness to GnRH to release LH in the postpartum cow (Randel et al., 1996).
However, PGs produced by the ovulatory follicle are obligatory for ovulation (Murdoch
et al., 1993). Ovulation involves a cascade of changes in follicles and most of the
mechanisms for ovulation in livestock have used the sheep model.
In sheep, luteolysis is induced by PGF23b1; and GnRH is used to time the LH surge for
collection of ovulatory follicles. Ovulation occurs 24 h after the LH surge (Murdoch,
1985). The LH surge through alterations in 32032;52032; cyclic adenosine
monophosphate (c-AMP) levels shifts steroidogenesis in the ovulatory follicle.
Androgens and oestradiol-173b2; levels decrease and progesterone levels increase, which
results in changes in microcirculation and morphology in the follicle wall (Murdoch and
Dunn, 1982a, Murdoch and Dunn, 1982b, Murdoch et al., 1983, Murdoch, 1988a,
Murdoch, 1992 and Cavender and Murdoch, 1988). Increases in c-AMP occur in the
follicle wall 4 h after the LH surge and remain elevated for 12 h, but decline by
162013;20 h (Murdoch et al., 1981 and Murdoch et al., 1986). The area of theca interna
occupied by blood vessels increases at 4 and 12 h after the LH surge and declines at
ovulation to levels lower than those prior to the LH surge, suggesting ischaemia may be a
factor in ovulation (Murdoch and Myers, 1983, Murdoch and Cavender, 1987 and
Murdoch and Cavender, 1989). Basophil/mast cell numbers in the theca interna decrease
by 4 h after the LH surge and neutrophils and eosinophils in ovulatory follicles increase
4 h prior to ovulation (Murdoch et al., 1981, Murdoch et al., 1986 and Murdoch and
Steadman, 1991). Follicular fluid volume increases and antihistamines given at the LH
surge decrease follicular fluid volume, but do not inhibit ovulation. However, subsequent
luteal function is decreased (Halterman and Murdoch, 1986 and Murdoch, 1990).
In sheep, concentrations of PGE2 and PGF23b1; in follicular fluid do not change after the
LH surge (Murdoch et al., 1981, Murdoch et al., 1986 and Murdoch et al., 1993). In the
follicle wall, COX-2 and both PGE2 and PGF23b1; increase at 8 h and remain elevated
through 12 h, and only PGE2 decreases by 162013;20 h (Murdoch et al., 1981, Murdoch
et al., 1986, Murdoch et al., 1991 and Murdoch et al., 1993). Intrafollicular injections of
LH or follicle stimulating hormone (FSH) in sheep increased PGE 2, but not PGF23b1;, in
granulosa cells, while both PGE2 and PGF23b1; were increased in the theca (Murdoch and
McCormick, 1991 and Murdoch et al., 1993). Intercellular space between mural and
cumulus cells increases by 12 h and is dependent on PGE 2, but not PGF23b1; (Murdoch et
al., 1993).
Indomethacin inhibition of ovulation can be overcome by PGE 2 or PGF23b1; (Murdoch,

1988b, Murdoch and Dunn, 1982b, Murdoch and McCormick, 1991 and Murdoch et al.,
1993). The theca converts PGE2 to PGF23b1; to activate collagenolysis, which is
dependent on follicular progesterone regulation of 9-keto-PGE 2-reductase to convert
PGE2 to PGF23b1; (Murdoch and Ferris, 1988 and Murdoch et al., 1993). Isoxazol, an
inhibitor of progesterone synthesis, also blocks ovulation, which can be restored by
progesterone or PGF23b1;, but not by PGE2 (Murdoch and Cavender, 1987 and Murdoch et
al., 1993).
Collagenolysis increases by 20 h after the LH surge and until follicular rupture (Murdoch
and McCormick, 1991, Murdoch and McCormick, 1992a and Murdoch, 2000).
Collagenolysis is not affected by isoxazol or indomethacin 16 h after the LH surge;
however, indomethacin or isoxazol decreases collagenolysis by 24 h (Murdoch et al.,
1993). Leukocyte infiltration into the follicle wall may be caused by a chemoattractant
produced by collagenolysis (Murdoch and McCormick, 1989, Murdoch and McCormick,
1991, Murdoch and McCormick, 1992a, Murdoch and McCormick, 1992b and Murdoch
and McCormick, 1993). Collagen, collagen -like peptides, LTB4 (a prostanoid), tumor
necrosis factor-3b1; (TNF3b1;), platelet activating factor (PAF), TXA2, and plasmin
increase leukocyte infiltration in the ovulatory follicle, while antisera to collagen-like
peptides prevents leukocyte infiltration (Murdoch, 1986, Murdoch, 1998a, Murdoch,
1999, Murdoch and McCormick, 1989, Murdoch and McCormick, 1991, Murdoch and
McCormick, 1992a, Murdoch and McCormick, 1992b, Murdoch and McCormick, 1993,
Alexander et al., 1990, Murdoch, 1997, Murdoch and Lund, 1999 and Gottsch et al.,
2000).
Nordihydroguaratic acid, a lipoxygenase inhibitor, decreases LT and leukocyte

infiltration into the follicle and blocks ovulation, although indomethacin also reduced
leukocyte infiltration (Espey, 1992, Carvalho et al., 1989 and Murdoch and McCormick,
1991). Remodeling of the extracellular matrix may require other proteases at ovulation,
which may be activated by TNF3b1; (Murdoch, 1999, Murdoch et al., 1999c, Johnson et
al., 1999a and Medan et al., 2003). Granulosa cell organelle arrangement, chromosomes
of follicular cells, and membrane potential of cells at the site of ovulation are altered by
PGs, resulting in local necrosis (Ackerman and Murdoch, 1993, Murdoch, 1994a,
Murdoch, 1994b, Murdoch, 1995a, Murdoch, 1995b, Murdoch, 1996a, Murdoch, 1996b,
Murdoch, 1996c, Murdoch, 1998a, Murdoch, 1998b, Murdoch, 1999, Murdoch, 2000,
Murdoch and Lund, 1999, Murdoch et al., 1999a, Murdoch et al., 1999b and Murdoch et
al., 1999c; Murdoch and Van Kirk, 2002; Murdoch and McDonnel, 2002).
As is observed in ewes, in the sow, cow, and mare, changes in c-AMP, steroids, LTs, and
PGs in the ovulatory follicle wall occur, but PGs may also increase in follicular fluid
(Tsang et al., 1979, Tsang et al., 1988, Evans et al., 1983, Ainsworth et al., 1984,
Ainsworth et al., 1990, Hunter and Poyser, 1985, Watson and Hinrichs, 1988, Watson
and Sertich, 1991, Algire et al., 1992, Murdoch et al., 1993, Liu et al., 1997 and Downey
et al., 1998). Indomethacin also blocks ovulation in the sow, cow, and mare (Ainsworth
et al., 1979, Downey and Ainsworth, 1980, DeSilva and Reeves, 1985 and Watson and
Sertich, 1991) and lipoxygenase inhibitors suppress porcine ovulation (Downey et al.,
1998).
Bovine granulosa cells produce both PGE2 and PGF23b1; (Bridges and Fortune, 2003).
Two increases in mRNA for progesterone receptors is induced by the LH surge in
granulosa and theca cells (Jo et al., 2002). Progesterone regulates follicular oxytocin
(OT), PG synthesis enzymes, and FP and EP 3 receptors (Sirois, 1994 and Tsai et al.,
1996). Whether this occurs through the protein kinase A or C (PKA; PKC) pathways is
unclear (Sirois, 1994 and Tsai et al., 1996). In cows, endothelin-1 (ET-1) and TNF3b1; can
also stimulate preovulatory granulosa cell PGE2 secretion (Acosta et al., 1998).
4. Luteolysis
After ovulation, the corpus luteum develops from the remainder of the follicle wall.
Granulosa cells differentiate into large luteal cells (LLC), which comprise 30% of the
steroidogenic cells, and secrete 70% of the progesterone, and LH does not stimulate LLC
progesterone secretion. The theca differentiates into small luteal cells (SLC), which
comprise 70% of the steroidogenic cells. SLC require LH stimulation for maximal
progesterone secretion, but secrete only 30% of the progesterone (Farin et al., 1989 and
Niswender et al., 1994). Loss of progesterone secretion by the corpus luteum occurs
before morphological regression (Ginther, 1974 and O2019;Shea and McCoy, 1988).
The primary site for initiation of luteolysis is through the LLC (Niswender et al., 1994).
Ovine luteolysis starts between days 112013;13 and between days 162013;17 post-
oestrus in the cow, sow, and mare (Ginther, 1967 and Ginther, 1974). Evidence of
luteolysis is seen by days 142013;15 post-oestrus in sheep and by days 172013;19 in the
sow, cow and mare with decreased jugular progesterone, decreased luteal weight, reduced
LLC and SLC size, and disruption of luteal vasculature (O2019;Shea and McCoy, 1988,
Geisert et al., 1992, Niswender et al., 1994, Niswender et al., 2000, Bazer et al., 1998 and
Kobayashi et al., 2001). Disruption of luteal vasculature may be via angiopoietin-2
Stouffer et al., 2001).
Understanding luteolysis requires an understanding of the anatomical relationships

between the ovary, uterus, and their associated vasculature. Hysterectomy early after
ovulation or autotransplantation of the ovary or uterus alone to the neck in sheep with
vascular connections to the jugular vein and carotid artery prevents luteal regression up
to, but not beyond, gestation for that species (Goding et al., 1972 and Ginther, 1974). The
ovine uterus must remain in situ throughout luteolysis, since hysterectomy on day 15
delays luteolysis for a few days (Moor et al., 1970). Removal of the uterine horn adjacent
to the luteal-containing ovary of ewes or cows delays luteolysis, while removal of the
opposite uterine horn does not. This indicates that the uterus secretes a luteolysin, which
is transferred locally from the uterine horn to the adjacent ovary (Ginther, 1974 and
Ginther, 1981).
The source of the luteolysin is the endometrium. Destruction of the endometrium or

ligation of the uterine vein adjacent to the luteal-containing ovary delays luteolysis
(Ginther, 1974). Infusion of mid-cycle endometrial extracts into the ovarian artery
hastens luteolysis (Caldwell and Moor, 1970). The luteolysin is secreted into the uterine
vein and is transferred locally to the ovarian artery of the luteal-containing ovary (ewe,
cow, sow), but delivery in mares is via systemic circulation, since little contact occurs
between the uterine vein and ovarian artery (Ginther, 1974).
An intrauterine device (IUD) adjacent or opposite to the side of ovulation on days

12013;2 post-oestrus hastens luteolysis in ewes. An IUD inserted on days 32013;4 post-
oestrus cause premature luteolysis only when placed in the uterine horn adjacent to the
luteal-containing ovary (Ginther, 1974). The luteolysin can be delivered between uterine
horns via venous anastomoses (Ginther, 1974). Further support for local delivery of the
luteolysin in the ewe and cow comes from cross-circulation studies using vascular
anastomoses combined with unilateral hysterectomy. Removal of the uterine horn
adjacent to the luteal-containing ovary delays luteolysis and prevents an IUD-induced
precocious luteolysis by the opposite uterine horn in sheep. If the uterine vein or ovarian
artery opposite to the side of hysterectomy is diverted to corresponding vessels on the
hysterectomized side, luteal function is not extended, with or without an IUD, in the
remaining uterine horn (Ginther, 1974).
Endometrial PGF23b1; secretion initiates luteolysis. Uterine venous PGF23b1; first increases
on days 112013;13 in ewes, days 132013;14 in sows, and days 162013;17 post-oestrus in
cows (Wilson et al., 1972a, Gleeson et al., 1974, Weems et al., 1975, Pexton et al., 1975a,
Pexton et al., 1975b, Pexton et al., 1975c, Weems, 1979, Christenson et al., 1994,
Thatcher et al., 1995 and Thatcher et al., 2001). Only 1 mg of PGF23b1; given
intramuscularly (IM) is required to induce luteolysis in the mare, but 25 mg of PGF 23b1;
given IM is required to induce luteolysis in cows (Douglas and Ginther, 1972 and
Ginther, 1981). This difference is presumed to be due to less catabolism by the lungs in
the mare, since PGF23b1; is destroyed by one pass through the lungs in most species (Piper
et al., 1970).
In sows, PGF23b1; is delivered locally from the uterus to the adjacent ovary, but is also
delivered by the systemic circulation (Niswender et al., 1970 and Ginther, 1974). In
cows, jugular venous progesterone decreases to less than 1 ng/mL within 4 h after luteal
resection, which is similar to PGF23b1;-induced luteolysis (Fogwell et al., 1978). This
indicates that PGF23b1;-induced luteolysis is a pharmacological effect, since progesterone
decreases slowly during natural luteolysis and PGF23b1; secretion is pulsatile, especially
on day 15 post-oestrus in sheep ( Zarco et al., 1988a). Bovine luteal tissue is usually not
responsive to PGF23b1; until after day 4 post-oestrus, although receptors for PGF23b1; are
present and responsiveness increases up to day 12 post-oestrus (Wiltbank et al., 1995 and
Weems et al., 1995). Sows are not responsive to LH or PGF23b1; until day 12 post-oestrus
(Gadsby et al., 1990 and Diaz et al., 2000).
Indomethacin or destruction of follicles, the source of oestrogen, at midcycle delays

luteolysis (Inskeep and Murdoch, 1980 and Fogwell et al., 1985). An IUD causes earlier
increases in uterine PGF23b1; secretion to shorten the inter-oestrous interval, which is
associated with oestrogen sensitization of the uterus (Pexton et al., 1975a). Oestradiol-
173b2; given IM at midcycle causes premature luteolysis in intact, but not in
hysterectomized, ewes or cows, (Ford et al., 1975). Initiation of uterine PGF23b1; secretion
in ruminants is triggered by oestradiol-173b2; after prior exposure to progesterone (Ford et
al., 1975).
Progesterone, oestradiol-173b2;, LH, and oxytocin (OT) are probably involved in uterine
PGF23b1; secretion through both COX-1 and COX-2 (Fogwell et al., 1985, Eggleston et
al., 1990 and Bazer et al., 1992). Indomethacin, a COX-1 inhibitor, delays luteolysis
(Fogwell et al., 1985).Uterine PGF23b1; secretion is increased by OT, oestrogen increases
the response to OT, and indomethacin inhibits OT-induced PGF 23b1; secretion (Sharma
and Fitzpatrick, 1974 and Silvia et al., 1991). Luteal OT may be responsible for uterine
PGF23b1; pulsatile secretion. Luteal OT is released by PGF23b1; indicating that a positive
feedback loop exists between the uterus and corpus luteum, which is modulated by
progesterone and oestradiol-173b2; (Silvia et al., 1991, Mirando et al., 1993, Carnahan et
al., 1996, Mann et al., 1998, Hu et al., 2001 and Okuda et al., 2002). However, bovine
luteal OT depletion or OT receptor antagonists do not delay luteolysis (Jaroszewki and
Kotwica, 1994, Kotwica et al., 1997 and Kotwica et al., 1999).
Although ovine endometrial progesterone receptors decline by day 12 post-oestrous,

progesterone receptor antagonists delay luteolysis (Morgan et al., 1993), possibly by
inactivating the endometrial PKC pathway (Whiteaker et al., 1995). Alternatively,
progesterone may modulate endometrial PGF23b1; secretion through COX prior to
decreases in progesterone receptors (Fogwell et al., 1985). Whether this is through COX-
1, COX-2, or both is unknown. Endometrial LH receptors may play a role in PGF 23b1;
secretion (Freidman et al., 1995 and Ziecik et al., 1986). Endometrial LH receptors
increase during luteolysis and LH increases PGF23b1; secretion late in the oestrous cycle
and may enhance luteolysis (Guthrie and Bolt, 1983, Stepien et al., 1999, Ziecik, 2000,
Ziecik, 2002, Ziecik et al., 2001, Stepien and Ziecik, 2002 and Shemesh et al., 2002).
Luteal progesterone secretion is regulated by LH via cAMP during the oestrous cycle.
Although PGF23b1; is a vasoconstrictor and reduces blood flow to the corpus luteum; loss
of luteal LH receptors and decreases in ovarian blood flow occur after the onset of
luteolysis and both cAMP and progesterone decline (Ford, 1984, Ford, 1995 and
Niswender et al., 1994). Whether PGF23b1; affects 3b2;-adrenergic receptor-stimulated
luteal progesterone secretion during luteolysis is unknown (Skarzynski and Okuda,
2000). Luteolytic effects of PGF23b1; occur via luteal membrane PGF23b1; receptors
(Gadsby et al., 1990, Pitzel et al., 1993, Niswender et al., 1994, Sakamoto et al., 1995,
Wiltbank et al., 1995 and Tsai et al., 2001).
Initiation of luteolysis is through PGF23b1; increases in cytosolic calcium and the LLC
phosphatidyl inositol PKC pathway (Wiltbank et al., 1991 and Weipz et al., 1993). The
PKC pathway is not involved in PGF23b1;-induced luteolysis in pigs (Christenson et al.,
1995). Luteolytic actions of PGF23b1; are through inhibition of lipoprotein uptake to limit
intracellular cholesterol transport in luteal cells and progesterone decreases before
steroidogenic enzymes decrease (Grusenmeyer and Pate, 1992, Rodgers et al., 1995 and
Niswender et al., 2000). The corpus luteum may participate in its own demise. Treatment
with PGF23b1; in vivo causes ovine luteal tissue incubated in vitro to secrete PGF 23b1;,
which may complete luteolysis (Tsai and Wiltbank, 1997a). Physiologic and morphologic
luteal regression may be facilitated through PG activation of the peroxisome proliferator
system to induce transcription factors in LLC (Viergutz et al., 2000).
Caution should be exercised when interpreting data on luteolysis with bovine luteal tissue
in vitro, since experiments with PGF23b1; produce conflicting results. Treatment of bovine
luteal cell suspensions with PGF23b1; in vitro increases progesterone before progesterone
decreases (Hansel et al., 1991). This may be due to conversion of PGE2 to PGF23b1; by 9-
keto-PGE2-reductase or disruption of gap junctions between SLC and LLC (Beaver and
Murdoch, 1992 and Grazul-Bilska et al., 1997). Interaction between SLC and LLC and
other luteal cells may be both luteotropic and luteolytic (Girsh et al., 1995). More
progesterone is secreted when SLC and LLC are co-cultured than the combined secretion
of progesterone by similar numbers of SLC and LLC cultured separately (Niswender et
al., 1994).
Species differences and reproductive status also affect luteal PG secretion in vitro.
Bovine luteal slices secrete PGF23b1; and PGE2 in vitro, which increases linearly with
time and the ratio is 1:1 (Weems et al., 1998b and Weems et al., 2002a). Ovine mid-cycle
luteal tissues do not secrete detectable PGE2 and PGF23b1; in vitro (Weems et al., 1997
and Kim et al., 2001).
In sows and cows, monocyte chemoattractant protein-1 causes macrophages and

leukocytes to invade luteal tissue (Hehnke et al., 1994, Tsai et al., 1997b, Penny, 2000,
Bauer et al., 2001 and Cavicchio et al., 2002). Macrophages and leucocytes secrete
cytokines and can activate luteal tissue cytokine secretion. Cytokine and their receptors
are expressed by luteal tissue for TNF3b1;, interleukin-13b2; (IL-13b2;), ET-1, insulin growth
factor binding protein-3, interferon (INF)-3b3;, and basic fibroblast growth factor during
luteolysis. These cytokines may enhance luteal PG secretion, apoptosis, and luteal matrix
remodeling (Richards and Almond, 1994a and Pate, 1995; Wuttke et al., 1998, Nicholson
et al., 1999, Penny et al., 1999, Friedman et al., 2000, Pitzel et al., 2000, Petroff et al.,
2001 and Davis and Rueda, 2002).
Receptors for TNF3b1; are detected on luteal tissue throughout the oestrous cycle and
early pregnancy and TNF3b1; stimulates bovine luteal PGF23b1; and PGE2 secretion, but at
different doses (Sakumoto et al., 2000, Miyamoto et al., 2002 and Okuda et al., 2003). In
ewes and cows, luteolysis has been reported to also require ET-1 (Meidan et al., 1999 and
Hinckley and Milvae, 2001). Receptors for ET-1 and ET-1 increase in bovine luteal
tissue by day 1 post-oestrous, increase up to day-10, and increase further after treatment
with PGF23b1;, but only during luteolysis (Flores et al., 2003).
Nitric oxide (NO) has been reported to play a role in luteolysis, since L-nitro-arginine
methyl ester (L-NAME), a NO synthase (NOS) inhibitor, delivered into the bovine
corpus luteum in vivo delayed decreases in progesterone (Jaroszewki and Hansel, 2000
and Skarzynski et al., 2003). However, ET-1, or NO donors alone, do not alter bovine
luteal tissue progesterone or PGF23b1; secretion in vitro (Weems et al., 2004a) and NO
inhibits PGF23b1;-induced luteolysis in vitro (Dong et al., 1999). Some cytokines, which
are increased in luteal tissue after the initiation of luteolysis, may be to resist luteolysis
rather than being involved in the luteolytic process.
Other molecules can also affect luteolysis. Inhibitors of LT synthesis delay luteolysis, by
unknown mechanisms (Cooke and Ahmad, 1994 and Hansel and Blair, 1996). Hydrogen
peroxide (H2O2) and GnRH are luteolytic and are as effective as PGF23b1; in vitro on rat
luteal cells (Behrman and Preston, 1989). Data on H2O2 and luteolysis in livestock are not
available, and luteal tissue GnRH receptors in livestock have not been detected (Brown
and Reeves, 1983). A GnRH-like protein in luteal and follicular tissue of ewes or cows
decreased LH-stimulated rat luteal progesterone secretion in vitro (Aten et al., 1987). It is
90% homologous to histone H2A. Histone H2A increased endometrial PGF23b1; secretion
in vitro (Aten and Behrman, 1989 and Weems et al., 1996).
5. Implantation
Oviductal blastocyst transport and development may be regulated by angiotensin II, atrial
natriuretic peptide, LH, progesterone, and oestradiol-173b2; stimulation of oviductal ET-
1 and PG secretion (Wijayagunawardane et al., 1998, Wijayagunawardane et al., 1999
and Wijayagunawardane et al., 2001). Horse embryos secrete PGE2 to permit entrance of
the embryo into the uterus, since unfertilized horse oocytes do not enter the uterus
(Weber et al., 1992). Implantation in rats occurs rapidly on day 4 post-breeding where
implantation is invasive, induces placental decidual tissue, and uterine weight increases
from a few hundred mg to grams within a few days (Weems et al., 1975 and Yee and
Kennedy, 1991).
Blastocyst implantation in livestock is gradual, occurs over weeks, and the endometrium
is not invaded (Weems et al., 1995 and Bazer et al., 1998). Caruncular weight in sheep
and cows increases during placentome formation during pregnancy, which is similar to
the increased decidual tissue weight during pregnancy in rats (Weems et al., 1975 and
Yee and Kennedy, 1991).
Endometrial stromal cells treated with human chorionic gonadotropin (hCG) differentiate
into decidual tissue (Han et al., 1999). In rats, oestradiol-173b2;, PGE2, PGI2, leukemia
inhibitory factor, epidermal growth factor (EGF) and LT appear to be involved in
implantation (Yee and Kennedy, 1991, Kennedy and Ross, 1993, Cheon et al., 2002 and
Lim et al., 2002). Oestradiol-173b2; increases on day 4 post-breeding to sensitize the
endometrium for implantation in rats. Implantation is inhibited by oestrogen receptor
antagonists and in the oestrogen receptor knock-out mice (Kennedy and Ross, 1993,
Cheon et al., 2002, Salamonsen et al., 2001 and Lim et al., 2002).
In rats, oestrogen regulates PGE2 production by the endometrium during implantation and
inhibition of PG synthesis prevents implantation, which is restored with PGE 2 (Yee and
Kennedy, 1991 and Kennedy and Ross, 1993). In rodents, EGF or IL-1 regulates PGE2
secretion and the peroxisome proliferator activated receptor (PPAR)394; can activate
PGI2 receptors (Lim et al., 2002). How LT are involved in implantation is unknown
(Cheon et al., 2002). Nitric oxide may regulate PG secretion for implantation, since NOS
inhibitors prevent implantation in rodents (Novaro et al., 1996, Chwalisz et al., 1999,
Chwalisz and Garfield, 2000, Saxena et al., 2000 and Tranguch et al., 2002). Placental
NOS increases on days 202013;40 in pigs (Self et al., 2003.)
Livestock blastocyst hatching and development may involve PG and LH (Scenna et al.,
2002 and Mishra et al., 2003). Embryos secrete PG, and LH influences embryonic
development, possibly through stimulating PG secretion (Marcus, 1981, Lewis et al.,
1982, Davis et al., 1983, Gandolfi et al., 1992 and Mishra et al., 2003). Embryos with low
cleavage rates secrete less PGE2. Addition of PGE2 increases bovine embryo cleavage
rates (Gurevich et al., 1993). Cow and sow, but not sheep, embryos secrete oestrogen,
which increases uterine blood flow during implantation (Ford, 1984 and Ford, 1995). In
sheep, circulating oestrogen increases as pregnancy progresses (Findlay and Cox, 1970)
and 90% is from the placenta by day 90 of pregnancy when all placentomes are formed
(Weems et al., 1994a and Weems et al., 1995).
In pigs, oestrogen regulates endometrial stromal EGF production, which stimulates PGE 2
secretion (Zhang and Davis, 1991). During implantation in sheep, the endometrium
secretes PGE2 (Beaver and Murdoch, 1992). Livestock embryos secrete a plethora of
prostaglandins, proteins, steroids, and growth factors, which may regulate endometrial or
blastocyst remodeling for implantation or maternal immunosuppression (Lewis et al.,
1982, Ford, 1984, Ford, 1995, Ott et al., 1998, Johnson et al., 1999b, Johnson et al., 2001,
Gray et al., 2001, Spencer and Bazer, 2002 and Choi et al., 2003).
6. Maternal recognition and maintenance of pregnancy

Embryos must be in the uterine horn adjacent to the luteal-containing ovary by day 12 in
ewes and day 16 post-oestrus in the cow, sow, and mare to prevent luteolysis (Moor and
Rowson, 1964, Niswender et al., 1970 and Ginther, 1974). Ovine embryos transferred on
day 13 fail to prevent luteolysis (Moor, 1968). Ovine or porcine embryo homogenates
infused into the uterine lumen extend luteal function (Rowson and Moor, 1967 and Ball
and Day, 1982). In sheep with a corpus luteum on each ovary, but an embryo in only one
uterine horn and the uterine horns separated surgically, only the corpus luteum adjacent
to the embryo-containing horn is maintained (Moor and Rowson, 1964).
Diversion of uterine venous or ovarian arterial blood from the embryo-containing side to
vasculature on the sterile side maintains luteal function adjacent to the sterile horn,
indicating an antiluteolysin is secreted into the uterine vein and is then transferred locally
to the ovarian artery (Ginther, 1974 and Ginther, 1981).
Local effects of an ovary on the adjacent uterine horn also occur. Progesterone is
delivered locally and systemically to the uterus of cows and ewes (Weems et al., 1988
and Weems et al., 1989b).This may explain why more cows become pregnant when
embryos are transferred to the anterior third of the uterine horn adjacent to the luteal-
containing ovary than to other regions of the uterus (Sreenan, 1976). These local vascular
routes also have negative effects on fertility, presumably via PG. Pregnancy rates in
pubertal ewe lambs to a single mating are about 28% until they reach 18 months of age
(Trapp and Slyter, 1979). However, removal of the oviductal vein adjacent to the luteal-
containing ovary on day 5 post-breeding increases pregnancy rates to those seen in
mature ewes, indicating a negative effect of the ovary on conception rates through a local
vascular route in young ewe lambs (Vincent et al., 1992).
At the first postpartum ovulation in cows, an embryo-toxic factor is secreted by the

corpus luteum, which is thought to be a PG delivered locally from an ovary to the
adjacent uterine horn, resulting in decreased postpartum fertility in cows. Fertility is
improved by inhibitors of PG synthesis (Inskeep, 1995).
Establishment and maintenance of pregnancy is dependent on changes in a plethora of

hormones. Changes in some of these hormones may result in pregnancy loss at any time
during pregnancy (Schrick and Inskeep, 1993, Geisert et al., 1994 and Weems et al.,
1995). Whether embryos suppress uterine PGF23b1; secretion to prevent luteolysis is
debatable (Weems et al., 1985b, Weems et al., 1992b, Weems et al., 1995, Bazer et al.,
1991 and Bazer et al., 1998). Endometrial PGF23b1; concentrations are not decreased
during early ovine pregnancy and PGF23b1; secretion is not reduced until the end of the
first trimester (Wilson et al., 1972b, Marcus, 1981, La Croix and Kann, 1982 and Weems
et al., 2003b).
Concentrations of PGF23b1; in uterine venous, or ovarian arterial blood, or in luteal tissue

during early pregnancy are not decreased in the ewe, cow, or sow (Pexton et al., 1975b,
Pexton et al., 1975c, Lewis et al., 1978, Ellinwood et al., 1979a, Ellinwood et al., 1979b,
Inskeep and Murdoch, 1980, Lukaszewska and Hansel, 1980, Silvia et al., 1984a, Silvia
et al., 1984b, Ottobre et al., 1984, Rawlings and Hyland, 1985, Stefanczyk et al., 1990
and Weems et al., 1995). Transport of PGF23b1; across the utero-ovarian vascular plexus
or luteal membranes of cows or binding to luteal membrane receptors is not decreased
during early pregnancy (Weipz et al., 1992 and Banu et al., 2003). Results of other
studies suggest that the number of peaks of PGF23b1; in jugular venous plasma during
early pregnancy is suppressed by INF-3c4; secreted by embryos to prevent luteolysis in
ewes and cows, based on the catabolite, 13-14-dihydro-15-keto-PGF 23b1;, [PGFM; Zarco
et al., 1988b, Bazer et al., 1991, Bazer et al., 1992, Bazer et al., 1998, Thatcher et al.,
1995 and Arnold et al., 2000]. However, basal concentrations of PGFM are increased
(Zarco et al., 1988b).
Differences in uterine venous PGF23b1; between these studies are probably irrelevant,
since less than 5% of PGF23b1; is transferred locally from the uterine vein to the adjacent
ovarian artery (Hixson and Hansel, 1974). Instead, ovine embryos impart a resistance to
PGF23b1;, since more PGF23b1; given IM is required during early pregnancy to induce
luteolysis than in nonbred ewes (Inskeep and Murdoch, 1980, Silvia and Niswender,
1984 and Silvia and Niswender, 1986). It has been suggested that ovine luteal tissue
catabolizes PGF23b1; during early pregnancy to prevent luteolysis (Silva et al., 2000). The
porcine uterus may secrete PGF23b1; into the uterine lumen rather than the uterine vein
during early pregnancy (Bazer and Thatcher, 1977). However, uterine venous PGF23b1;
increases in pigs on days 112013;13 post-breeding, but decreases through day 16, while
PGF23b1; is pulsatile from days 132013;16 in nonpregnant pigs (Christenson et al., 1994).
Embryos probably stimulate production of a blood-borne antiluteolysin or luteotropin for
delivery by the same local vascular route as PGF23b1;, to prevent luteolysis (Weems et al.,
1995).
Molecules produced by embryos that may have roles in establishing pregnancy directly or
indirectly proteins [INF-3c4;; pregnancy specific protein B (PSPB); a 35-kDa protein];
early pregnancy factor (EPF), or lipids (PAF, PGE 1, PGE2, or PGI2). EPF increases at
fertilization and does not relate to the timing of embryo transfer, the half life of PGI 2 is
too short, and local transfer of PG is slow (Hixson and Hansel, 1974 and Weems et al.,
1995). Proteins are too large for local delivery from the uterine vein to the adjacent
ovarian artery; INF-3c4; is not found in blood; and the 35-kDa protein does not inhibit
PGF23b1;-induced luteolysis (Bazer et al., 1991, Bazer et al., 1998, Roberts et al., 1992
and Wiltbank et al., 1992). Infusions of INF-3c4; into the uterine lumen or via systemic
routes extend ovine and bovine luteal function for only a few days (Davis et al., 1992,
Meyer et al., 1995, Ott et al., 1997, Binelli et al., 2001 and Thatcher et al., 2001).
Porcine interferons do not delay luteolysis in pigs (Lefevre et al., 1998). It was reported
that INF-3c4; decreased endometrial OT and oestrogen receptors, but increased
endometrial progesterone receptors to reduce uterine PGF23b1; secretion during early
pregnancy to prevent luteolysis in the ewe and cow (Spencer et al., 1995a, Spencer et al.,
1995b, Spencer et al., 1996, Spencer and Bazer, 1995, Spencer and Bazer, 2002, Flint,
1995, Mann et al., 1999 and Mann and Lamming, 2001). However, ovine oestradiol-
173b2; does not increase during early pregnancy and inhibition of bovine OT during the
oestrous cycle does not delay luteolysis (Jaroszewki and Kotwica, 1994, Ford, 1995,
Kotwica et al., 1997 and Kotwica et al., 1999). In addition, porcine or equine embryos do
not suppress OT-induced endometrial PGF23b1; secretion (Carnahan et al., 1996 and
Vanderwall et al., 2003). Although PSPB increases in blood by day 21 post-breeding and
increases as pregnancy progresses, infusions of PSPB into the ovine uterine lumen delays
luteolysis only a few days (Ruder et al., 1988 and Sasser et al., 1989; Weems,
unpublished data). These embryonic proteins are immunosuppressive and may suppress
the maternal immune system (Ford, 1984 and Ford, 1995).
Research efforts have focused on PGE1 and PGE2, which are structurally similar to
PGF23b1; and could be delivered locally from the uterine vein to the adjacent ovarian
artery to prevent luteolysis. Both PGE1 and PGE2 are vasodilators and increase luteal
progesterone secretion via cAMP (Marsh, 1971, Fitz et al., 1982, Fitz et al., 1984a, Fitz et
al., 1984b, Niswender et al., 1994, Niswender et al., 2000 and Richards et al., 1994b).
Ovarian and uterine blood flow increases during early pregnancy (Ford, 1984 and Ford,
1995). PGE1 or PGE2 are not luteolytic when large doses are given into the jugular vein
of ewes on days 9 and 10 post-oestrus (Weems et al., 1985b). Receptors for PGE are on
luteal membranes and PGE2 prevents PGF23b1;-induced luteolysis in vitro (Fitz et al.,
1984a, Fitz et al., 1984b, Weipz et al., 1992, Weems et al., 1995, Feng and Almond, 1998
and Feng and Almond, 1999).
A single injection of PGE2 or PGE1 into the luteal-containing ovine ovary prevented a
PGF23b1;-induced luteolysis, increased progesterone for 24 and 48 h, respectively, but
only PGE1 doubled LH bound to receptors and increased unbound LH receptors when the
corpus luteum was threatened with luteolysis (Reynolds et al., 1981 and Weems et al.,
1985e). In cows, PGE1 increased progesterone secretion for 72 h when given IM
(Kimball and Lauderdale, 1975). Chronic infusions of PGE1 or PGE2 into the uterine
lumen of nonbred ewes prevented a natural or premature luteolysis induced by an IUD or
oestradiol-173b2; for up to 48 days. The delay in luteolysis occurred only when PGE 1 or
PGE2 were infused into the uterine lumen adjacent to the luteal-containing ovary (Huie et
al., 1981, Magness et al., 1981, Hoyer et al., 1985, Weems et al., 1985a, Weems et al.,
1985b, Weems et al., 1985c, Weems et al., 1985d and Weems et al., 1995).
When both PGE2 and oestradiol-173b2; are infused into the uterine lumen during the
oestrous cycle of cows, luteolysis is delayed, but not when oestradiol-17 3b2; or PGE2 are
infused alone (Reynolds et al., 1983). Intrauterine infusions of PGE2 into pigs delayed
luteolysis in some studies, but not others (Schneider et al., 1983, Okrasa et al., 1985,
Akinlosotu et al., 1986, Akinlosotu et al., 1988 and Wiesak et al., 1992). Sulfproston, a
synthetic PGE2 analogue, is not luteotropic or antiluteolytic in pigs (Sander et al., 1982).).
However, implants of PGE2 into a pig corpus luteum prevent luteolysis only in that
corpus luteum, while unimplanted luteal cohorts in the same ovary undergo luteolysis
(Ford and Christenson, 1991 and Christenson et al., 1995 In mares, which produce an
embryonic protein gonadotropin, PGE1 does not delay luteolysis (Weems et al., 1985b).
Ovine endometrial PGE1 and PGE2 increase two-fold on day 13 post-breeding (Wilson et
al., 1972b). These increases in PGE2 may be mediated through INF-3c4; downregulation
of 9-keto-PGE2 reductase to inhibit conversion of PGE2 to PGF23b1; during early
pregnancy, but not PGE1, since PGE1 is derived from di-homo-3b3;-linolenic acid, not
AA (Lands, 1979, Beaver and Murdoch, 1992 and Asselin and Fortier, 2000). The ovine
uterine venous PGE2:PGF23b1; ratio on day-8 is 0.1 and increases to 1:1 by day 13 post-
breeding (Vincent and Inskeep, 1986). Progesterone causes premature luteolysis when
given daily, beginning early in the oestrous cycle of sheep. Transfer of day 10 ovine
embryos to a day 6 uterus is detrimental to pregnancy. However, when the uterus is
primed daily with progesterone early in the oestrous cycle, a day 10 embryo transferred
to ewes on day 6 advances uterine PGE2 secretion (Vincent et al., 1986).
In cows, endometrial prostaglandin E-synthase mRNA is expressed from days 18 to 50

post-breeding of pregnancy (Binta et al., 2003). In bovine caruncular crypts, COX-2 and
EP2 receptors are increased and co-expressed during early pregnancy and may be
regulated by INF-3c4; (Parent et al., 2002 and Arosh et al., 2003). Endometrial stroma
secretes PGE2, which may be regulated by EGF and INF-3c4;, while the luminal
epithelium secretes PGF23b1; (Zhang and Davis, 1991, Zhang et al., 1992, Davis and Blair,
1993 and Asselin et al., 1997). Alternatively, LH stimulates PGE2 secretion by ovine
caruncular endometrium up to day 50 of pregnancy, while after day 50 PSPB regulates
placental PGE2 secretion and may also be involved in the asynchronous implantation
(Weems et al., 2003b). Endometrial LH receptors late in the luteal phase may not be for
luteolysis (Shemesh et al., 2002 and Ziecik, 2002), but to regulate PGE2 secretion if an
embryo is present (Weems et al., 2003b).
Other molecules may also be antiluteolytic. Adenosine, a potent immunosuppressant in
rodents, may suppress luteolysis and the maternal immune system during pregnancy
(Priebe and Nelson, 1991). Adenosine prevents PGF23b1;-induced luteolysis in vitro
(Behrman et al., 1986), and in vivo when infused chronically into the uterine lumen or
ovary adjacent to the luteal-containing ovary of sheep (Weems et al., 1989a and Weems
et al., 1992c). Adenosine also enhances the gonadotropic response to hCG, PGE 1, and
PGE2 by bovine or ovine luteal cell suspensions in vitro (Weems et al., 1991 and Weems
et al., 1992a).
Progesterone is required throughout pregnancy (Weems et al., 1995). Fetal growth

stretches the uterus causing it to contract (Nathanielsz, 1988). Progesterone relaxes the
myometrium by decreasing gap junctions to reduce contractions to prevent fetal
expulsion (Nathanielsz, 1988 and Wray, 1993). In sheep, but not cows or pigs, circulating
progesterone increases after the first trimester (Fylling, 1970, Stabenfeldt et al., 1970,
Sarda et al., 1973, Rickets and Flint, 1980, Randel and Erb, 1971 and Hamon and Heap,
1990). In cows, the placenta does not secrete progesterone when the corpus luteum is
functional, but the placenta can secrete progesterone to maintain pregnancy after day 150
if the corpus luteum is regressed (Conley and Ford, 1987 and Fuchs et al., 1996). Ovaries
are not necessary after day 55 in sheep, since the placenta secretes sufficient progesterone
to maintain pregnancy (Casida and Warwick, 1945, Weems et al., 1992d and Weems et
al., 1999b). In ewes, half of the progesterone circulating at day 90 is from the corpus
luteum and half is from the placenta (Weems et al., 1992d and Weems et al., 1995).
Ovine corpora lutea and placentae at day 90 weigh 600 and 700,000 mg, respectively,
indicating that the placenta contains only a few steroidogenic cells (Gross and Williams,
1988, Weems et al., 1992d and Weems et al., 1995). Progesterone increases until day
130, and decreases by day 135 when steroidogenic enzymes decrease (Stabenfeldt et al.,
1970 and Rodgers et al., 1988).
Circulating oestradiol-173b2;, PGE2, and PSPB increase as pregnancy progresses and

increase further in ewes with twin fetuses (Findlay and Cox, 1970, Hallford et al., 1990,
Weems et al., 1993, Weems et al., 1994a, Weems et al., 1994b and Weems et al., 1995).
Ninety per cent of circulating oestradiol-173b2; is secreted by the ovine placenta at day 90
of pregnancy (Weems et al., 1994a). This is due to inhibition of ovarian follicular growth
during pregnancy (Weems et al., 1994a) and the decreased LH availability as pregnancy
progresses (Nalbandov and Casida, 1939, Little et al., 1982 and Nett, 1987). This
decreased LH availability and follicular growth during pregnancy may contribute to the
delay in resumption of postpartum oestrous cycles in sheep and cows (Inskeep, 1995 and
Weems et al., 1995).
In sheep, uterine PGF23b1; secretion declines by day 50 and PGF23b1; in the uterine vein at
day 90 averages 200 pg/mL (Weems et al., 1993 and Weems et al., 2003b). This is due to
an inhibitor of COX produced by the placenta, which is regulated by progesterone
receptors (Rice et al., 1987, Gross et al., 1988 and Weems et al., 2002b). Concentration
of PGE2 in the uterine vein averages 6 ng/mL by day 90 (Weems et al., 1993). In 90 day
intact or ovariectomized (OVX) pregnant ewes, PGF23b1; does not abort ewes, but luteal
tissue regresses without affecting placental progesterone secretion (Weems et al., 1992d
and Weems et al., 1995). Exogenous PGF23b1; increases oestradiol-173b2; in 90 day
pregnant ewes, which is followed by concomitant increases in PGF23b1;, PGE2, and PSPB
64 h later (Weems et al., 1992d, Weems et al., 1994a and Weems et al., 1994b).
However, 90 day pregnant ewes given a single injection of PGF23b1; and oestrogen every
6 h for three days produces an increase in oestradiol-173b2; and PGF23b1; linearly and ewes
start to abort. This does not occur when either hormone is given alone (Bridges et al.,
1999a and Bridges et al., 1999b). This explains why uterine PGF23b1; secretion needs to
be suppressed, since oestrogen secretion increases as pregnancy progresses (Findlay and
Cox, 1970, Weems et al., 1994a and Weems et al., 1995). Trilostane, a 3-3b2;-hydroxy-
steroid dehydrogenase inhibitor, decreased progesterone and increased oestradiol-17 3b2;
and PGF23b1;, while RU-486, a progesterone receptor antagonist, increased PGF 23b1;
secretion to abort 90 day pregnant ewes (Weems et al., 1999a, Weems et al., 1999c and
Weems et al., 2002b).
A model has been proposed whereby oestradiol-173b2; regulates placental PSPB secretion,
PSPB regulates placental PGE2, and, in turn, PGE2 regulates placental progesterone
secretion to maintain pregnancy in sheep (Weems et al., 1995). Aromatase inhibitors in
vivo decrease oestradiol-173b2; first, followed by decreases in PSPB and then PGE2 before
progesterone decreases in 90 day pregnant ewes (Weems et al., 2001). Oestradiol-173b2;
increases PSPB and indomethacin decreases PGE 2 and progesterone secretion in vivo in
90 day pregnant ewes (Bridges et al., 1999a and Bridges et al., 1999b). After OVX of 90
day pregnant ewes, circulating progesterone decreases, but is followed by increases in
oestradiol-173b2;, then PSPB, and then PGE2 before progesterone increases (Weems et al.,
1999b). Placental tissue from 90 day OVX pregnant ewes secrete two-three fold more
progesterone, oestradiol-173b2;, PGE2, and PSPB than intact ewes in vitro, but not
PGF23b1; (Weems et al., 1999b). In vitro, PSPB increases bovine endometrial and 602013;
90 day ovine placental PGE2 secretion, but PGE1 or PGE2 stimulate placental
progesterone secretion only when PG synthesis is inhibited (Del Vecchio et al., 1990,
Weems et al., 2003b and Weems et al., 2004c). It is interesting that recurrent human
abortions are associated with antibodies to phospholipids, the source of AA for PG
synthesis (Lands, 1979, Roth, 1963, Brown, 1990, Gleicher et al., 1992 and Cowchock et
al., 1992). Calcium ions may regulate ruminant placental progesterone secretion
(Shemesh, 1990 and De La Llosa-Hermier et al., 1991). However, calcium channel
agonists did not affect bovine placental progesterone or PG secretion (Weems et al.,
2004a).
Hypophysectomy of pregnant ewes at day 50 does not cause the pregnancy to be lost.
This indicates that regulation of luteal progesterone secretion is not regulated by LH
(Casida and Warwick, 1945 and Denamur et al., 1973). Pituitary LH content, LH in
blood, and LH pulse height and frequency decrease as pregnancy progresses in ruminants
(Nalbandov and Casida, 1939, Little et al., 1982, Schallenberger et al., 1985 and Nett,
1987). Indomethacin in vivo reduces bovine progesterone secretion during the oestrous
cycle and in 90 day pregnant sheep (Milvae and Hansel, 1985 and Bridges et al., 1999a).
Concentrations of PGE2 in the ovarian and uterine vein of sheep are similar at day 90 of
pregnancy (Weems et al., 1993). Ovine luteal tissue of pregnancy, but not the oestrous
cycle, secretes PGE2 in vitro (Kim et al., 2001, Weems et al., 1997 and Weems et al.,
2003a).
Bovine luteal tissue of pregnancy or the oestrous cycle secretes both PGE 2 and PGF23b1;
in vitro, which increases with time (Weems et al., 1998b). During the oestrous cycle, LH
regulates luteal progesterone secretion (Niswender et al., 1994). However, PGE1 or PGE2,
not LH, regulates 90 day ovine or 200 day bovine luteal progesterone secretion during
pregnancy (Weems et al., 1997, Weems et al., 1998a and Kim et al., 2001). Moreover,
indomethacin decreases 90 day ovine luteal progesterone and PGE 2 secretion in vitro,
which can be restored with PGE2, but not LH (Kim et al., 2001). Ovine luteal
progesterone secretion is regulated by LH until day 50 of pregnancy, and PGE 2 regulates
luteal progesterone secretion after day 50 (Weems et al., 2003b). PSPB regulates luteal
and placental PGE2 secretion after day 50 (Weems et al., 1997, Weems et al., 2003a and
Weems et al., 2003b).
NO may play a role in bovine luteal PGE2 secretion. NO donors increase PGE2, but not
PGF23b1; secretion in vitro, indicating that NO is not luteolytic (Weems et al., 2004a).
Chronic infusion of NO donors into the ovine uterine lumen or ovarian vascular pedicle
adjacent to the luteal-containing ovary delays luteolysis (Weems et al., 2004d). NO
donors prevent PGF23b1;-induced luteolysis in vitro (Dong et al., 1999). NO appears to be
involved in maintenance of uterine blood flow, implantation, maintenance of pregnancy,
and NO declines prior to parturition (Yallampalli and Garfield, 1993, Buhimschi et al.,
1995, Buhimschi et al., 2000, Purcell et al., 1997, Sladek et al., 1997, Yallampalli et al.,
1998, Zhang et al., 1999, Zhang et al., 2000, Chwalisz et al., 1999, Chwalisz and
Garfield, 2000, Vedemikov et al., 2000, Yallampalli and Dong, 2000 and Lowe, 2000).
Arginine is the main source of NO and arginine increases in porcine uterine fluids during
gestation, which may be to assist in maintenance of blood flow to the uterus during
pregnancy (Ford, 1984, Ford, 1995, Sladek et al., 1997 and Kwon et al., 2003).
7. Parturition
In sheep, parturition is initiated by the fetal hypothalamic/hypophyseal/adrenal axis.
Destruction of any part of this axis delays parturition (Liggins, 1969, Challis and Lye,
1986 and Thorburn, 1991). Similar relationships exist for regulating parturition in cows
(Wagner et al., 1974 and Bazer and First, 1983). As parturition approaches, fetal
hypothalamic corticotrophin releasing hormone (CRH) secretion increases to increase
fetal hypophyseal adrenocorticotropic hormone (ACTH) secretion and then fetal adrenal
cortisol secretion before maternal progesterone decreases. Infusions of CRH, ACTH, or
cortisol into fetal circulation a few days prior to parturition advances parturition (Challis
and Lye, 1986, Challis et al., 2000 and Thorburn and Rice, 1990). Reduction in
progesterone receptor activity with RU-486 facilitates parturition in cows (Li et al.,
1991). This is followed by increases in oestradiol-17 3b2;, OT, PGF23b1;, and PGE2, leading
to changes in myometrial activity from contractures to contractions (Nathanielsz, 1988,
Thorburn, 1991, Janszen et al., 1993 and Challis et al., 2000).
Oestrogen increases OT and uterine and placental PGF23b1; secretion, while PGF23b1;
increases uterine/placental oestradiol-173b2; secretion to reinforce uterine contractility
(Fuchs et al., 1992 and Fuchs et al., 1997). Myometrial contractions may be increased by
OT without increasing PGF23b1; in cows (Taverne et al., 2001).With increases in
oestrogen and decreases in progesterone, myometrial gap junctions are reestablished to
propagate impulses throughout the myometrium for forceful contractions (Wray, 1993).
Uterine venous and circulating PGE2 increase as pregnancy progresses. It has been
suggested that blood levels of PGE2 reach levels that the lung cannot catabolize so that is
initiated (Thorburn and Rice, 1990 and Thorburn, 1991). However, PGE synthase is not
co-regulated with COX in sheep placentae at term and cortisol increases COX, but not
PGE synthase expression (Martin et al., 2002).
Bovine placental slices treated with PGE2 increased PGE2 secretion even in the presence
of PG synthesis inhibitors, but without changes in 8-epi-PGE 2, which is not dependent on
enzymatic production via COX (Weems et al., 1999d). Bovine placental OT receptors
increase at parturition and are associated with increased PGF 23b1; secretion to facilitate
parturition (Fuchs et al., 1992). Decreases in uterine NOS and NO occur just prior to the
onset of parturition (Purcell et al., 1997). Cervical dilation is associated with increases in
water content, collagen degradation, and collagen content, which may be facilitated by
PGE2 and relaxin (Ellwood et al., 1980, Ledger et al., 1985, Hall and Anthony, 1993,
Platz-Christensen et al., 1997, Kankover and Wiercinski, 1999 and Breeveld-Dwarkasing
et al., 2003). Parturition can be induced with PGF23b1;, but the incidence of a retained
placenta is more than 90% (Weems et al., 1992d and Weems et al., 1995).
8. Postpartum period
Major postpartum events are myometrial contraction for expulsion of lochia and
elimination of uterine and cervical microbial contamination; uterine and cervical
involution; endometrial repair; and resumption of ovarian cyclicity. Times for postpartum
uterine involution in livestock are: beef cow 2013; 302013;35 days; dairy cow 2013;
452013;50 days; ewe 2013; 282013;30 days; sow 2013; 282013;30 days; and mare 2013;
212013;28 days (Senger, 2003). Subclinical or clinical endometritis or mastitis increases
postpartum PG secretion and delays resumption of oestrous cyclicity and reduces
postpartum fertility (Manns et al., 1985, Del Vecchio et al., 1992, Bekana et al., 1996a,
Bekana et al., 1996b, Archbald et al., 1998, Hockett et al., 2000, Seals et al., 2002a, Seals
et al., 2002b, Lewis, 2003 and Sales and Jabbour, 2003) This is exacerbated by induced
parturition where the incidence of retained placenta is nearly 100% (Weems et al., 1999a,
Weems et al., 1999c and Weems et al., 2002b).
Uterine infections are associated with increased uterine endotoxins, PG, and early
pregnancy loss (Mateus et al., 2003, Stewart et al., 2003 and Grigsby et al., 2003).
Intrauterine inoculation of pregnant ewes with Chlamydia psittaci increases PGF23b1; and
decreases PGE2 in uterine venous plasma (Howie, 1989); however, endotoxin does not
affect bovine placental PG secretion in vitro (Weems et al., 2004b). This suggests that
microbes colonize and destroys the placenta resulting in abortion.
Administration of PGF23b1; postpartum shortens the postpartum interval in beef cows (Del
Vecchio et al., 1988, Randel et al., 1988 and Tolleson and Randel, 1988) and may
influence days to first oestrus in dairy cows (Young et al., 1984, Young and Anderson,
1986, McClary et al., 1989 and White and Dobson, 1990), by enhancing lochia
evacuation. On day 32 or days 292013;42 after calving, PGF23b1; treatment decreased
postpartum intervals (Randel et al., 1988 and Tolleson and Randel, 1988).
Uterine manipulation 292013;35 days after calving increases endogenous PGF 23b1;
secretion (Wann and Randel, 1990 and Velez et al., 1991) and decreases the postpartum
interval in beef cows (Tolleson and Randel, 1988, Velez et al., 1991 and Vujosevic et al.,
1988). Earlier postpartum return to oestrus in dairy cows may be due to PGF 23b1;
reduction in pyometra, chronic endometritis, and luteinized cysts during the postpartum
period, which are common in dairy cows (Revah et al., 1989 and Young et al., 1984).
Interaction between the uterus, the hypothalamic2013;pituitary2013;ovarian axis, and

PGF23b1; may be implicated in the postpartum return to oestrus (Tolleson and Randel,
1988). Suggestions that PGF23b1; may be involved in GnRH release were proposed in
1970 (Zor et al., 1970). In 1973, it was reported that PGF23b1; stimulated LH release in
oestrous cycling ewes (Carlson et al., 1973). This does not occur in anestrous or OVX
ewes (Chamley and Christie, 1973). Pituitary LH release can be induced by PG in male
rats (Ratner et al., 1974). Postpartum beef (Randel et al., 1988) and dairy (Peters, 1989)
cows have increased circulating LH following PGF23b1;administration. Exogenous
PGF23b1; facilitates GnRH-stimulated LH release in multiparous and primiparous suckled
beef cows in a dose related manner (Randel et al., 1996). Reduction of the postpartum
period in cows treated with PGF23b1;, or induced to secrete PGF23b1;, may be due to a
combination of mechanisms. Uterine involution or follicular development may be
hastened which may advance ovarian cyclicity. Also, PGF23b1; may act upon the
hypothalamic2013;pituitary axis to stimulate LH secretion and advance ovarian cyclicity
(Randel et al., 1996).
Evidence exists that PGF23b1; is involved in short luteal phases, which often accompany
postpartum return to ovarian cyclicity. Premature uterine secretion of PGF 23b1; causes a
short luteal phase of the first postpartum ovulation in cows (Garverick et al., 1992).
Further results implicate subluteolytic concentrations of PGF 23b1; in embryonic loss after
the first postpartum ovulation through an embryotoxin released by the corpus luteum
(Inskeep, 1995, Buford et al., 1996 and Costine et al., 2001). This embryotoxin is a PG,
presumably delivered locally from the ovary to the adjacent uterine horn. An inhibitor of
PG synthesis and removal of the corpus luteum improves fertility in bred cows with
norgestomet implants (Inskeep, 1995 and Buford et al., 1996).
It is clear that PGF23b1; can have both positive and negative influences on postpartum
reproductive efficiency. When exogenous PGF23b1; or when endogenous PGF23b1; release
is stimulated prior to return of postpartum ovarian cyclicity, the time to return to oestrus
has been reported to be reduced. When PGF23b1; secretion is not coordinated with ovarian
cyclicity, as in the short luteal phase postpartum, the increased PGF 23b1; results in a
shortened oestrus cycle and decreased conception rates when these cows are
bred(Garverick et al., 1992, Inskeep, 1995, Buford et al., 1996 and Costine et al., 2001).
9. Synchronization of oestrus
Oestrus is synchronized in cows with PGF23b1;; progestins; progestins + oestrogens;
progestins + PGF23b1;; PGF23b1; + GnRH; or PGF23b1; + oestrogen + GnRH.
Synchronization of oestrus with PGF23b1; advances luteal regression to synchronize
oestrus for breeding to produce uniform calves or to improve herd genetics by artificial
insemination (AI). The response by bovine corpora lutea to PGF 23b1; on days 12013;4
post-oestrus is reduced and responsiveness increases as the corpus luteum matures. This
led to two injections of PGF23b1; 11 days apart being used to improve detection of oestrus
in more cows. With or without oestrous detection, timed AI around 80 h after the second
injection of PGF23b1; gives acceptable first service conception rates. Progestins given as
ear implants for nine days and oestradiol-cypionate injected intramuscularly to regress
the corpus luteum has been used because oestrus is detected in more cows.
Controlled internal drug-releasing devices (CIDR) to deliver progesterone have been

employed to synchronize cows. Use of progestins and the potential development of a
cystic follicle led to the addition of GnRH or oestrogen to synchronization protocols with
improvement in some studies, but not others (Beal, 1998).
Bovine luteal slices secrete PGE2 and PGF23b1; linearly with time in vitro at a 1:1 ratio
(Weems et al., 1998b and Weems et al., 2002b). Luteal slices from cows synchronized
with a norgestomet ear implant during the induced oestrous cycle cultured in vitro
resulted in 90% and 57% of luteal cultures secreting no PGE 2 at 4 and 8 h of culture,
respectively, but PGF23b1; secretion was not inhibited. Subsequently, cows were
synchronized with PGF23b1;, a CIDR, or cows were not synchronized. Induced corpora
lutea were collected, weighed, and luteal slices were incubated in vitro.
Synchronization with PGF23b1; lowered luteal weights during, but had no effect on luteal
PGE2 or PGF23b1; secretion in vitro per unit mass. A CIDR did not reduce induced luteal
weights or basal progesterone secretion in vitro, but inhibited NO or ET-1-induced, but
not basal PGE2 secretion when compared to cows whose oestrus was not synchronized
(Weems et al., 2004a). Successful synchronization regimens may depend on stage of
follicular development when treatments are initiated. Cows have two or three follicular
waves and three wave cows have longer inter-oestrous intervals (Ginther et al., 1989,
Ginther et al., 2001, Stock and Fortune, 1993, Savio et al., 1993, Mihm et al., 1994,
Ahmad et al., 1997, Vasconcelos et al., 1999 and Bridges et al., 1999c). Premature
ovulation of a dominant follicle on day 10 or 13 post-oestrus decreased the size of the
ovulatory follicle, reduced subsequent luteal function, and compromised conception rates
(Mussard et al., 2003).
To obtain high pregnancy rates in cows, control of follicular growth is important during
oestrous synchronization (Diskin et al., 2002). Further assessment is needed to
understand the effects of oestrous synchronization on subsequent luteal function and
maintenance of pregnancy.
Acknowledgments
The authors thank their former graduate students and collaborators as well as Drs. Roy
Butcher and Roy Fogwell for review of the manuscript. We acknowledge the support of
USDA Hatch Projects 259 (Hawaii, C. Weems), 3128 (Texas, R. Randel) and USDA
CSREES Special Grants Program administered by the Pacific Basin Advisory Group to
C. Weems (90-34135-5343; 95-34135-1776; 98-34135-6783; 2002-34135-12791). The
manuscript is Journal Series No. 4668 of the College of Tropical Agriculture and Human
Resources. We dedicate this review to Drs. Hal Behrman, Roy Butcher, Ralph Erb, Keith
Inskeep, Jim Lauderdale, Ed Pexton, Laird Wilson Jr., and Mr. Nathan Douglas Weems.
References
Ackerman and Murdoch, 1993 R.C. Ackerman and W.J. Murdoch, Prostaglandin F23b1;-
induced apoptosis of ovarian surface epithelial cells, Prostaglandins 45 (1993), pp.
4752013;485. Abstract-EMBASE | Abstract-MEDLINE | $Order Document
Acosta et al., 1998 T.J. Acosta, A. Miyamoto, T. Ozawa, M.P. Wijayagunawardane and
K. Sato, Local release of steroid hormones, prostaglandin E 2, and endothelin-1 from
bovine mature follicles in vitro: effects of luteinizing hormone, endothelin-1, and
cytokines, Biology of Reproduction 59 (1998), pp. 4372013;443. Abstract-EMBASE |
Abstract-Elsevier BIOBASE | Abstract-MEDLINE | $Order Document
Ahmad et al., 1997 N. Ahmad, E. Townsend, R.A. Dailey and E.K. Inskeep, Relationship
of hormonal patterns and fertility to occurrence of two or three waves of ovarian follicles,
before and after breeding, in beef cows and heifers, Animal Reproduction Science 49
(1997), pp. 132013;28. SummaryPlus | Full Text + Links | PDF (1215 K)
Ainsworth et al., 1979 L. Ainsworth, B.K. Tsang, B.R. Downey, R.D. Baker, G.J. Marcus
and D.T. Armstrong, Effects of indomethacin on ovulation and luteal function in gilts,
Biology of Reproduction 21 (1979), pp. 4012013;409.
Ainsworth et al., 1984 L. Ainsworth, B.K. Tsang, G.J. Marcus and B.R. Downey,
Prostaglandin production by dispersed granulosa and theca interna cells from porcine
preovulatory follicles, Biology of Reproduction 31 (1984), pp. 1142013;115.
Ainsworth et al., 1990 L. Ainsworth, B.K. Tsang, B.R. Downey and G.J. Marcus, The
synthesis and actions of steroids and prostaglandins during follicular maturation in the
pig, Journal of Reproduction and Fertility 40 (1990) (Suppl.), pp. 1372013;148.
Akinlosotu et al., 1986 B.A. Akinlosotu, J.R. Diehl and T. Gimenez, Sparing effects of
intrauterine treatment with prostaglandin E2 on luteal function in gilts, Prostaglandins 32
(1986), pp. 2912013;299. Abstract-MEDLINE | $Order Document
Akinlosotu et al., 1988 B.A. Akinlosotu, J.A. Diehl and T. Gimenez, Prostaglandin E 2
counteracts the effect of PGF23b1; in indomethacin-treated gilts, Prostaglandins 35
(1988), pp. 812013;93. Abstract-EMBASE | Abstract-MEDLINE | $Order Document
Alexander et al., 1990 B.M. Alexander, E.A. VanKirk and W.J. Murdoch, Secretion of
platelet activating factor by periovulatory ovine follicles, Life Science 47 (1990), pp.
8652013;868. Abstract | Abstract + References | PDF (231 K)
Algire et al., 1992 J.E. Algire, A. Srikandakumar, L. Guilbault and B.A. Downey,
Preovulatory changes in follicular prostaglandins and their role in ovulation in cattle,
Canadian Journal of Veterinary Research 56 (1992), pp. 672013;79.
Archbald et al., 1998 L.F. Archbald, I.F. Tsai, W.W. Thatcher, T. Tran, K. Wolfsdorf and
C. Risco, Use of plasma concentrations of 13,14-dihydro, 15-keto-PGF 23b1; (PGFM) in
the diagnosis of sub-clinical endometritis and its relationship to fertility in the postpartum
dairy cow, Theriogenology 49 (1998), pp. 14252013;1436. SummaryPlus | Full Text +
Links | PDF (577 K)
Arnold et al., 2000 D.R. Arnold, M. Binelli, J. Vonk, A.P. Alexenko, M. Drost, C.J.
Wilcox and W.W. Thatcher, Intracellular regulation of endometrial PGF 23b1; and PGE2
production in dairy cows during early pregnancy and following treatment with
recombinant interferon-tau, Domestic Animal Reproduction 18 (2000), pp. 1992013;216.
SummaryPlus | Full Text + Links | PDF (312 K)
Arosh et al., 2003 J.A. Arosh, S.K. Banu, M. Parent, P. Chapdelaine and M.A. Fortier,
Dynamic expression and regulation of prostaglandin receptors EP 2, EP3, EP4, and FP, as
well as cyclooxygenases 1 and 2 in caruncles, intercaruncles, and fetal membranes during
bovine pregnancy, Biology of Reproduction 68 (2003) (Suppl. 1, Abstract), pp.
2482013;249.
Asselin et al., 1997 E. Asselin, D. Lacroix and M.A. Fortier, INF-tau increases PGE 2
production and COX-2 gene expression in the bovine endometrium, Molecular Cellular
Endocrinology 132 (1997), pp. 1172013;126. Abstract | PDF (891 K)
Asselin and Fortier, 2000 E. Asselin and M.A. Fortier, Detection and regulation of the
messenger for a putative endometrial 9-keto-prostaglandin E 2 reductase: effect of
oxytocin and interferon-tau, Biology of Reproduction 62 (2000), pp. 1252013;131.
Abstract-EMBASE | Abstract-Elsevier BIOBASE | Abstract-MEDLINE | $Order
Document
Aten and Behrman, 1989 R.F. Aten and H.R. Behrman, A GnRH binding inhibitor
protein from bovine ovaries: purification and identification as histone-H2A, Journal of
Biological Chemistry 264 (1989), pp. 110652013;11073.
Aten et al., 1987 R.F. Aten, J.J. Ireland, C.W. Weems and H.R. Behrman, Presence of
GnRH-like protein hormone in bovine and ovine ovaries, Endocrinology 120 (1987), pp.
Ball and Day, 1982 G.D. Ball and B.N. Day, Local effects of PGF23b1; and embryonic
extracts on luteal function in swine, Journal of Animal Science 54 (1982), pp.
1502013;159.
Banu et al., 2003 S.K. Banu, J.A. Arosh, P. Chapdelaine and M.A. Fortier, Expression
and regulation of prostaglandin transporter in corpus luteum and utero-ovarian plexus
during the oestrous cycle and pregnancy, Biology of Reproduction 68 (2003) (Suppl. 1,
Abstract), p. 175.
Bauer et al., 2001 M. Bauer, I. Reibiger and K. Spanel-Borowski, Leucocyte proliferation

in the bovine corpus luteum, Reproduction 121 (2001), pp. 2972013;305. Abstract-
EMBASE | Abstract-Elsevier BIOBASE | Abstract-MEDLINE | $Order Document |
Full Text via CrossRef
Bazer and Thatcher, 1977 F.W. Bazer and W.W. Thatcher, Theory of maternal
recognition of pregnancy in swine based on estrogen controlled endocrine versus
exocrine secretion of prostaglandin F23b1; by uterine endometrium, Prostaglandins 14
(1977), pp. 3972013;401.
Bazer and First, 1983 F.W. Bazer and N.L. First, Pregnancy and parturition, Journal of
Animal Science 57 (1983) (Suppl. 2), pp. 4252013;460. Abstract-MEDLINE | $Order
Document
Bazer et al., 1991 F.W. Bazer, W.W. Thatcher, P.J. Hansen, M.A. Mirando, T.L. Ott and
T.L. Plante, Physiological mechanisms of pregnancy recognition in ruminants, Journal of
Reproduction and Fertility 43 (1991) (Suppl.), pp. 392013;46.
Bazer et al., 1992 F.W. Bazer, M.A. Mirando, T.L. Ott, J. Harney, D. Dubois, T. Schalue,
C. Pontzer, C. Hostetler, H. Johnson and T. Ogle, Roles of ovine trophoblastic protein-1
and estradiol-173b2;/prolactin in the establishment of pregnancy in sheep and pigs,
Reproduction Fertility and Development 4 (1992), pp. 3352013;340. Abstract-EMBASE |
Abstract-MEDLINE | $Order Document
Bazer et al., 1998 F.W. Bazer, T.L. Ott and T.E. Spencer, Maternal recognition of
pregnancy: comparative aspects, Trophoblast Research 12 (1998), pp. 3752013;386.
Beal, 1998 W.E. Beal, Current estrous synchronisation and artificial insemination
programs for cattle, Journal of Animal Science 76 (1998) (Suppl. 3), pp. 302013;38.
Beaver and Murdoch, 1992 C.J. Beaver and W.J. Murdoch, Ovarian and uterine
prostaglandin E2-9-ketoreductase activity in cyclic and pregnant ewes, Prostaglandins 44
(1992), pp. 342013;45.
Behrman et al., 1986 H.R. Behrman, R.F. Aten, J.L. Luborsky, M.O. Polan, J.G.O. Miller
and L.K. Soodak, Purines, prostaglandins and peptides-nature and cellular mechanisms of
action of local assist and assassin agents in the ovary, Journal of Animal Science 62
(1986) (Suppl. 2), pp. 142013;24. Abstract-MEDLINE | $Order Document
Behrman and Preston, 1989 H.R. Behrman and S. Preston, Luteolytic actions of hydrogen
peroxide in rat ovarian cells, Endocrinology 124 (1989), pp. 2892013;297.
Bekana et al., 1996a M. Bekana, P. Jonsson and H. Kindahl, Intrauterine bacterial

findings and hormonal profiles in post-partum cows with normal puerperium, Acta
Veterinaria Scandanavia 37 (1996), pp. 2512013;263. Abstract-MEDLINE | $Order
Document
Bekana et al., 1996b M. Bekana, K. Odensvik and H. Kindahl, Prostaglandin F 2 alpha

metabolite and progesterone profiles in post-partum cows with retained foetal
membranes, Acta Veterinaria Scandinavia 37 (1996), pp. 1712013;185. Abstract-
MEDLINE | $Order Document
Bergstrom et al., 1968 S. Bergstrom, L.A. Carlson and J.R. Weeks, The prostaglandins: a
family of biologically active lipids, Pharmacological Reviews 20 (1968), pp. 12013;19.
Binelli et al., 2001 M. Binelli, W.W. Thatcher, R. Mattos and P.S. Baruselli,
Antiluteolytic strategies to improve fertility in cattle, Theriogenology 56 (2001), pp.
14512013;1463. Abstract | Abstract + References | PDF (952 K)
Binta et al., 2003 H. Binta, A.R. Fuchs, M. Drost, W. Thatcher, N. Denslow, N. Nichols,
S.-M.T. Chang, E. Johnson and M.J. Fields, Expression of oxytocin receptor,
prostaglandin (PG) H synthase (S)-2, PGE-S, and PGF-S in endometrium during the
estrous cycle and peri-implantation period, Biology of Reproduction 68 (2003) (Suppl. 1,
Abstract), p. 204.
Breeveld-Dwarkasing et al., 2003 V.N.A. Breeveld-Dwarkasing, J.M. Koppele, R.A.

Bank, G.C. van der Weijden, M.A. Taverne and F.M.F. van Dissel-Emiliani, Changes in
water content, collagen breakdown, collagen content and concentration in repeated
biopsies of the cervix of pregnant cows, Biology of Reproduction 69 (2003), pp.
16082013;1614. Abstract-EMBASE | Abstract-Elsevier BIOBASE | Abstract-
MEDLINE | $Order Document | Full Text via CrossRef
Bridges et al., 1999a P.J. Bridges, Y.S. Weems, L.M. Kim, R.G. Sasser, B.R. LeaMaster,
D.L. Vincent and C.W. Weems, Expression of oxytocin receptor, prostaglandin (PG) H
synthase (S)-2, PGE-S, and PGF-S in endometrium during the estrous cycle and peri-
implantation period, Prostaglandins and Other Lipid Mediators 58 (1999), pp.
1132013;124. SummaryPlus | Full Text + Links | PDF (256 K)
Bridges et al., 1999b P.J. Bridges, Y.S. Weems, L.M. Kim, R.G. Sasser, B.R. LeaMaster,
D.L. Vincent and C.W. Weems, Effect of prostaglandin F23b1; (PGF23b1;), indomethacin,
tamoxifen or estradiol-173b2; on prostaglandin E (PGE), PGF23b1; and estradiol-173b2;
secretion in 882013;90 day pregnant sheep, Prostaglandins and Other Lipid Mediators
58 (1999), pp. 1672013;178. SummaryPlus | Full Text + Links | PDF (428 K)
Bridges et al., 1999c P.J. Bridges, P.E. Lewis, W.R. Wagner and E.K. Inskeep, Follicular
growth, estrus and pregnancy after fixed-time insemination in cows treated with
intravaginal progesterone inserts and estradiol benzoate, Theriogenology 52 (1999), pp.
5732013;583. Abstract | PDF (570 K)
Bridges and Fortune, 2003 P.J. Bridges and J.E. Fortune, Progesterone mediates
gonadotrophin-induced secretion of prostaglandins E and F23b1; (PGE, PGF) by follicular
cells from bovine preovulatory follicles, Biology of Reproduction 68 (2003) (Suppl. 1,
Abstract), p. 47.
Brown, 1990 H.L. Brown, Antiphospholipid antibodies and recurrent pregnancy loss,
Clinical Obstetrics and Gynecology 34 (1990), pp. 172013;22.
Brown and Reeves, 1983 J.L. Brown and J.J. Reeves, Absence of specific gonadotropin
releasing hormone receptors in ovine, bovine and porcine ovaries, Biology of
Reproduction 29 (1983), pp. 11792013;1182. Abstract-EMBASE | Abstract-
Buford et al., 1996 W.I. Buford, N. Ahmad, F.N. Schrick, R.L. Butcher, P.E. Lewis and
E.K. Inskeep, Embryotoxicity of a regressing corpus luteum in beef cows supplemented
with progestogen, Biology of Reproduction 54 (1996), pp. 5312013;537. Abstract-
EMBASE | Abstract-MEDLINE | $Order Document
Buhimschi et al., 1995 I.A. Buhimschi, C. Yallampalli, Y. Dong and R.E. Garfield,
Involvement of a nitric oxide-cyclic guanosine monophosphate pathway in control of
human uterine contractility during pregnancy, American Journal of Obstetrics and
Gynecology 172 (1995), pp. 15772013;1584. SummaryPlus | Full Text + Links | PDF
(677 K)
Buhimschi et al., 2000 I.A. Buhimschi, C. Yallampalli, C.S. Buhimschi, G.E. Sadde and
R.E. Garfield, Distinct regulation of nitric oxide and cyclic guanosine monophosphate
production by steroid hormones in the rat uterus, Molecular Human Reproduction 6
| Full Text via CrossRef
Caldwell and Moor, 1970 B. Caldwell and R.M. Moor, Further studies on the role of the
uterus in regulation of corpus luteum function in sheep, Journal of Reproduction and
Fertility 26 (1970), pp. 1332013;144.
Carlson et al., 1973 J.C. Carlson, B. Barcikowski and J.A. McCracken, PGF 23b1; and the
release of LH in sheep, Journal of Reproduction and Fertility 34 (1973), pp.
Carnahan et al., 1996 K.G. Carnahan, B.C. Prince and M.A. Mirando, Exogenous
oxytocin stimulates uterine secretion of prostaglandin F23b1; in cyclic and early pregnant
swine, Biology of Reproduction 55 (1996), pp. 8382013;843. Abstract-Elsevier
BIOBASE | Abstract-MEDLINE | $Order Document
Carvalho et al., 1989 C.B. Carvalho, B.S. Yeik and W.J. Murdoch, Significance of
follicular cyclooxygenase and lipoxygenase pathways of metabolism of arachidonate in
sheep, Prostaglandins 37 (1989), pp. 5532013;558. Abstract-EMBASE | Abstract-
Casida and Warwick, 1945 L.E. Casida and J. Warwick, The necessity of the corpus
luteum for maintenance of pregnancy in the ewe, Journal of Animal Science 4 (1945), pp.
342013;39.
Cavender and Murdoch, 1988 J.L. Cavender and W.J. Murdoch, Morphological studies
on the microcirculatory system of periovulatory ovine follicles, Biology of Reproduction
39 (1988), pp. 9892013;997. Abstract-MEDLINE | $Order Document
Cavicchio et al., 2002 V. Cavicchio, J. Pru, B. Davis, J. Davis, B. Rueda and D.

Townson, Secretion of monocyte chemoattractant protein-1 by endothelial cells of the
bovine corpus luteum: regulation by cytokines but not prostaglandin F 23b1;,
Endocrinology 143 (2002), pp. 35822013;3589. Abstract-EMBASE | Abstract-Elsevier
BIOBASE | Abstract-MEDLINE | $Order Document | Full Text via CrossRef
Challis and Lye, 1986 J. Challis and S. Lye, Parturition, Oxford Reviews Reproductive
Biology 8 (1986), pp. 612013;76.
Challis et al., 2000 J.R.G. Challis, S.G. Matthews, W. Gibbs and S.J. Lye, Endocrine and
paracrine regulation of birth at term and preterm, Endocrine Reviews 21 (2000), pp.
5142013;550. Abstract-EMBASE | Abstract-Elsevier BIOBASE | $Order Document |
Chamley and Christie, 1973 W.A. Chamley and M. Christie, Failure of prostaglandin
F23b1; to affect LH secretion in the ovariectomized ewe, Prostaglandins 3 (1973), pp.
4042013;412.
Cheon et al., 2002 Y.P. Cheon, Q. Li, X. Xu, F.J. Demayo, I.C. Bachi and M.K. Bachi, A
genomic approach to identify novel progesterone regulated pathways in the uterus during
implantation, Molecular Endocrinology 16 (2002), pp. 28532013;2871. Abstract-
Choi et al., 2003 Y. Choi, G.A. Johnson, T.E. Spencer and F.W. Bazer, Pregnancy and
interferon-tau regulate major histocompatibility complex class I and 3b2; 2-microglobulin
expression in the ovine uterus, Biology of Reproduction 68 (2003), pp. 17032013;1710.
Document
Christenson et al., 1994 L.K. Christenson, D.B. Farley, L.H. Anderson and S.P. Ford,
Luteal maintenance during early pregnancy in the pig: role for prostaglandin E 2,
Prostaglandins 47 (1994), pp. 612013;75. Abstract-EMBASE | Abstract-Elsevier
Christenson et al., 1995 L.K. Christenson, D.B. Farley and S.P. Ford, Evaluation of
biochemical and structural changes in individual porcine corpora lutea during
prostaglandin F23b1;-induced luteolysis with an in vivo implant system, Domestic Animal
Endocrinology 12 (1995), pp. 412013;50. SummaryPlus | Full Text + Links | PDF (670
K)
Chwalisz et al., 1999 K. Chwalisz, E. Winterhager, T. Thienel and R.E. Garfield,

Synergistic role of nitric oxide and progesterone during the establishment of pregnancy in
the rat, Human Reproduction 14 (1999), pp. 5422013;552. Abstract-EMBASE | Abstract-
Chwalisz and Garfield, 2000 K. Chwalisz and R.E. Garfield, Role of nitric oxide in
implantation and menstruation, Human Reproduction 3 (2000) (Suppl.), pp. 962013;111.
Abstract-EMBASE | Abstract-MEDLINE | $Order Document
Conley and Ford, 1987 A.J. Conley and S.P. Ford, Effect of prostaglandin F 23b1;-induced
luteolysis in vivo and in vitro progesterone production by individual placentomes in
cows, Journal of Animal Science 65 (1987), pp. 5002013;507. Abstract-MEDLINE |
$Order Document
Cooke and Ahmad, 1994 R.G. Cooke and N. Ahmad, Potential role for lipoxygenase
products of arachidonic acid in prostaglandin F23b1;-stimulated oxytocin release from the
ovine corpus luteum, Journal of Endocrinology 142 (1994), pp. 472013;52. Abstract-
EMBASE | Abstract-Elsevier BIOBASE | Abstract-MEDLINE | $Order Document
Costine et al., 2001 B.A. Costine, B.L. Sayre and E.K. Inskeep, Embryo-toxicity of a
regressing corpus luteum in ewes, Reproduction 122 (2001), pp. 8832013;887. Abstract-
Cowchock et al., 1992 F.S. Cowchock, E.A. Reece, D. Balaban, D.W. Branch and L.
Plouffe, Repeated fetal losses associated with antiphospholipid antibodies: a collaborative
randomized trial comparing prednisone with low-dose heparin treatment, American
Journal of Obstetrics and Gynecology 166 (1992), pp. 13182013;1327.
Davis et al., 1983 D.L. Davis, P.L. Pakrasi and B.N. Day, Prostaglandins in swine
blastocysts, Biology of Reproduction 28 (1983), pp. 11142013;1122.
Davis and Blair, 1993 D. Davis and R. Blair, Uterine secretions and products of primary
cultures of endometrial cell in pigs, Journal of Reproduction and Fertility 48 (1993)
(Suppl.), pp. 1432013;155. Abstract-MEDLINE | $Order Document
Davis and Rueda, 2002 J.S. Davis and B.R. Rueda, The corpus luteum: an ovarian
structure with maternal instincts and suicidal tendencies, Frontiers Bioscience 7 (2002),
pp. 19492013;1979.
Davis et al., 1992 M.A. Davis, T.L. Ott, M.A. Mirando, M.T. Moser and F.W. Bazer,
Effect of recombinant alpha interferons on fertility and interestrous interval in sheep,
Theriogenology 38 (1992), pp. 8672013;875. Abstract | Full Text + Links | PDF (639 K)
De La Llosa-Hermier et al., 1991 M.P. De La Llosa-Hermier, J. Martal, A. Ricour and C.

Hermier, Evidence for modulation of progesterone secretion by calcium and protein
kinase C activators in ovine chorionic cells, Placenta 12 (1991), pp. 5112013;519.
Del Vecchio et al., 1988 R.P. Del Vecchio, R.D. Randel, D.A. Neuendorff and L.A.
Peterson, Effects of alfaprostol, lasalocid and once-daily suckling on postpartum interval
in Brahman and Brahman crossbred cattle, Theriogenology 30 (1988), pp. 7972013;809.
Abstract | Full Text + Links | PDF (660 K)
Del Vecchio et al., 1990 R.P. Del Vecchio, R.G. Sasser and R.D. Randel, Effect of
pregnancy specific protein B on prostaglandins F23b1; and E2 release by day 16 perfused
bovine endometrial tissue, Prostaglandins 40 (1990), pp. 2712013;282. Abstract-
Del Vecchio et al., 1992 R.P. Del Vecchio, D.J. Matsas, T.J. Inzana, D.P. Sponenberg
and G.S. Lewis, Effect of intrauterine bacterial infusions and subsequent endometritis on
prostaglandin F23b1; metabolite concentrations in postpartum beef cows, Journal of
Animal Science 70 (1992), pp. 31582013;3162. Abstract-MEDLINE | $Order Document
Denamur et al., 1973 R. Denamur, G. Kann and R.V. Short, How does the corpus luteum
of the sheep know that there is an embryo in the uterus In: C.G. Pierrepont, Editors,
Endocrinology of Pregnancy and Parturition, Alpha Omega Publishing Company,
Cardiff, Wales, UK (1973), pp. 242013;38.
DeSilva and Reeves, 1985 M. DeSilva and J.J. Reeves, Indomethacin inhibition of
ovulation in the cow, Journal of Reproduction and Fertility 75 (1985), pp. 5472013;553.
Diaz et al., 2000 F.J. Diaz, T.D. Crenshaw and M.C. Wiltbank, Prostaglandin F2 (alpha)
induces distinct physiological responses in porcine corpora lutea after acquisition of
luteolytic capacity, Biology of Reproduction 63 (2000), pp. 15042013;1512. Abstract-
Diskin et al., 2002 M.G. Diskin, E.J. Austin and J.F. Roche, Exogenous hormonal
manipulation of ovarian activity in cattle, Domestic Animal Endocrinology 23 (2002), pp.
Dong et al., 1999 Y.L. Dong, P.R. Gangula, L. Fang and C. Yallampalli, Nitric oxide
reverses prostaglandin-induced inhibition on ovarian progesterone secretion in rats,
Human Reproduction 14 (1999), pp. 272013;32. Abstract-EMBASE | Abstract-
Douglas and Ginther, 1972 R. Douglas and O.J. Ginther, Effect of prostaglandin F23b1; on
length of diestrus in mares, Prostaglandins 2 (1972), pp. 2652013;271.
Downey and Ainsworth, 1980 B.R. Downey and L. Ainsworth, Reversal of indomethacin
blockade on ovulation in gilts by prostaglandins, Prostaglandins 19 (1980), pp.
172013;24.
Downey et al., 1998 B.R. Downey, J.E. Mootoo and S.E. Doyle, A role for lipoxygenase
metabolites of arachidonic acid in porcine ovulation, Animal Reproduction Science 49
(1998), pp. 2692013;279. Abstract | PDF (563 K)
Eggleston et al., 1990 D.L. Eggleston, C. Wilken, E.A. Van Kirk, E.L. Belden and W.J.
Murdoch, Progesterone induces expression of endometrial messenger RNA encoding for
cyclooxygenase (sheep), Prostaglandins 39 (1990), pp. 6752013;683. Abstract-EMBASE
| Abstract-MEDLINE | $Order Document
Ellinwood et al., 1979a W.E. Ellinwood, T.M. Nett and G.D. Niswender, Maintenance of
the corpus luteum of early pregnancy in the ewe: I. Luteotropic properties of embryonic
homogenates, Biology of Reproduction 21 (1979), pp. 2812013;292.
Ellinwood et al., 1979b W.E. Ellinwood, T.M. Nett and G.D. Niswender, Maintenance of
the corpus luteum of early pregnancy in the ewe: II. Prostaglandin secretion by the
endometrium in vitro and in vivo, Biology of Reproduction 21 (1979), pp. 8452013;856.
Ellwood et al., 1980 D.A. Ellwood, M.D. Mitchell, A.B. Anderson and A.C. Turnbull,
Specific changes in the in vitro production of prostanoids by the ovine cervix at
parturition, Prostaglandins 19 (1980), pp. 4792013;488. Abstract-EMBASE | Abstract-
Espey, 1992 L. Espey, A review of factors that could influence membrane potentials of
ovarian follicular cells during mammalian ovulation, Acta Endocrinologica 126 (1992)
(Suppl.), pp. 12013;16.
Evans et al., 1983 G. Evans, M. Dobias, G.J. King and D.T. Armstrong, Production of
prostaglandins by porcine preovulatory follicular tissues and their roles in intrafollicular
function, Biology of Reproduction 28 (1983), pp. 3222013;331.
Farin et al., 1989 C.E. Farin, H.R. Sawyer and G.D. Niswender, Analysis of cell types in
the corpus luteum of the sheep, Journal of Reproduction and Fertility 37 (1989) (Suppl.),
pp. 1812013;192.
Feng and Almond, 1998 S.M. Feng and G.W. Almond, Effects of LH, prostaglandin 8-
bromo-cyclic AMP and forskolin on progesterone secretion by pig luteal cells, Journal of
Reproduction and Fertility 113 (1998), pp. 832013;89. Abstract-EMBASE | Abstract-
Elsevier BIOBASE | Abstract-MEDLINE | $Order Document
Feng and Almond, 1999 S.M. Feng and G.W. Almond, PGE receptor characteristics on
porcine luteal cells during the estrous cycle, Prostaglandins and Other Lipid Mediators
Findlay and Cox, 1970 J.K. Findlay and R.I. Cox, Oestrogens in the plasma of the sheep
fetus, Journal of Endocrinology 46 (1970), pp. 2812013;286.
Fitz et al., 1982 T. Fitz, M. Mayan, H. Sawyer and G. Niswender, Characterization of two
steroidogenic cell types in ovine corpus luteum, Biology of Reproduction 27 (1982), pp.
Fitz et al., 1984a T.A. Fitz, P.B. Hoyer and G.D. Niswender, Interactions of
prostaglandins with subpopulations of ovine luteal cells: I. Stimulatory effects of
prostaglandins E1, E2 and I2, Prostaglandins 28 (1984), pp. 1192013;126. Abstract-
Fitz et al., 1984b T.A. Fitz, E.J. Mock, M.H. Mayan and G.D. Niswender, Interactions of
prostaglandins with subpopulations of ovine luteal cells: II. Inhibitory effects of PGF 23b1;
and protection by PGE2, Prostaglandins 28 (1984), pp. 1272013;138. Abstract-EMBASE
Flint, 1995 A.P.F. Flint, Interferon, the oxytocin receptor and the maternal recognition of
pregnancy in ruminants and non-ruminants: a comparative approach, Reproduction,
Fertility and Development 7 (1995), pp. 3132013;318. Abstract-MEDLINE | $Order
Document
Flores et al., 2003 J.A. Flores, E. Choudhary, B.A. Costine, M.E. Wilson and E.K.
Inskeep, The bovine luteal endothelin system regulated by PGF23b1; during the late luteal
phase, Biology of Reproduction 68 (2003) (Suppl. 1, Abstract), p. 306.
Fogwell et al., 1978 R. Fogwell, C. Weems, G. Lewis, R. Butcher and E.K. Inskeep,
Secretion of steroids and induced luteal regression in beef heifers: effects of PGF 23b1; and
removal of corpora lutea, Journal of Animal Science 46 (1978), pp. 17182013;1723.
Fogwell et al., 1985 R. Fogwell, J. Cowley, J. Wortman, K. Ames and J.J. Ireland, Luteal
function in cows following destruction of ovarian follicles at midcycle, Theriogenology
23 (1985), pp. 3892013;398. Abstract | Full Text + Links | PDF (550 K)
Ford, 1984 S.P. Ford, Maternal recognition of pregnancy in the ewe, cow, and sow:
vascular and immunological aspects, Theriogenol ogy 23 (1984), pp. 1452013;161.
Ford, 1995 S.P. Ford, Control of blood flow to the gravid uterus of domestic livestock
species, Journal of Animal Science 73 (1995), pp. 18522013;1860. Abstract-
Ford et al., 1975 S.P. Ford, C.W. Weems, R. Pitts, J.E. Pexton and E.K. Inskeep, Effects
of estradiol-173b2; and progesterone on prostaglandin F in sheep uteri and uterine venous
plasma, Journal of Animal Science 41 (1975), pp. 14072013;1413. Abstract-
Ford and Christenson, 1991 S.P. Ford and L. Christenson, Direct effects of oestradiol-
173b2; and prostaglandin E2 in protecting pig corpora lutea from a luteolytic dose of
prostaglandin F23b1;, Journal of Reproduction and Fertility 93 (1991), pp. 2032013;209.
Freidman et al., 1995 S. Freidman, M. Gurevich and M. Shemesh, Bovine cyclic

endometrium contains high-affinity luteinizing hormone/human chorionic gonadotropin
binding sites, Biology of Reprod uction 52 (1995), pp. 10202013;1026. Abstract-
Friedman et al., 2000 A. Friedman, S. Weiss, N. Levy and R. Meidan, Role of tumor
necrosis factor alpha and its type I receptor in luteal regression: induction of programmed
cell death in bovine corpus luteum-derived endothelial cells, Biology of Reproduction 63
(2000), pp. 19052013;1912. Abstract-EMBASE | Abstract-Elsevier BIOBASE | Abstract-
Fuchs et al., 1992 A.R. Fuchs, H. Helmer, Behrens, H.C. Liu, L.S. Antonian, S.M. Chang
and M.J. Fields, Oxytocin and bovine parturition: a steep rise in endometrial oxytocin
receptors precedes onset of labor, Biology of Reproduction 47 (1992), pp. 9372013;946.
Fuchs et al., 1996 A.R. Fuchs, M. Rollyson, M.J. Fields, J.N. Enix and R.D. Randel,
Oxytocin induces prostaglandin F23b1; release in pregnant cows: influence of gestational
age and oxytocin receptor concentration, Biology of Reproduction 54 (1996), pp.
6472013;653. Abstract-EMBASE | Abstract-Elsevier BIOBASE | Abstract-MEDLINE |
$Order Document
Fuchs et al., 1997 A.R. Fuchs, Fuchs, M.J. Fields, S.M. Chang, W.W. Thatcher, C.C.
Willard and R.D. Randel, Oxytocin antagonist [1-D(CH2)5, Tyr(ME)2,Thr4,Tyr-
NH2(9)] ornithine vasotocin inhibits oxytocin-induced prostaglandin F 2 alpha release in
late pregnant cows, Biology of Reproduction 57 (1997), pp. 4362013;441. Abstract-
Fylling, 1970 P. Fylling, The effect of pregnancy, ovariectomy, and parturition on plasma
progesterone levels in sheep, Acta Endocrinologica 65 (1970), pp. 2732013;281.
Gadsby et al., 1990 J. Gadsby, A. Balapure, J. Britt and T. Fitz, Prostaglandin F23b1;
receptors on enzyme-dissociated pig luteal cells throughout the estrous cycle,
Endocrinology 126 (1990), pp. 7872013;795. Abstract-EMBASE | Abstract-
Gandolfi et al., 1992 F. Gandolfi, S. Brevini, S. Modina and L. Passoni, Early embryonic
signals: embryo-maternal interactions before implantation, Animal Reproduction Science
28 (1992), pp. 2692013;276. Abstract | Full Text + Links | PDF (467 K)
Garverick et al., 1992 H.A. Garverick, W.G. Zollers and M.F. Smith, Mechanisms
associated with corpus luteum lifespan in animals having normal or subnormal luteal
function, Animal Reproduction Science 28 (1992), pp. 1112013;124. Abstract | Full Text
+ Links | PDF (796 K)
Geisert et al., 1992 R.D. Geisert, E.C. Short and M.T. Zavy, Maternal recognition of
pregnancy, Animal Reproduction Science 28 (1992), pp. 2872013;298. Abstract | Full
Text + Links | PDF (713 K)
Geisert et al., 1994 R. Geisert, R. Short and G. Morgan, Establishment of pregnancy in

domestic farm species In: M. Zavy and R. Geisert, Editors, Embryonic Mortality in
Domestic Species, CRC Press, Boca Raton, FL (1994), pp. 232013;51.
Ginther, 1967 O.J. Ginther, Local utero-ovarian relationships, Journal of Animal Science
26 (1967), pp. 5782013;589.
Ginther, 1974 O.J. Ginther, Internal regulation of physiological processes through local
venoarterial pathways: a review, Journal of Animal Science 39 (1974), pp. 5502013;564.
Ginther, 1981 O.J. Ginther, Local verses systemic utero-ovarian relationships in farm
animals, Acta Veterinaria Scandinavia 77 (1981) (Suppl.), pp. 1032013;117.
Ginther et al., 1989 O.J. Ginther, L. Knopf and J.P. Kastelic, Temporal associations
among ovarian events in cattle during oestrous cycles with two and three follicular
waves, Journal of Reproduction and Fertility 87 (1989), pp. 2232013;230. Abstract-
Ginther et al., 2001 O.J. Ginther, M.A. Berg, D.R. Bergfelt, F.X. Donadeu and K. Kot,
Follicle selection in monovular species, Biology of Reproduction 65 (2001), pp.
6372013;647.
Girsh et al., 1995 E. Girsh, Y. Greber and R. Meidan, Luteotrophic and luteolytic
interactions between bovine small and large luteal-like cells and endothelial cells,
Biology of Reproduction 52 (1995), pp. 9542013;962. Abstract-EMBASE | Abstract-
Gleeson et al., 1974 A.R. Gleeson, G.D. Thorburn and R.J. Cox, Prostaglandin F
concentrations in the utero-ovarian venous plasma of the sow during the late luteal phase
of the oestrous cycle, Prostaglandins 5 (1974), pp. 5212013;529. Abstract-MEDLINE |
$Order Document
Gleicher et al., 1992 N. Gleicher, L. Harlow and M. Zilberstein, Regulatory effect of

autophospholipid antibodies on signal transduction: a possible model for autoantibody-
induced reproductive failure, American Journal of Obstetrics and Gynecology 167
(1992), pp. 6372013;647.
Goding et al., 1972 J. Goding, D. Baird, I. Cumming and J.A. McCracken, Functional
assessment of autotransplanted endocrine glands, Acta Endocrinologica 158 (1972)
Gottsch et al., 2000 M.L. Gottsch, E.A. Van Kirk and W.J. Murdoch, Tumour necrosis
factor alpha up-regulates matrix metalloproteinase-2 activity in periovulatory follicles:
metamorphic and endocrine implications, Reproduction, Fertility and Development 12
Gray et al., 2001 C.A. Gray, K.M. Taylor, W.S. Ramsey, J.R. Hill, F.W. Bazer, F.F.
Bartol and T.E. Spencer, Endometrial glands are required for preimplantation conceptus
elongation and survival, Biology of Reproduction 64 (2001), pp. 16082013;1613.
Document
Grazul-Bilska et al., 1997 A.T. Grazul-Bilska, L.P. Reynolds and D.A. Redmer, Gap
junctions in the ovary, Biology of Reproduction 57 (1997), pp. 9472013;957. Abstract-
Grigsby et al., 2003 P.L. Grigsby, J.J. Hirst, J.P. Scheerlinck, D.J. Phillips and G. Jenkin,
Fetal responses to maternal and intra-amniotic lipopolysaccharide administration in
sheep, Biology of Reproduction 68 (2003), pp. 16952013;1702. Abstract-EMBASE |
Gross and Williams, 1988 T. Gross and W. Williams, Bovine placental prostaglandin
synthesis: cell synthesis as modulated by the binucleate cell, Biology of Reprod uction 38
(1988), pp. 10272013;1038.
Gross et al., 1988 T.S. Gross, W.W. Thatcher, P. Hansen, J. Johnson and S. Helmer,
Presence of an endometrial inhibitor of prostaglandin synthesis during early pregnancy in
the cow, Prostaglandins 35 (1988), pp. 3592013;367.
Grusenmeyer and Pate, 1992 D. Grusenmeyer and J. Pate, Localization of prostaglandin

F2 alpha inhibition of lipoprotein use by bovine luteal cells, Journal of Reproduction and
Fertility 94 (1992), pp. 3112013;318. Abstract-EMBASE | Abstract-MEDLINE | $Order
Document
Gurevich et al., 1993 M. Gurevich, E. Harel-Markowitz, S. Marcus, L.S. Shore and M.

Shemesh, Prostaglandin production by the oocyte cumulus complex around the time of
fertilization and the effect of prostaglandin E on the development of the early bovine
embryo, Reproduction, Fertility, and Development 5 (1993), pp. 1832013;281.
Guthrie and Bolt, 1983 H.D. Guthrie and D.J. Bolt, Changes in plasma estrogen,
luteinizing hormone and 13,14-dihydro-15-keto-prostaglandin F 23b1; during blockade of
luteolysis in pigs after human chorionic gonadotropin treatment, Journal of Animal
Science 52 (1983), pp. 9932013;1000. Abstract-MEDLINE | $Order Document
Hall and Anthony, 1993 J.A. Hall and R.V. Anthony, Influence of ovarian steroids on
relaxin-induced distensibility and compositional changes in the porcine cervix, Biology of
Reproduction 48 (1993), pp. 13482013;1357.
Hallford et al., 1990 D. Hallford, T. Ross, B. Oetting, J. Sachse, C. Heird and R.G.
Sasser, Estimation of fetal numbers in ewes using ultrasound or serum progesterone or
pregnancy specific protein B concentrations, Journal of Animal Science 68 (1990)
(Suppl. 1, Abstract), p. 181.
Halterman and Murdoch, 1986 S.D. Halterman and W.J. Murdoch, Ovarian function in
ewes treated with antihistamines, Endocrinology 119 (1986), pp. 24172013;2421.
Hamon and Heap, 1990 M.H. Hamon and R.B. Heap, Progesterone and estrogen
concentrations in plasma of Barbary sheep compared with those of domestic sheep and
goats during pregnancy, Journal of Reproduction and Fertility 90 (1990), pp.
2072013;213.
Han et al., 1999 S.W. Han, Z.M. Lei and Ch.V. Rao, Treatment of human endometrial
stromal cells with chorionic gonadotropin promotes their morphological and functional
differentiation into deciduas, Molecular Cellular Endocrinology 25 (1999), pp. 72013;16.
Hansel et al., 1991 W. Hansel, H.W. Alila, J.P. Dowd and R.A. Milvae, Differential
origin and control mechanisms in small and large bovine luteal cells, Journal of
Reproduction and Fertility 43 (1991) (Suppl.), pp. 772013;89. Abstract-MEDLINE |
$Order Document
Hansel and Blair, 1996 W. Hansel and R. Blair, Role of lipoxygenase products of
arachidonic acid on bovine corpus luteum function, Reproduction Domestic Animals 31
(1996), pp. 4272013;429.
Hehnke et al., 1994 K.E. Hehnke, L.H. Christensen, S.P. Ford and M. Taylor,
Macrophage infiltration into the porcine corpus luteum during prostaglandin F 23b1;-
induced luteolysis, Biology of Reproduction 50 (1994), pp. 102013;15. Abstract-
Hinckley and Milvae, 2001 S.T. Hinckley and R.A. Milvae, Endothelin-1 mediates
prostaglandin F23b1;-induced luteal regression in the ewe, Biology of Reproduction 64
Hixson and Hansel, 1974 J.R. Hixson and W. Hansel, Evidence for preferential transfer
of prostaglandin F23b1; to the ovarian artery following intrauterine administration in cattle,
Hockett et al., 2000 M. Hockett, F.M. Hopkins, M.J. Lewis, A.M. Saxton, H.H. Dowlen,
S.P. Oliver and F.N. Schrick, Endocrine profiles of dairy cows following experimentally
induced clinical mastitis during early lactation, Animal Reproduction Science 15 (2000),
pp. 2412013;251. SummaryPlus | Full Text + Links | PDF (194 K)
Howie, 1989 A. Howie, The effect of chlamydial infection on the initiation of premature
labor. Serial measurements of intrauterine prostaglandin E 2 in amniotic fluid and
uteroovarian vein, using catheterized sheep experimentally infected with an ovine
abortion strain of Chlamydia psittaci, Prostaglandins, Leukotrienes, and Essential Fatty
Acids 37 (1989), pp. 2032013;214.
Hoyer et al., 1985 G.L. Hoyer, R.R. Magness, J.M. Huie, G. Whysong and C.W. Weems,
Effects of prostaglandin E1 (PGE1) on estradiol-173b2;-induced luteolysis in the
nonpregnant ewe, Prostaglandins, Leukotrienes, and Medicine 17 (1985), pp.
Hu et al., 2001 J. Hu, T. Ludwig, U. Salli, F. Stormshak and M. Mirando, Autocrine and
paracrine action of oxytocin in pig endometrium, Biology of Reproduction 64 (2001), pp.
Huie et al., 1981 J.M. Huie, R.R. Magness, L.P. Reynolds, G.L. Hoyer, T. Huecksteadt,
M. Colcord, B. Stalcup, G.L. Whysong and C.W. Weems, Effect of chronic ipsilateral or
contralateral intrauterine infusion of prostaglandin E 1 (PGE1) on luteal function of
unilaterally ovariectomized ewes, Prostaglandins 21 (1981), pp. 9452013;955. Abstract-
Hunter and Poyser, 1985 R.H.F. Hunter and W.L. Poyser, Ovarian follicular fluid
concentrations of prostaglandins E2, F23b1; and PGI2 during the pre-ovulatory period in
pigs, Reproduction, Nutrition, and Development 25 (1985), pp. 9092013;917. Abstract-
Inskeep and Murdoch, 1980 E.K. Inskeep and W.J. Murdoch, Relation of ovarian
function to uterine and ovarian secretion of prostaglandins during the estrous cycle and
early pregnancy in the ewe and cow, International Review of Physiology 22 (1980), pp.
3252013;356. Abstract-MEDLINE | $Order Document
Inskeep, 1995 E.K. Inskeep, Factors that affect fertility during estrous cycles with short
or normal luteal phases in postpartum cows, Journal of Reproduction and Fertility 49
(1995) (Suppl.), pp. 4932013;503. Abstract-MEDLINE | $Order Document
Janszen et al., 1993 B.P. Janszen, M.M. Bevers, M.M. Ravenshorst, G.C. van der
Weijden, S.J. Dieleman and M.A. Taverne, Relationship between prostaglandin-induced
luteolysis and temporary inhibition of myometrial activity in late pregnant cows with ear
implants containing progestagen, Journal of Reproduction and Fertility 97 (1993), pp.
Jaroszewki and Kotwica, 1994 J.J. Jaroszewki and J. Kotwica, Reduction of ovarian
oxytocin content from early luteal phase does not affect the corpus luteum secretory
function in cattle, Reproduction, Nutrition, and Development 34 (1994), pp.
1752013;182.
Jaroszewki and Hansel, 2000 J. Jaroszewki and W. Hansel, Intraluteal administration of a

nitric oxide synthase blocker stimulates progesterone, oxytocin secretion and prolongs
the life span of the bovine corpus luteum, Proceedings Society Experimental Biology
Medicine 224 (2000), pp. 502013;55.
Jo et al., 2002 M. Jo, C.M. Komar and J.E. Fortune, Gonadotropin surge induces two
separate increases in messenger RNA for progesterone receptor in bovine preovulatory
follicles, Biology of Reproduction 67 (2002), pp. 19812013;1988. Abstract-EMBASE |
Abstract-Elsevier BIOBASE | Abstract-MEDLINE | $Order Document | Full Text via
CrossRef
Johnson et al., 1999a M.L. Johnson, J. Murdoch, E.A. Van Kirk, J.E. Kaltenbach and
W.J. Murdoch, Tumor necrosis factor alpha regulates collagenolytic activity in
preovulatory ovine follicles: relationship to cytokine secretion by the oocyte-cumulus
complex, Biology of Reproduction 61 (1999), pp. 15812013;1585. Abstract-EMBASE |
Johnson et al., 1999b G.A. Johnson, T.E. Spencer, T.R. Hansen, K.J. Austin, R.C.
Burghardt and F.W. Bazer, Expression of the interferon tau inducible ubiquitin cross-
reactive protein in the ovine uterus, Biology of Reproduction 61 (1999), pp. 3122013;318.
Document
Johnson et al., 2001 G.A. Johnson, F.W. Bazer, L.A. Jaeger, H. Ka, J.E. Garlow, C.
Pfarrer, T.E. Spencer and R.C. Burghardt, Muc-1, integrin, and osteopontin expression
during the implantation cascade in sheep, Biology of Reproduction 65 (2001), pp.
$Order Document
Kankover and Wiercinski, 1999 M. Kankover and J. Wiercinski, Prostaglandin E 2-9-keto

reductase from bovine term placenta, Prostaglandins, Leukotrienes and Essential Fatty
Acids 61 (1999), pp. 292013;32.
Kennedy and Ross, 1993 T.G. Kennedy and H.E. Ross, Effects of prostaglandin E 2 on
rate of decidualization in rats, Prostaglandins 46 (1993), pp. 2432013;250. Abstract-
Kim et al., 2001 L. Kim, Y.S. Weems, P.J. Bridges, B.R. LeaMaster, D.L. Vincent and
C.W. Weems, Effects of indomethacin, luteinizing hormone (LH), prostaglandin E 2
(PGE2), trilostane, mifepristone, ethamoxytriphetol (MER-25), or pregnancy on secretion
of prostaglandin E (PGE), prostaglandin F23b1; (PGF23b1;) and progesterone by ovine
corpora lutea of pregnancy or the estrous cycle, Prostaglandins and Other Lipid
Mediators 63 (2001), pp. 1892013;203. SummaryPlus | Full Text + Links | PDF (665 K)
Kimball and Lauderdale, 1975 F. Kimball and J. Lauderdale, Prostaglandins E 1 and F23b1;
specific binding sites in bovine corpora lutea: comparison with luteolytic effects,
Prostaglandins 10 (1975), pp. 3132013;319.
Kobayashi et al., 2001 S. Kobayashi, B. Berisha, W.M. Amselgruber, D. Schams and A.

Miyamoto, Production and localization of angiotensin II in the bovine early corpus
luteum: a possible interaction with luteal angiogenic factors and prostaglandin F 2 alpha,
Journal of Endocrinology 170 (2001), pp. 3692013;380. Abstract-EMBASE | Abstract-
Elsevier BIOBASE | Abstract-MEDLINE | $Order Document | Full Text via CrossRef
Kotwica et al., 1997 J. Kotwica, D.J. Skarzynski, M. Bogacki, P. Melin and B. Starostka,
The use of an oxytocin antagonist to study the function of ovarian oxytocin during
luteolysis in cattle, Theriogenology 48 (1997), pp. 12872013;1299. Abstract | PDF (601
K)
Kotwica et al., 1999 G. Kotwica, A. Franczak, S. Okrasa and J. Kotwica, Effect of an

oxytocin antagonist on prostaglandin F23b1; secretion and the course of luteolysis in sows,
Acta Veterinaria Hungary. 47 (1999), pp. 612013;75.
Kwon et al., 2003 H. Kwon, T.E. Spencer, F.W. Bazer and G. Wu, Developmental
changes in amino acids in ovine fetal fluids, Biology of Reproduction 68 (2003), pp.
La Croix and Kann, 1982 M.C. La Croix and G. Kann, Comparative studies of
prostaglandins F23b1; and E2 in late cyclic and early pregnant sheep: In vitro synthesis by
endometrium and conceptus: Effects of in vivo indomethacin treatment on establishment
of pregnancy, Prostaglandins 23 (1982), pp. 5072013;526. Abstract-EMBASE |
Lands, 1979 W.E.M. Lands, The biosynthesis and metabolism of prostaglandins, Annual
Review of Physiology 41 (1979), pp. 6332013;654.
Ledger et al., 1985 W.L. Ledger, M.A. Webster, A.B.M. Anderson and A.C. Turnbull,
Effect of inhibition of prostaglandin synthesis on cervical softening and uterine activity
during ovine parturition resulting from progesterone withdrawal induced by epostane,
Journal of Endocrinology 105 (1985), pp. 2222013;231.
Lefevre et al., 1998 F. Lefevre, Martinat-Botte, A. Locatelli, P. de Niu, M. Terqui and C.

La Bonnardiere, Intrauterine infusion of high doses of pig trophoblast interferons has no
antiluteolytic effect in cyclic gilts, Biology of Reproduction 58 (1998), pp.
Lewis et al., 1978 G.S. Lewis, J.E. Pexton, R.L. Fogwell and E.K. Inskeep,
Concentrations of prostaglandins E2 and F23b1; and their relationship to luteal function in
early pregnant ewes, Journal of Animal Science 47 (1978), pp. 13142013;1322.
Lewis et al., 1982 G.S. Lewis, W.W. Thatcher, F.W. Bazer and J.J. Curl, Metabolism of
arachidonic acid in vitro by bovine blastocysts and endometrium, Biology of
Reproduction 26 (1982), pp. 4312013;439. Abstract-EMBASE | Abstract-MEDLINE |
$Order Document
Lewis, 2003 G.S. Lewis, Role of ovarian progesterone and potential role of prostaglandin
F23b1; and prostaglandin E2 in modulating the uterine response to infectious bacteria in
postpartum ewes, Journal of Animal Science 81 (2003), pp. 2852013;293. Abstract-
Li et al., 1991 Y. Li, P. Perezgrovas, O.S. Gazal, C. Schwabe and L.L. Anderson,
Antiprogesterone, RU-486, facilitates parturition in cattle, Endocrinology 129 (1991), pp.
7652013;772.
Liggins, 1969 G.C. Liggins, The foetal role in the initiation of parturition in the ewe In:
G.E.W. Wolstenholme and M. O2019;Connor, Editors, Ciba Foundation Symposium,
Churchill Publishers, London, England (1969), pp. 2182013;234.
Lim et al., 2002 H. Lim, H. Song, B. Paria, J. Reese, S. Das and S. Dey, Molecules in
blastocyst implantation: uterine and embryonic perspectives, Vitamins and Hormones 64
Little et al., 1982 D. Little, C. Rahe, J. Fleeger and P.G. Harms, Episodic release of LH
during gestation in the cow, Journal of Reproduction and Fertility 66 (1982), pp.
Liu et al., 1997 J. Liu, P.D. Carriere, M. Dore and J. Sirois, Prostaglandin G/H synthase-2
is expressed in bovine preovulatory follicles after the endogenous surge of luteinizing
hormone, Biology of Reproduction 57 (1997), pp. 15242013;1531. Abstract-EMBASE |
Lowe, 2000 D.T. Lowe, Nitric oxide dysfunction in the pathophysiology of pre-
eclampsia, Nitric Oxide 4 (2000), pp. 4412013;458. Abstract | Abstract + References |
PDF (134 K)
Lukaszewska and Hansel, 1980 J. Lukaszewska and W. Hansel, Corpus luteum

maintenance during early pregnancy in the cow, Journal of Reproduction and Fertility 59
Magness et al., 1981 R.R. Magness, J.M. Huie, G.L. Hoyer, T. Huecksteadt, L. Reynolds,
L. Seperich and C.W. Weems, Effects of chronic ipsilateral or contralateral intrauterine
infusion of prostaglandin E2 (PGE2) on luteal function of unilaterally ovariectomized
ewes, Prostaglandins and Medicine 6 (1981), pp. 3892013;401. Abstract-EMBASE |
Mann et al., 1998 G.E. Mann, G.E. Lamming and J.H. Payne, Role of early luteal phase
progesterone in control of the timing of the luteolytic signals in cows, Journal of
Mann et al., 1999 G.E. Mann, G.E. Lamming, R.S. Robinson and D.C. Wathes, The
regulation of interferon-tau production and uterine hormone receptors during early
pregnancy, Journal of Reproduction and Fertility 54 (1999) (Suppl.), pp. 3172013;328.
Mann and Lamming, 2001 G.E. Mann and G.E. Lamming, Relationship between
maternal endocrine environment, early embryo development and inhibition of the
luteolytic mechanism in cows, Reproduction 121 (2001), pp. 1752013;180. Abstract-
Manns et al., 1985 J.G. Manns, J.R. Nkuuhe and F. Bristol, Prostaglandin concentrations
in uterine fluid of cows with pyometra, Canadian Journal of Comparative Medicine 49
Marcus, 1981 G.J. Marcus, Prostaglandins formation by the sheep embryo and
endometrium as an indication of maternal recognition of pregnancy, Biology of
$Order Document
Marsh, 1971 J. Marsh, The effect of prostaglandins on the adenyl cyclase of the bovine
corpus luteum, Annals New York Academy of Science 180 (1971), pp. 4162013;425.
Martin et al., 2002 R.L. Martin, W.L. Whittle, A.C. Holloway, S. Gyomoreg, W. Gibb,
S.J. Lye and J.R.G. Challis, Ontogeny and regulation of ovine placental prostaglandin E 2
synthase, Biology of Reproduction 67 (2002), pp. 8682013;873. Abstract-EMBASE |
CrossRef
Mateus et al., 2003 L. Mateus, L. Lopes da Costa, P. Diniz and A.J. Ziecik, Relationship
between endotoxin and prostaglandin (PGE2 and PGFM) concentrations and ovarian
function in dairy cows with puerperal endometritis, Animal Reproduction Science 76
McClary et al., 1989 D.G. McClary, M.R. Putman, J.C. Wright and J.L. Sartin Jr., Effect
of early postpartum treatment with prostaglandin F23b1; on subsequent fertility in the dairy
cow, Theriogenology 31 (1989), pp. 5652013;570. Abstract | Full Text + Links | PDF
(380 K)
Medan et al., 2003 P. Medan, P.J. Bridges, C.M. Komar, A.G. Beristan, R.
Rajamahendran, J.E. Fortune and C.D. MacCalman, Expression of messenger ribonucleic
acid for ADAMTS subtype changes in the periovulatory follicle after the gonadotropin
surge and during luteal development and regression in cattle, Biology of Reproduction 69
(2003), pp. 15062013;1514.
Meidan et al., 1999 R. Meidan, R.A. Milvae, S. Weiss, N. Levy and A. Friedman,
Intraovarian regulation of luteolysis, Journal of Reproduction and Fertility 54 (1999)
Meyer et al., 1995 M.D. Meyer, P.J. Hansen, W.W. Thatcher, M. Drost, L. Badinga,
R.M. Roberts, J. Li, T.L. Ott and F.W. Bazer, Extension of corpus luteum lifespan and
reduction of uterine secretion of prostaglandin F23b1; of cows in response to recombinant
interferon-tau, Journal of Dairy Science 78 (1995), pp. 19212013;1931. Abstract-
Mihm et al., 1994 M.A. Mihm, L. Baguisi, M.P. Boland and J.F. Roche, Association
between the duration of dominance of the ovulatory follicle and pregnancy rate in beef
heifers, Journal of Reproduction and Fertility 102 (1994), pp. 1232013;130. Abstract-
Milvae and Hansel, 1985 R.A. Milvae and W. Hansel, Inhibition of bovine luteal function
by indomethacin, Journal of Animal Science 60 (1985), pp. 5282013;531. Abstract-
Mirando et al., 1993 M. Mirando, W. Becker and S. Whiteaker, Relationship among

endometrial oxytocin receptors, oxytocin-stimulated phosphoinositide hydrolysis and
prostaglandin F2 alpha secretion in vitro, and plasma concentrations of ovarian steroids
before and during corpus luteum regression in cyclic heifers, Biology of Reproduction 48
(1993), p. 874. Abstract-EMBASE | Abstract-MEDLINE | $Order Document
Mishra et al., 2003 S. Mishra, Z.M. Lei and Ch.V. Rao, A novel role of luteinizing
hormone in the embryo development in cocultures, Biology of Reproduction 68 (2003),
pp. 14552013;1462. Abstract-EMBASE | Abstract-Elsevier BIOBASE | Abstract-
Miyamoto et al., 2002 Y. Miyamoto, R. Sakumoto, Y. Sakabe, M. Miyake, A. Okano and

K. Okuda, Tumour necrosis factor-alpha receptors are present in the corpus luteum
throughout the oestrus cycle and during the early gestation period in pigs, Reproduction
Domestic Animals 37 (2002), pp. 1052013;110. Abstract-Elsevier BIOBASE | Abstract-
Moor and Rowson, 1964 R.M. Moor and L.E.A. Rowson, Influence of the embryo and
uterus on luteal function in the sheep, Nature 10 (1964), pp. 5222013;531.
Moor, 1968 R.M. Moor, Effect of embryo removal on corpus luteum function, Journal of
Animal Science 27 (1968) (Suppl. 2), pp. 972013;118. Abstract-MEDLINE | $Order
Document
Moor et al., 1970 R.M. Moor, M.F. Hay, R.V. Short and L.E.A. Rowson, The corpus
luteum of the sheep: effect of uterine removal during luteal regression, Journal of
Reproduction and Fertility 21 (1970), pp. 3192013;324.
Morgan et al., 1993 G.L. Morgan, R.D. Geisert, J.P. McCann, F.W. Bazer, T.L. Ott, M.A.
Mirando and M. Stewart, Failure of luteolysis and extension of the interoestrus interval in
sheep treated with the progesterone antagonist mifepristone (RU-486), Journal of
Murdoch, 1985W.J. Murdoch, Follicular determinants of ovulation in the ewe, Domestic

Animal Endocrinology 2 (1985), pp. 1052013;113.
Murdoch, 1986 W.J. Murdoch, Accumulation of thromboxane B 2 within periovulatory

ovine follicles: relationship to adhesion of platelets to endothelium, Prostaglandins 32
Murdoch, 1988a W.J. Murdoch, Distribution of cellular associations within the
granulosal compartment of periovulatory ovine follicles: relationship to maturation of the
oocyte and regulation by prostaglandins, Cell Tissue Research 252 (1988), pp.
Murdoch, 1988b W.J. Murdoch, Effect of a steroidal (prednisolone) and nonsteroidal

(indomethacin) anti-inflammatory agent on ovulation and follicular accumulation of
prostaglandin F23b1; in sheep, Prostaglandins 37 (1988), pp. 3312013;334.
Murdoch, 1990 W.J. Murdoch, Localization and hormonal regulation of ovarian

production of histamine in the sheep, Life Science 46 (1990), pp. 19612013;1965.
Abstract | Abstract + References | PDF (241 K)
Murdoch, 1992 W.J. Murdoch, Comparative morphometry and steroidogenic function of

antral ovine follicles destined for ovulation or atresia, Domestic Animal Endocrinology 9
(1992), pp. 2192013;224. Abstract | Abstract + References | PDF (529 K)
Murdoch, 1994a W.J. Murdoch, Ovarian surface epithelium during ovulatory and
anovulatory ovine estrous cycles, Anatomical Record 240 (1994), pp. 3222013;326.
Document
Murdoch, 1994b W.J. Murdoch, Immunoregulation of mammalian fertility, Life Science

55 (1994), pp. 18712013;1876.
Murdoch, 1995a W.J. Murdoch, Endothelial cell death in preovulatory follicles: possible
implication in the biomechanics of rupture, Journal of Reproduction and Fertility 105
Murdoch, 1995b W.J. Murdoch, Programmed cell death in preovulatory ovine follicles,
Murdoch, 1996a W.J. Murdoch, Microtubular dynamics in granulosa cells of

periovulatory follicles and granulose-derived (large) lutein cells of sheep: relationships to
the steroidogenic folliculo-shift and functional luteolysis, Biology of Reproduction 54
(1996), pp. 11352013;1140. Abstract-EMBASE | Abstract-MEDLINE | $Order
Document
Murdoch, 1996b W.J. Murdoch, Differential effects of indomethacin on the sheep ovary:
prostaglandin biosynthesis, intracellular calcium, apoptosis, and ovulation,
Prostaglandins 52 (1996), pp. 4972013;506. Abstract-EMBASE | Abstract-MEDLINE |
$Order Document
Murdoch, 1996c W.J. Murdoch, Ovarian surface epithelium, ovulation and
carcinogenesis, Biological Reviews of Cambridge Philosophical Society 71 (1996), pp.
$Order Document
Murdoch, 1997 W.J. Murdoch, Role of tumor necrosis factor-alpha in the ovulatory
mechanism of ewes, Journal of Animal Science 75 (1997), pp. 16012013;1605. Abstract-
Murdoch, 1998a W.J. Murdoch, Regulation of collagenolysis and cell death by plasmin
within the formative stigma of preovulatory follicles, Journal of Reproduction and
Fertility 113 (1998), pp. 3312013;338.
Murdoch, 1998b W.J. Murdoch, Perturbation of sheep ovarian surface epithelial cells by
ovulation: evidence for roles of progesterone and poly (ADP-ribose) polymerase in the
restoration of DNA integrity, Journal of Endocrinology 156 (1998), pp. 5032013;508.
Abstract-EMBASE | Abstract-MEDLINE | $Order Document | Full Text via CrossRef
Murdoch, 1999 W.J. Murdoch, Plasma-tumour necrosis factor interaction in the ovulatory
process, Journal of Reproduction and Fertility 54 (1999) (Suppl.), pp. 3532013;358.
Murdoch, 2000 W.J. Murdoch, Proteolytic and cellular death mechanisms in ovulatory
ovarian rupture, Biological Signals and Receptors 9 (2000), pp. 1022013;114. Abstract-
Murdoch and Dunn, 1982a W.J. Murdoch and T.G. Dunn, Alterations in follicular steroid
hormones during the preovulatory period in the ewe, Biology of Reprod uction 27 (1982),
pp. 3002013;307. Abstract-EMBASE | Abstract-MEDLINE | $Order Document
Murdoch and Dunn, 1982b W.J. Murdoch and T.G. Dunn, Luteal function after ovulation
blockade by intrafollicular injection of indomethacin in the ewe, Journal of Reproduction
and Fertility 69 (1982), pp. 6712013;675.
Murdoch and Myers, 1983 W.J. Murdoch and D.A. Myers, Effect of treatment of estrous
ewes with indomethacin on the distribution of ovarian blood to the periovulatory follicle,
Murdoch and Cavender, 1987 W.J. Murdoch and J.L. Cavender, Mechanisms of
ovulation, Advances Contraceptive Delivery Systems 3 (1987), pp. 3532013;367.
Murdoch and Ferris, 1988 W.J. Murdoch and M.L. Ferris, Prostaglandin E2-9-keto-
reductase activity of preovulatory ovine follicles, Journal of Animal Science 66 (1988),
pp. 29242013;2929. Abstract-MEDLINE | $Order Document
Murdoch and Cavender, 1989 W.J. Murdoch and J.L. Cavender, Effect of indomethacin
on the vascular architecture of preovulatory ovine follicles: possible implication in the
luteinized unruptured follicle syndrome, Fertility and Sterility 51 (1989), pp.
Murdoch and McCormick, 1989 W.J. Murdoch and R.J. McCormick, Production of low
weight molecular chemoattractants for leucocytes by periovulatory ovine follicles,
Murdoch and McCormick, 1991 W.J. Murdoch and R.J. McCormick, Dose-dependent
effects of indomethacin on ovulation in sheep: relationship to follicular prostaglandin
production, steroidogenesis, collagenolysis and leukocyte chemotaxis, Biology of
$Order Document
Murdoch and Steadman, 1991 W.J. Murdoch and L.E. Steadman, Investigations
concerning the relationship of ovarian eosinophilia to ovulation and luteal function in the
sheep, American Journal of Reproductive Immunology 25 (1991), pp. 812013;87.
Murdoch and McCormick, 1992a W.J. Murdoch and R.J. McCormick, Enhanced
degradation of collagen within apical vs. basal wall of ovulatory follicles, American
Journal of Physiology 263 (1992), pp. E2212013;225.
Murdoch and McCormick, 1992b W.J. Murdoch and R.J. McCormick, Sequence analysis
of chemoattractant peptides secreted by periovulatory ovine follicles, Biochemistry
Biophysics Research Communication 184 (1992), pp. 8482013;852. Abstract | Full Text
Murdoch and McCormick, 1993 W.J. Murdoch and R.J. McCormick, Mechanisms and
physiological implications of leukocyte chemoattraction into periovulatory ovine
follicles, Journal of Reproduction and Fertility 97 (1993), pp. 3752013;380. Abstract-
Murdoch and Lund, 1999 W.J. Murdoch and S.A. Lund, Prostaglandin-independent
anovulatory mechanism of indomethacin action: Inhibition of tumor necrosis factor
alpha-induced sheep ovarian cell apoptosis, Biology of Reproduction 61 (1999), pp.
Murdoch and McDonnel, 2002 W.J. Murdoch and A.C. McDonnel, Roles of the ovarian
surface epithelium in ovulation and carcinogenesis, Reproduction 123 (2002), pp.
$Order Document | Full Text via CrossRef
Murdoch and Van Kirk, 2002 W.J. Murdoch and E.A. Van Kirk, Steroid hormonal
regulation of proliferative, p53 tumor suppressor, and apoptotic responses of sheep
ovarian surface epithelial cells, Molecular and Cellular Endocrinology 186 (2002), pp.
Murdoch et al., 1981 W.J. Murdoch, R.A. Dailey and E.K. Inskeep, Preovulatory changes
in prostaglandins E2 and F23b1; in ovine follicles, Journal of Animal Science 53 (1981),
pp. 1922013;205. Abstract-MEDLINE | $Order Document
Murdoch et al., 1983 W.J. Murdoch, K.J. Nix and T.G. Dunn, Dynamics of ovarian blood
supply to periovulatory follicles of the ewe, Biology of Reproduction 28 (1983), pp.
Murdoch et al., 1986 W.J. Murdoch, T.A. Peterson, E.A. Van Kirk, D.L. Vincent and
E.K. Inskeep, Interactive roles of progesterone, prostaglandins and collagenase in the
ovulatory mechanisms of the ewe, Biology of Reproduction 35 (1986), pp.
Murdoch et al., 1991 W.J. Murdoch, R.G. Slaughter and T.H. Ji, In situ hybridization
analysis of ovarian endoperoxide synthase mRNA throughout the periovulatory period of
the ewe, Domestic Animal Endocrinology 8 (1991), pp. 4552013;457. Abstract | Abstract
+ References | PDF (217 K)
Murdoch et al., 1993 W.J. Murdoch, T.R. Hansen and L.A. McPherson, Role of
eicosanoids in vertebrate ovulation, Prostaglandins 46 (1993), pp. 852013;115. Abstract-
Murdoch et al., 1999a W.J. Murdoch, C. Wilken and D.A. Young, Sequence of apoptosis
and inflammatory necrosis within the formative ovulatory site of sheep follicles, Journal
of Reproduction and Fertility 117 (1999), pp. 3252013;329. Abstract-EMBASE |
Murdoch et al., 1999b J. Murdoch, E.A. Van Kirk and W.J. Murdoch, Hormonal control
of urokinase plasminogen activator secretion by sheep ovarian surface epithelial cells,
Murdoch et al., 1999c W.J. Murdoch, E.A. Van Kirk and J. Murdoch, Plasmin cleaves
tumor necrosis factor alpha exodomain from sheep follicular endothelium: implication in
the ovulatory process, Biology of Reproduction 60 (1999), pp. 11662013;1171. Abstract-
Mussard et al., 2003 M. Mussard, C.R. Burke, C.L. Gasser, E.J. Behlke, K.A.
Colliflower, D.F. Grum and M.L. Day, Ovulatory response, luteal function, and fertility
in cattle induced to ovulate dominant follicles of early or late maturity, Biology of
Reproduction 68 (2003) (Suppl. 1, Abstract), p. 332.
Nalbandov and Casida, 1939 A. Nalbandov and L.E. Casida, Gonadotropic activity of
pituitaries from cows in different stages of pregnancy, Proceedings American Society of
Animal Production (1939), pp. 3402013;341.
Narumiya, 1995 S. Narumiya, Structures, properties and distributions of prostanoid

receptors In: B. Samuelsson, P.W. Ramwell, R Paoletti, G. Folco, E. Granstrom and S.
Nicosia, Editors, Advances in Prostaglandin, Thromboxane, and Leukotriene Research
vol. 23 (1995), pp. 172013;22. Abstract-MEDLINE | $Order Document
Nathanielsz, 1988 P.W. Nathanielsz, The regulation of the switch from myometrial
contractures to contractions in late pregnancy: studies in the sheep and monkey In: P.D.
Gluckman, B.W. Johnson and P.W. Nathanielsz, Editors, Advances in Fetal Physiology
vol. 8, Perinatology Press, Ithaca, NY, USA (1988), pp. 4092013;428.
Nett, 1987 T.M. Nett, Function of the hypothalamic2013;hypophyseal axis during

gestation in ewes and cows, Journal of Reproduction and Fertility 34 (1987) (Suppl.), pp.
Nicholson et al., 1999 W.C. Nicholson, Z. Ge, D.M. Plotner, C.E. Farin and J.E. Gadsby,
Insulin-like growth factor (IGF)-1, IGF-1 receptor, and IGF binding protein-3 messenger
ribonucleic acids and protein in corpora lutea from prostaglandin F (2 alpha)-treated gilts,
Niswender et al., 1970 G.D. Niswender, P.J. Dzuik, C.C. Kaltenbach and H.W. Norton,
Local effects of embryos and the uterus on corpora lutea in gilts, Journal of Animal
Science 30 (1970), pp. 2252013;234.
Niswender et al., 1994 G.D. Niswender, J.L. Juengel, W.J. McGuire, C.J. Belfiore and
M.C. Wiltbank, Luteal function: the estrous cycle and early pregnancy, Biology of
$Order Document
Niswender et al., 2000 G.D. Niswender, J.L. Juengel, P.J. Silva, M.K. Rollyson and E.W.
McIntush, Mechanisms controlling the function and life span of the corpus luteum,
Physiological Reviews 80 (2000), pp. 12013;29. Abstract-EMBASE | Abstract-Elsevier
Novaro et al., 1996 V. Novaro, V. Rettori, E.T. Gonzalez, A. Jawerbaum, A. Faletti, G.

Canteros and M.A. Gimeno, Interaction between uterine PGE and PGF23b1; production
and the nitridergic system during embryonic implantation in the rat, Prostaglandins 51
Okrasa et al., 1985 S.O. Okrasa, J.E. Tilton and R.M. Weigl, Utero-ovarian venous
concentrations of prostaglandin E2 (PGE2) and prostaglandin F23b1; (PGF23b1;) following
PGE2 intrauterine infusions, Prostaglandins 30 (1985), pp. 8512013;859.
Okuda et al., 2002 K. Okuda, A.Y. Miyamoto and D.J. Skarzynski, Regulation of
endometrial prostaglandin F23b1; synthesis during luteolysis and early pregnancy in cattle,
Domestic Animal Endocrinology 23 (2002), pp. 2552013;264. SummaryPlus | Full Text
Okuda et al., 2003 K. Okuda, M.M. Bah, K.M. Deptula, I. Woclawek-Potocka, A.

Karzekwa, W. Pilawski and D.J. Skarzynski, Tumor necrosis factor-3b1; inversely
regulates prostaglandin F23b1; and E2 production in heifers, Biology of Reproduction 68
(2003) (Suppl. 1, Abstract), p. 351.
O2019;Shea and McCoy, 1988 J.D. O2019;Shea and K. McCoy, Weight, composition,
mitosis, cell death, content of progesterone and DNA in the corpus luteum of pregnancy
in the ewe, Journal of Reproduction and Fertility 83 (1988), pp. 1072013;119.
Ott et al., 1997 T.L. Ott, J.G. Fleming, T.E. Spencer, M.M. Joyce, P. Chen, C.N. Green,
D. Zhu, T.H. Welsh Jr., P.G. Harms and F.W. Bazer, Effects of exogenous recombinant
ovine interferon-tau, circulating progesterone, cortisol, luteinizing hormone, and antiviral
activity; interestrous interval; rectal temperature; and uterine response to oxytocin in
cyclic ewes, Biology of Reproduction 57 (1997), pp. 6212013;629. Abstract-EMBASE |
Ott et al., 1998 T. Ott, J. Yin, A. Wiley, H. Kim, B. Gerami-Naini, T. Spencer, F. Bartol,
R. Burghardt and F. Bazer, Effects of the estrous cycle and early pregnancy on uterine
expression of Mx protein in sheep (Ovis aries), Biology of Reproduction 59 (1998), pp.
Ottobre et al., 1984 J.S. Ottobre, D.L. Vincent, W.J. Silvia and E.K. Inskeep, Aspects of
regulation of uterine secretion of prostaglandins during the oestrous cycle and early
pregnancy, Animal Reproduction Science 7 (1984), pp. 752013;100. Abstract | Full Text
Parent et al., 2002 J. Parent, P. Chapdelaine, J. Sirois and M.A. Fortier, Expression of
microsomal prostaglandin E synthase in bovine endometrium: coexpression with
cyclooxygenase type 2 and regulation by interferon-tau, Endocrinology 143 (2002), pp.
Pate, 1995 J.L. Pate, Involvement of immune cells in regulation of ovarian function,
Journal of Reproduction and Fertility 49 (1995) (Suppl.), pp. 3652013;377. Abstract-
Penny, 2000 L. Penny, Monocyte chemoattractant protein 1 in luteolysis, Reviews

Reproduction 5 (2000), pp. 632013;66. Abstract-EMBASE | Abstract-Elsevier BIOBASE
| Abstract-MEDLINE | $Order Document | Full Text via CrossRef
Penny et al., 1999 L. Penny, D. Armstrong, T. Bramley, R. Webb, R. Collins and E.
Watson, Immune cells and cytokine production in the bovine corpus luteum throughout
the oestrous cycle and after induced luteolysis, Journal of Reproduction and Fertility 115
Peters, 1989 A.R. Peters, Effect of prostaglandin F23b1; on hormone concentrations in

dairy cows after parturition, Veterinary Record 124 (1989), pp. 3712013;372.
Petroff et al., 2001 J.G. Petroff, B.K. Petroff and J.L. Pate, Mechanisms of cytokine-
induced death of cultured bovine luteal cells, Reproduction 121 (2001), pp. 7532013;760.
Pexton et al., 1975a J.E. Pexton, S.P. Ford, L. Wilson, R.L. Butcher and E.K. Inskeep,
Prostaglandins F in uterine tissue and venous plasma of ewes with intrauterine devices,
Journal of Animal Science 41 (1975), pp. 1442013;155.
Pexton et al., 1975b J.E. Pexton, C.W. Weems and E.K. Inskeep, Prostaglandins F in
uterine venous plasma, ovarian arterial and venous plasma and in ovarian and luteal
tissue of pregnant and nonpregnant ewes, Journal of Animal Science 41 (1975), pp.
1542013;159.
Pexton et al., 1975c J.E. Pexton, C.W. Weems and E.K. Inskeep, Prostaglandins F in
uterine and ovarian venous plasma from nonpregnant and pregnant ewes collected by
cannulation, Prostaglandins 9 (1975), pp. 5012013;509.
Piper et al., 1970 P.J. Piper, J.R. Vane and J.H. Wyllie, Inactivation of prostaglandins by
the lungs, Nature 225 (1970), pp. 6042013;660.
Pitzel et al., 1993 L. Pitzel, H. Jarry and W. Wuttke, Different steroidogenic response of
young and aged porcine small and large luteal cells to prostaglandin F 2 alpha, oxytocin
and estradiol, Experimental Clinical Endocrinology 101 (1993), pp. 2552013;261.
Document
Pitzel et al., 2000 L. Pitzel, S. Ludemann and W. Wuttke, Secretion and gene expression
of metalloproteinases and gene expression of their inhibitors in porcine corpora lutea at
different stages of the luteal phase, Biology of Reproduction 62 (2000), pp.
Platz-Christensen et al., 1997 J.J. Platz-Christensen, P. Pernevi and H. Bokstrom,

Prostaglandin E and F23b1; concentration in the cervical mucus and mechanism of cervical
ripening, Prostaglandins 53 (1997), pp. 2532013;261. SummaryPlus | Full Text + Links
| PDF (1134 K)
Priebe and Nelson, 1991 T.S. Priebe and J.A. Nelson, Adenosine and immune system
function In: J.W. Phillips, Editors, Adenosine and Adenine Nucleotides as Regulators of
Cellular Function, CRC Press, Boca Raton, FL, USA (1991), pp. 1412013;156.
Purcell et al., 1997 T.L. Purcell, I.A. Buhimschi, R. Given, K.E. Chwalisz and R.E.
Garfield, Inducible nitric oxide synthase is present in the rat placenta at the
fetal2013;maternal interface and decreases prior to labour, Molecular Human
Reproduction 3 (1997), pp. 4852013;491. Abstract-MEDLINE | $Order Document | Full
Text via CrossRef
Ramwell et al., 1977 P.W. Ramwell, E.M.K. Leovey and A.C. Sintetos, Regulation of the
arachidonic acid cascade, Biology of Reproduction. 16 (1977), pp. 702013;88.
Randel and Erb, 1971 R.D. Randel and R.E. Erb, Reproductive steroids in the bovine:
changes and interrelationships from 0 to 260 days of pregnancy, Journal of Animal
Science 33 (1971), pp. 1152013;123. Abstract-MEDLINE | $Order Document
Randel et al., 1988 R.D. Randel, R.P. Del Vechhio, D.A. Neuendorff and L.A. Peterson,
Effect of alfaprostol on postpartum reproduction in Brahman cows and heifers,
Theriogenology 29 (1988), pp. 6572013;670.
Randel et al., 1996 R.D. Randel, M.A. Lammoglia, A.W. Lewis, D.A. Neuendorff and
M.J. Guthrie, Exogenous PGF23b1; enhanced GnRH-induced LH release in postpartum
cows, Theriogenology 45 (1996), pp. 6432013;654. SummaryPlus | Full Text + Links |
PDF (703 K)
Ratner et al., 1974 A. Ratner, M.C. Wilson, L. Srivastava and G.R. Peake, Stimulatory
effect of PGE1 on rat anterior pituitary cyclic AMP and luteinizing hormone release,
Rawlings and Hyland, 1985 N.C. Rawlings and J.H. Hyland, Prostaglandin F and E
levels in the conceptus, uterus and plasma during early pregnancy in the ewe,
$Order Document
Revah et al., 1989 I. Revah, R. Lomas, L. Zarzo and C. Galina, Evaluacion del
tratamiento rutinario con prostaglandin F23b1; en el dia 302013;40 post-parto sobre la
actividad ovarica y la eficiencia reproductive de vacas Holstein, Veterinaria Mexico 20
(1989), pp. 1352013;143.
Reynolds et al., 1981 L.P. Reynolds, J. Stigler, G.L. Hoyer, R.R. Magness, J.M. Huie, T.
Huecksteadt, H.R. Behrman and C.W. Weems, Effects of PGE1 or PGE2 on PGF23b1;-
induced luteolysis in nonbred ewes, Prostaglandins 21 (1981), pp. 9572013;972.
Reynolds et al., 1983 L.P. Reynolds, D. Robertson and S.P. Ford, Effects of intrauterine
infusion of oestradiol-173b2; and protaglandin E2 on luteal function in nonpregnant
heifers, Journal of Reproduction and Fertility 69 (1983), pp. 7032013;709. Abstract-
Rice et al., 1987 G.E. Rice, M.H. Wong and G.D. Thornburn, Ovine myometrial cytosol
inhibits phospholipase A2 activity in vitro, Prostaglandins 34 (1987), pp. 5632013;571.
Richards and Almond, 1994a R.G. Richards and G.W. Almond, Tumour necrosis factor-
alpha differentially alters progesterone and prostaglandin F 2 alpha production by porcine
luteal cells, Journal of Endocrinology 143 (1994), pp. 752013;83. Abstract-EMBASE |
Richards et al., 1994b R.G. Richards, J.E. Gadsby and G.W. Almond, Differential effects
of LH and PGE2 on progesterone secretion by small and large porcine luteal cells,
Journal of Reproduction and Fertility 102 (1994), pp. 272013;34. Abstract-EMBASE |
Rickets and Flint, 1980 A.P. Rickets and A.P.F. Flint, Onset of synthesis of progesterone
by ovine placenta, Journal of Endocrinology 86 (1980), pp. 3372013;345.
Roberts et al., 1992 R.M. Roberts, J.C. Cross and D.W. Leaman, Interferons as hormones
of pregnancy, Endocrine Reviews 13 (1992), pp. 4322013;452. Abstract-EMBASE |
Abstract-MEDLINE | $Order Document | Full Text via CrossRef
Rodgers et al., 1988 R.J. Rodgers, Rodgers, M.R. Waterman, R.E. Simpson and R.R.
Magness, Immunoblot analysis of cholesterol side-chain cleavage cytochrome P-450 and
adrenodoxin in corpora lutea of cyclic and late pregnant sheep, Journal of Reproduction
and Fertility 83 (1988), pp. 8432013;850. Abstract-EMBASE | Abstract-MEDLINE |
$Order Document
Rodgers et al., 1995 R.J. Rodgers, C.A. Vella, F.M. Young, X.C. Tian and J.E. Fortune,
Concentrations of cytochrome P450 cholesterol side-chain cleavage enzyme and 3 beta-
hydroxysteroid dehydrogenase during prostaglandin F2 alpha-induced luteal regression in
cattle, Reproduction Fertility and Development 7 (1995), pp. 12132013;1216. Abstract-
Roth, 1963 D.C. Roth, The frequency of spontaneous abortions, International Journal of
Fertility 8 (1963), pp. 4312013;442.
Rowson and Moor, 1967 L.E.A. Rowson and R.M. Moor, The influence of embryonic
tissue homogenate into the uterus on the life span of the corpus luteum in the sheep,
Journal of Reproduction and Fertility 13 (1967), pp. 5112013;519.
Ruder et al., 1988 C.A. Ruder, J. Stellflug, J. Dahman, K. Ivani and R.G. Sasser,
Detection of pregnancy in sheep by radioimmunoassay of sera for pregnancy specific
protein B, Theriogenology 29 (1988), pp. 9052013;912. Abstract | Full Text + Links |
PDF (449 K)
Sakamoto et al., 1995 K. Sakamoto, K. Miwa, T. Ezashi, E. Okuda-Ashitaka, T. Houtani,

T. Sugimoto, S. Ito and O. Hayaishi, Expression of mRNA encoding the prostaglandin F 2
alpha receptor in bovine corpora lutea throughout the oestrous cycle and pregnancy,
Sakumoto et al., 2000 R. Sakumoto, B. Berisha, N. Kawate, D. Schams and K. Okuda,

Tumor necrosis factor-3b1; and its receptor in bovine corpus luteum throughout the
estrous cycle, Biology of Reproduction 62 (2000), pp. 1922013;199. Abstract-EMBASE |
Salamonsen et al., 2001 L.A. Salamonsen, N. Guiying, E. Dimitriadis, L. Robb and J.K.
Findlay, Genes involved in implantation, Reproduction, Fertility, and Development 13
Sales and Jabbour, 2003 K. Sales and H. Jabbour, Cyclooxygenase and prostaglandins in
reproductive tract physiology and pathology, Prostaglandins and Other Lipid Mediators
Samuelsson et al., 1971 Samuelsson, B., Granstrom, E., Green, K., Hamberg, M., 1971.
Metabolism of prostaglandins. IN: Prostaglandins, EDS. Ramwell, P.W., Shaw, J.E.
Annals New York Academy of Science 180, 1382013;163.
Sander et al., 1982 H. Sander, T. Brunn, W. Elger and F. Ellendorff, Prostaglandin E

(Sulproston) is neither luteolytic nor luteotropic in the pig, Prostaglandins 24 (1982), pp.
6912013;698.
Sarda et al., 1973 F.R. Sarda, H. Robertson and J. Smeaton, Sequential changes in plasma
progesterone levels in the ewe during the estrous cycle and during pregnancy in intact
and ovariectomized sheep, Canadian Journal of Animal Science 53 (1973), pp.
252013;35.
Sasser et al., 1989 R. Sasser, J. Crock and C. Ruder-Montegomery, Characteristics of

pregnancy specific protein B in cattle, Journal of Reproduction and Fertility 37 (1989)
(Suppl), pp. 2892013;304.
Savio et al., 1993 J.D. Savio, W.W. Thatcher, L. Badinga, R.L. de la Sota and D.
Wolfensen, Regulation of dominant follicle turnover during the estrous cycle in cows,
Saxena et al., 2000 D. Saxena, S. Purohit, G. Kumer and M. Laloraya, Increased
inducible nitric oxide synthase in the uterus and embryo at implantation, Nitric Oxide 4
Scenna et al., 2002 F.N. Scenna, M.E. Hockett, H.E. Blackmon and F.N. Schrick,
Development of in vivo-derived embryos cultured with prostaglandin F 23b1;,
Theriogenology 57 (2002) (Abstract), p. 512.
Schallenberger et al., 1985 E. Schallenberger, J. Ramp and D.L. Walters, Gonadotropins

and ovarian steroids in cattle: II. Pulsatile changes of concentrations in the jugular vein
throughout pregnancy, Acta Endocrinologica 108 (1985), pp. 3222013;330. Abstract-
Schneider, 1976 W.P. Schneider, The chemistry of prostaglandins In: S.M.M. Karim,
Editors, Prostaglandins: Chemical and Biochemical Aspects, University Park Press,
Baltimore, MD, USA (1976), pp. 12013;23.
Schneider et al., 1983 T.M. Schneider, J.E. Tilton, S. Okrasa, J. Mah, R. Weigl and G.L.
Williams, The effect of intrauterine infusions of prostaglandin E 2 on luteal function in
nonpregnant gilts, Theriogenology 20 (1983), pp. 5092013;517.
Schrick and Inskeep, 1993 F.N. Schrick and E.K. Inskeep, Determination of early
pregnancy in ewes using transrectal ultrasonography, Theriogenology 40 (1993), pp.
2952013;306. Abstract | Full Text + Links | PDF (1101 K)
Seals et al., 2002a R.C. Seals, I. Matamoros and G.S. Lewis, Relationship between
postpartum changes in 13,14-dihydro-15-keto-PGF23b1; concentrations in Holstein cows
and their susceptibility to endometritis, Journal of Animal Science 80 (2002), pp.
Seals et al., 2002b R.C. Seals, M.C. Wulster-Radcliffe and G.S. Lewis, Modulation of the
uterine response to infectious bacteria in postpartum ewes, American Journal of
Reproductive Immunology 47 (2002), pp. 572013;63. Abstract-EMBASE | Abstract-
Elsevier BIOBASE | Abstract-MEDLINE | $Order Document | Full Text via CrossRef
Self et al., 2003 J. Self, G. Johnson, F. Bazer and T. Spencer, Developmental changes in
placental nitric oxide synthesis in pigs, Biology of Reproduction 68 (2003) (Suppl. 1,
Abstract), p. 153.
Senger, 2003 P.L. Senger, Pathways to Pregnancy and Parturition, Current Concepts Inc.,
Washington State University, Pullman, WA, USA (2003) pp. 3282013;345.
Sharma and Fitzpatrick, 1974 S.C. Sharma and R.J. Fitzpatrick, Effect of oestradiol-173b2;
and oxytocin treatment on prostaglandin F23b1; release in anestrous ewes, Prostaglandins
6 (1974), pp. 972013;103.
Shemesh, 1990 M. Shemesh, Production and regulation of progesterone in bovine corpus
luteum and placenta in mid and late gestation: a personal review, Reproduction, Fertility,
and Development 2 (1990), pp. 1292013;144.
Shemesh et al., 2002 M. Shemesh, D. Mizrachi, L.S. Shore, J. Reed and S.-M.T. Chang,
Expression of functional luteinizing hormone (LH) receptor and its messenger
ribonucleic acid in bovine endometrium: LH augmentation of cAMP and inositol
phosphate in vitro and human chorionic gonadotrophin (hCG) augmentation of peripheral
prostaglandin in vivo, Reproductive Biology 1 (2002), pp. 132013;32.
Silvia and Niswender, 1984 W.J. Silvia and G.D. Niswender, Maintenance of the corpus
luteum of early pregnancy in the ewe: III. Differences between pregnant and nonpregnant
ewes in luteal responsiveness to prostaglandin F23b1;, Journal of Animal Science 59
(1984), pp. 7562013;764.
Silvia and Niswender, 1986 W.J. Silvia and G.D. Niswender, Maintenance of the corpus
luteum of early pregnancy in the ewe: IV. Changes in luteal sensitivity to prostaglandin
F23b1; throughout early pregnancy, Journal of Animal Science 63 (1986), pp.
Silvia et al., 1984a W.J. Silvia, T.A. Fitz, M. Mayan and G.D. Niswender, Cellular and
molecular mechanisms involved in luteolysis and maternal recognition of pregnancy in
the ewe, Animal Reproduction Science 7 (1984), pp. 572013;74. Abstract | Full Text +
Links | PDF (913 K)
Silvia et al., 1984b W.J. Silvia, J.S. Ottobre and E.K. Inskeep, Concentrations of
prostaglandins E2, F23b1;, and 6-keto-prostaglandin F13b1; in the utero-ovarian venous
plasma of nonpregnant and early pregnant ewes, Biology of Reproduction 30 (1984), pp.
Silvia et al., 1991 W.J. Silvia, G.S. Lewis, J.A. McCracken, W.W. Thatcher and L.
Wilson jr., Hormonal regulation of uterine secretion of prostaglandin F 23b1; during
luteolysis in ruminants, Biology of Reproduction 45 (1991), pp. 6552013;663. Abstract-
Silva et al., 2000 P. Silva, J. Juengel, M. Rollyson and G. Niswender, Prostaglandin

metabolism in the ovine corpus luteum: catabolism of prostaglandin F 23b1; (PGF23b1;)
coincides with resistance of the corpus luteum to PGF23b1;, Biology of Reproduction 63
Sirois, 1994 J. Sirois, Induction of prostaglandin endoperoxide synthase-2 by human

chorionic gonadotropin in bovine preovulatory follicles in vivo, Endocrinology 135
Skarzynski and Okuda, 2000 D.J. Skarzynski and K. Okuda, Different actions of
noradrenaline and nitric oxide on the output of prostaglandins and progesterone in
cultured bovine luteal cells, Prostaglandins and Other Lipid Mediators 60 (2000), pp.
Skarzynski et al., 2003 D. Skarzynski, J. Jaroszewki, M. Bah, K. Deptula, B.

Barszczewska, B. Gawronska and W. Hansel, Administration of nitric oxide synthase
inhibitor counteracts prostaglandin F23b1;-induced luteolysis in cattle, Biology of
Reproduction 68 (2003), pp. 16742013;1681. Abstract-EMBASE | Abstract-Elsevier
Sladek et al., 1997 S.M. Sladek, R.R. Magness and K.P. Conrad, Nitric oxide and
pregnancy, American Journal of Physiology 272 (1997), pp. R4212013;463.
Smith and DeWitt, 1996 W.L. Smith and D.L. DeWitt, Prostaglandin endoperoxide
synthases-1 and -2, Advances in Immunology 62 (1996), pp. 1672013;215. Abstract-
Spencer and Bazer, 1995 T.E. Spencer and F.W. Bazer, Temporal and spatial alterations
in uterine estrogen receptor and progesterone receptor gene expression during the estrous
cycle and early pregnancy in the ewe, Biology of Reproduction 53 (1995), pp.
12432013;1527.
Spencer and Bazer, 2002 T.E. Spencer and F.W. Bazer, Biology of progesterone action
during pregnancy recognition and maintenance of pregnancy, Frontiers Bioscience 7
(2002), pp. d18792013;1898.
Spencer et al., 1995a T.E. Spencer, W.C. Becker, P. George, M.A. Mirando, T.F. Ogle
and F.W. Bazer, Ovine interferon-tau regulates expression of endometrial receptors for
estrogen and oxytocin but not progesterone, Biology of Reproduction 53 (1995), pp.
$Order Document
Spencer et al., 1995b T.E. Spencer, W.C. Becker, P. George, M.A. Mirando, T.F. Ogle
and F.W. Bazer, Ovine interferon-tau inhibits estrogen receptor up-regulation and
estrogen-induced luteolysis in cyclic ewes, Endocrinology 136 (1995), pp.
Spencer et al., 1996 T.E. Spencer, T.L. Ott and F.W. Bazer, Tau-interferon: pregnancy
signal in ruminants, Proceedings Experimental Biology and Medicine 213 (1996), pp.
Sreenan, 1976 J.M. Sreenan, Egg transfer in the cow: effect of site of transfer,
Proceedings 8th International Congress on Animal Reproduction and A.I. 3 (1976), pp.
2692013;272.
Stabenfeldt et al., 1970 G. Stabenfeldt, M. Drost and C. Franti, Peripheral progesterone
levels in the ewe during pregnancy and parturition, Endocrinology 90 (1970) (1970), pp.
1442013;149.
Stefanczyk et al., 1990 S. Stefanczyk, T. Krzymowski, N. Einer-Jensen, T. Kaminski and

J. Kotwica, Local transfer of prostaglandin F23b1; from the uterine lumen into the venous
and arterial blood and into the uterine, mesometrial and ovarian tissue on day 18 of
pregnancy in the pig, Animal Reproduction Science 23 (1990), pp. 2232013;235.
Stepien et al., 1999 A. Stepien, M. Shemesh and A.J. Ziecik, Luteinizing hormone
receptor kinetic and LH-induced prostaglandin production throughout the oestrous cycle
in porcine endometrium, Reproduction, Nutrition, and Development 39 (1999), pp.
$Order Document
Stepien and Ziecik, 2002 A. Stepien and A.J. Ziecik, Second messenger systems in the
action of LH and oxytocin on porcine endometrial cells in vitro, Theriogenology 57
Stewart et al., 2003 A.B. Stewart, E.K. Inskeep, E.C. Townsend and R.A. Dailey, Effects
of gram-positive bacterial pathogens in ewes: peptidoglycan as a potential mediator of
interruption of early pregnancy, Reproduction 125 (2003), pp. 2952013;299. Abstract-
Stock and Fortune, 1993 A. Stock and J.E. Fortune, Ovarian follicular dominance:
relationship between prolonged growth of the ovulatory follicle and endocrine
parameters, Endocrinology 132 (1993), pp. 11082013;1114. Abstract-EMBASE |
Abstract-MEDLINE | $Order Document | Full Text via CrossRef
Stouffer et al., 2001 R.L. Stouffer, J.C. Martinez-Chequer, T.A. Molskness, F. Xu and
T.M. Hazzard, Regulation and action of angiogenic factors in the primate ovary, Archives
of Medical Research 32 (2001), pp. 5672013;575. SummaryPlus | Full Text + Links |
PDF (198 K)
Taverne et al., 2001 M.M. Taverne, N.C. de Schwartz, M. Kankofer, M.M. Bevers, H.A.
van Oord, D. Schams, S. Gutjahr and G.C. van der Weijden, Uterine responses to
exogenous oxytocin before and after pre-partum luteolysis in the cow, Reproduction
Domestic Animals 36 (2001), pp. 2672013;272. Abstract-Elsevier BIOBASE | Abstract-
Thatcher et al., 1995 W.W. Thatcher, M.D. Meyer and G. Danet-Desnoyers, Maternal
recognition of pregnancy, Journal of Reproduction and Fertility 49 (1995) (Suppl.), pp.
Thatcher et al., 2001 W. Thatcher, A. Guzeloglu, R. Mattos, M. Binelli, T. Hansen and J.
Pru, Uterine-conceptus interaction and reproductive failure in cattle, Theriogenology 56
Thorburn, 1991 G.D. Thorburn, The placenta, prostaglandins and parturition: a review,
Reproduction, Fertility, and Development 3 (1991), pp. 2772013;299.
Thorburn and Rice, 1990 G.D. Thorburn and G. Rice, Placental PGE2 and the initiation
of parturition in the sheep In: M. Mitchell, Editors, Eicosanoids in Reproduction, CRC
Press, Boca Raton, FL, USA (1990), pp. 742013;82.
Tolleson and Randel, 1988 D.R. Tolleson and R.D. Randel, Effects of alfaprostol and
uterine palpation on postpartum interval and pregnancy rate to embryo transfer in
Brahman influenced beef cows, Theriogenology 29 (1988), pp. 5552013;564.
Tranguch et al., 2002 S. Tranguch, N. Steuerwald and Y.M. Huet-Hudson, Nitric oxide
synthase production and nitric oxide regulation of preimplantation embryo development,
Biology of Reproduction 68 (2002), pp. 15382013;1544. Full Text via CrossRef
Trapp and Slyter, 1979 M.J. Trapp and A.L. Slyter, Pregnancy rates in ewes at first
breeding, Journal of Animal Science 59 (1979) (Suppl. 2), pp. 4652013;479.
Tsai et al., 1996 S.J. Tsai, M.C. Wiltbank and K.J. Bodensteiner, Distinct mechanisms
regulate induction of messenger ribonucleic acid for prostaglandin (PG) G/H synthase-2,
PGE (EP3) receptor, and PGF2 alpha receptor in bovine preovulatory follicles,
Endocrinology 137 (1996), pp. 33482013;3355. Abstract-MEDLINE | $Order
Document | Full Text via CrossRef
Tsai and Wiltbank, 1997a S.J. Tsai and M.C. Wiltbank, Prostaglandin F23b1; induces
expression of prostaglandin G/H synthase-2 in the ovine corpus luteum: a potential
positive feedback loop during luteolysis, Biology of Reproduction 57 (1997), pp.
Tsai et al., 1997b S.J. Tsai, J.L. Juengel and M.C. Wiltbank, Hormonal regulation of
monocyte chemoattractant protein-1 messenger ribonucleic acid expression in corpora
lutea, Endocrinology 138 (1997), pp. 45172013;4520. Abstract-EMBASE | Abstract-
Tsai et al., 2001 S.J. Tsai, K. Kot, O.J. Ginther and M.C. Wiltbank, Temporal gene
expression in bovine corpora lutea after treatment with prostaglandin F 2 alpha based on
serial biopsies in vivo, Reproduction 121 (2001), pp. 9052013;913. Abstract-EMBASE |
CrossRef
Tsang et al., 1979 B.K. Tsang, L. Ainsworth, B.R. Downey and D.T. Armstrong,
Preovulatory changes in cyclic AMP and prostaglandin concentrations in follicular fluid
of gilts, Prostaglandins 17 (1979), pp. 1412013;149.
Tsang et al., 1988 B.K. Tsang, J. Arodi, M. Li and L. Ainsworth, Gonadotropic

regulation of prostaglandin production by ovarian follicular cells of the pig, Biology of
Reproduction 38 (1988), pp. 6272013;638.
Vane and Botting, 1995 J.R. Vane and R.M. Botting, A better understanding of anti-
inflammatory drugs based on isoforms of cyclooxygenase (COX-1 and COX-2),
Advances in Prostaglandin, Thromboxane, and Leukotriene Research 23 (1995), pp.
Vanderwall et al., 2003 D. Vanderwall, J. Williams and G. Woods, Equine conceptuses

did not suppress OT-induced endometrial PGF23b1; secretion, Biology of Reproduction 68
(2003) (Suppl. 1, Abstract), p. 161.
Vasconcelos et al., 1999 J.L. Vasconcelos, R.W. Silcox, G.J.M. Rosa, J.R. Pursley and
M.C. Wiltbank, Synchronisation rate, size of the ovulatory follicle, and pregnancy rate
after synchronisation of ovulation beginning on different days of the estrous cycle in
lactating dairy cows, Theriogenology 52 (1999), pp. 10672013;1078. Abstract | PDF (685
K)
Vedemikov et al., 2000 Y. Vedemikov, A.S. Syal, T. Okawa, V. Jain, G.R. Sadde and
R.E. Garfield, The role of cyclic nucleotides in the spontaneous contractility and
responsiveness to nitric oxide of the rat uterus at mid-gestation and term, American
Journal of Obstetrics and Gynecology 182 (2000), pp. 6122013;619.
Velez et al., 1991 J.S. Velez, R.D. Randel and D.A. Neuendorff, Effect of uterine
manipulation on postpartum fertility and plasma 13, 14-dihydro-15-keto prostaglandin
F23b1; on Brahman cows and first calf heifers, Theriogenology 36 (1991), pp.
9872013;997.
Viergutz et al., 2000 T. Viergutz, B. Loehrke, R. Poehland, F. Becker and W. Kanitz,

Relationship between different stages of the corpus luteum and the expression of the
peroxisome proliferators-activated receptor gamma protein in bovine large lutein cells,
CrossRef
Vincent and Inskeep, 1986 D.L. Vincent and E.K. Inskeep, Role of progesterone in
regulating utero-ovarian venous concentrations of PGF23b1; and PGE2 during the estrous
cycle and early pregnancy in ewes, Prostaglandins 31 (1986), pp. 7152013;733.
Vincent et al., 1986 D.L. Vincent, S. Meredith and E.K. Inskeep, Advancement of uterine
secretion of prostaglandin E2 by treatment with progesterone and transfer of
asynchronous embryos, Endocrinology 119 (1986), pp. 5272013;529. Abstract-EMBASE
Vincent et al., 1992 D.L. Vincent, K.R.C. Gemmer, M. Thamotharan, Y. Tanaka, C.N.
Lee and C.W. Weems, Oviductal vein resection affects pregnancy rates in sheep, Chinese
Journal of Physiology 34 (1992), pp. 2432013;254.
Vujosevic et al., 1988 J. Vujosevic, M. Zeremski, I. Koremic and D. Kuzmanov, The

effect of injection of synthetic prostaglandin F23b1; (Estrumate) in disturbed puerperium
on some reproductive measurements in cows, Proceedings 10th International Congress
on Animal Reproduction and A.I. III (1988), pp. 4332013;435.
Wagner et al., 1974 W.C. Wagner, F.N. Thompson, L.E. Evans and E.C.I. Molokwu,
Hormonal mechanisms controlling parturition, Journal of Animal Science 39 (1974)
(Suppl.), pp. 392013;57.
Wann and Randel, 1990 R.A. Wann and R.D. Randel, Effect of uterine manipulation 35
days after parturition on plasma concentrations of 13,14-dihydo-15-keto prostaglandin
F23b1; in multiparous and primiparous Brahman cows, Journal of Animal Science 68
Watson and Hinrichs, 1988 E.D. Watson and K. Hinrichs, Changes in the concentrations
of steroids and prostaglandin F in preovulatory follicles of the mare after administration
of hCG, Journal of Reproduction and Fertility 84 (1988), pp. 5572013;565.
Watson and Sertich, 1991 E.D. Watson and P.L. Sertich, Concentrations of arachidonate
metabolites, steroids and histamine in preovulatory horse follicles after administration of
human chorionic gonadotropin and the effect of intrafollicular injection of indomethacin,
Journal of Endocrinology 129 (1991), pp. 1312013;139. Abstract-EMBASE | Abstract-
Weber et al., 1992 J. Weber, D. Freeman, D. Vanderwall and G. Woods, PGE2 secretion
by oviductal transport-stage specific equine embryos, Biology of Reproduction 45 (1992),
pp. 5402013;543.
Weems et al., 1975 C.W. Weems, J.E. Pexton, R.L. Butcher and E.K. Inskeep,
Prostaglandins F in uterine tissue and venous plasma of pseudopregnant rats: effect of
deciduomata, Biology of Reproduction 13 (1975), pp. 2822013;288. Abstract-EMBASE |
Weems, 1979 C.W. Weems, Prostaglandins F in uterine and ovarian compartments and in
plasma from the uterine vein, ovarian artery and vein, and abdominal aorta of
pseudopregnant rats with and without deciduomata, Prostaglandins 17 (1979), pp.
Weems et al., 1985a C.W. Weems, G.L. Hoyer, R.R. Magness, G.L. Whysong and M.
Huie, Effects of prostaglandin E2 (PGE2) on estradiol-173b2;-induced luteolysis in the
nonpregnant ewe, Prostaglandins 29 (1985), pp. 2332013;241. Abstract-EMBASE |
Weems et al., 1985b C.W. Weems, M. Huie, G.L. Hoyer, R.R. Magness and G.L.
Whysong, Maternal recognition of pregnancy in ruminants In: W.C. Davis, J.N. Shelton
and C.W. Weems, Editors, Characterization of the Bovine Immune System and the Genes
Regulating Expression of Immunity With Particular Reference to Their Role in Disease
Resistance, Washington State University Press, Pullman, WA (1985), pp. 1932013;210.
Weems et al., 1985c C.W. Weems, M. Huie, R.R. Magness, G.L. Hoyer and G.L.
Whysong, Inhibition of an IUD-induced precocious luteolysis by prostaglandin E 1
(PGE1) in sheep, Prostaglandins 30 (1985), pp. 5732013;580. Abstract-EMBASE |
Weems et al., 1985d C.W. Weems, M. Huie, R.R. Magness, G.L. Hoyer and G.L.
Whysong, Prostaglandin E2 (PGE2) inhibits an IUD-induced premature luteolysis in
sheep, Prostaglandins, Leukotrienes, and Medicine 20 (1985), pp. 2672013;273.
Weems et al., 1985e C.W. Weems, L.P. Reynolds, J.M. Huie, G.L. Hoyer and H.R.
Behrman, Effects of prostaglandins E1 or E2 (PGE1; PGE2) on luteal function and binding
of luteinizing hormone in nonpregnant ewes, Prostaglandins 29 (1985), pp. 1612013;173.
Weems et al., 1988 C. Weems, Y. Weems, C. Lee and D. Vincent, Distribution of

progesterone to the uterus and associated vasculature of cattle, Endocrinologia Japonica
35 (1988), pp. 1512013;156.
Weems et al., 1989a C.W. Weems, Y.S. Weems, C.N. Lee, J. Hall, B. DeGroot and D.L.
Vincent, Effects of chronically administered dibutyryl-cAMP or adenosine on luteal
function in sheep, Endocrinologia Japononica 36 (1989), pp. 1512013;159.
Weems et al., 1989b C.W. Weems, Y.S. Weems, C.N. Lee and D.L. Vincent,
Progesterone in uterine and arterial tissue and in jugular and uterine venous plasma of
sheep, Biology of Reproduction 41 (1989), pp. 12013;6. Abstract-EMBASE | Abstract-
Weems et al., 1991 C.W. Weems, Y.S. Weems, C.N. Lee and D.L. Vincent, The purine
adenosine amplifies the response to gonadotropins and inhibits PGF 23b1; suppression of
gonadotropin stimulation of bovine luteal cells in vitro, Chinese Journal of Physiology 34
(1991), pp. 3552013;371.
Weems et al., 1992a C.W. Weems, Y.S. Weems, C.N. Lee and D.L. Vincent, Adenosine
facilitates the response to hCG, PGE1 or PGE2 and inhibits the response to PGF23b1; by
hCG-stimulated ovine luteal cells in vitro, Prostaglandins 43 (1992), pp. 2232013;237.
Weems et al., 1992b C.W. Weems, D.L. Vincent and Y.S. Weems, Roles of
prostaglandins (PG), F23b1;, E1, E2, adenosine, oestradiol-173b2;, histone-H2A and
progesterone of conceptus, uterine or ovarian origin during early and mid-pregnancy in
the ewe, Reproduction Fertility and Development 4 (1992), pp. 2892013;295. Abstract-
Weems et al., 1992c C.W. Weems, Y.S. Weems, C.N. Lee and D.L. Vincent, Purine
modulation of prostaglandin F23b1; (PGF23b1;)-induced premature luteolysis in vivo in
sheep, Second Messengers and Phosphoproteins 14 (1992), pp. 12013;9. Abstract-
Weems et al., 1992d Y.S. Weems, D.L. Vincent, Y. Tanaka, K. Miller-Patrick, K.D.
Nusser, D. Ashimine and C.W. Weems, Effects of prostaglandin F 23b1; on sources of
progesterone and pregnancy in intact, ovariectomized, and hysterectomized 902013;100
day pregnant ewes, Prostaglandins 43 (1992), pp. 2032013;222. Abstract-EMBASE |
Weems et al., 1993 Y.S. Weems, D.L. Vincent, Y. Tanaka, K.D. Nusser, K.S.
Ledgerwood and C.W. Weems, Effect of prostaglandin F23b1; on uterine or ovarian
secretion of prostaglandins E and F23b1; (PGE; PGF23b1;) in vivo in 902013;100 day
hysterectomized, intact or ovariectomized pregnant ewes, Prostaglandins 46 (1993), pp.
Weems et al., 1994a Y.S. Weems, D.L. Vincent, K.D. Nusser, Y. Tanaka, K. Miller-
Patrick, K.S. Ledgerwood and C.W. Weems, Effects of prostaglandin F 23b1; (PGF23b1;) on
secretion of estradiol-173b2; and cortisol in 902013;100 day hysterectomized, intact or
ovariectomized pregnant ewes, Prostaglandins 48 (1994), pp. 1392013;157.
Weems et al., 1994b Y.S. Weems, R.G. Sasser, D.L. Vincent, K.D. Nusser, Y. Tanaka,
K. Miller-Patrick, K.S. Ledgerwood and C.W. Weems, Effects of prostaglandin F 23b1;
(PGF23b1;) on secretion of pregnancy specific protein B (PSPB) by 902013;100 day intact
and ovariectomized pregnant ewes, Prostaglandins 48 (1994), pp. 3772013;387.
Document
Weems et al., 1995 C.W. Weems, Y.S. Weems and D.L. Vincent, Maternal recognition
of pregnancy and maintenance of gestation in sheep In: G. Enne, G.F. Greppi and A.
Lauria, Editors, Proceedings XXX International Simposia Societa Italiana Zootechnica,
Milan, Italy, Elsevier, Amsterdam, Holland (1995), pp. 2772013;293.
Weems et al., 1996 C.W. Weems, Y.S. Weems, L.N. French, J.T. French, R.G. Sasser, T.
Welsh, P.G. Harms and R.D. Randel, Effect of histone-H2A (H-H2A), platelet activating
factor (PAF), and pregnancy specific protein B (PSPB) on secretion of prostaglandins E
and F23b1; (PGE; PGF23b1;) by nonpregnant bovine endometrium and H-H2A on basal
release of luteinizing hormone (LH) by bovine pituitary cells in vitro, Chinese Journal of
Physiology 39 (1996), pp. 12013;8.
Weems et al., 1997 Y.S. Weems, P.J. Bridges, R.G. Sasser, B.R. LeaMaster, D.L.
Vincent and C.W. Weems, Prostaglandin E1 (PGE1) or prostaglandin E2 (PGE2), not
luteinizing hormone (LH), regulates secretion of progesterone by the ovine corpus luteum
of pregnancy, Prostaglandins 53 (1997), pp. 3372013;353. SummaryPlus | Full Text +
Links | PDF (889 K)
Weems et al., 1998a Y.S. Weems, M.A. Lammoglia, H. Vera-Avila, R.D. Randel, R.G.
Sasser and C.W. Weems, Effect of luteinizing hormone (LH), PGE 2, 8-Epi-PGE1, 8-Epi-
PGE2, trichosanthin and pregnancy specific protein B (PSPB) on secretion of
progesterone in vitro by the corpus luteum (CL) from nonpregnant and pregnant cows,
Prostaglandins 55 (1998), pp. 282013;43.
Weems et al., 1998b Y.S. Weems, M.A. Lammoglia, H. Vera-Avila, R.D. Randel, R.G.
Sasser and C.W. Weems, Effect of luteinizing hormone (LH), PGE 2, 8-Epi-PGE1, 8-Epi-
PGE2, trichosanthin and pregnancy specific protein B (PSPB) on secretion of
prostaglandin E (PGE) or PGF23b1; in vitro by the corpus luteum (CL) from nonpregnant
and pregnant cows, Prostaglandins 55 (1998), pp. 3592013;376. SummaryPlus | Full
Text + Links | PDF (426 K)
Weems et al., 1999a Y.S. Weems, D.L. Vincent, C. Lemme and C.W. Weems, Trilostane,
but not prostaglandin F23b1; (PGF23b1;) or cortisol, aborts 902013;100 day lutectomized
pregnant sheep, Prostaglandins and Other Lipid Mediators 58 (1999), pp. 772013;86.
Weems et al., 1999b Y.S. Weems, P.J. Bridges, B.R. LeaMaster, R.G. Sasser, D.L.
Vincent and C.W. Weems, Secretion of progesterone, estradiol-17 3b2;, prostaglandins
(PG) E (PGE), F23b1; (PGF23b1;), and pregnancy specific protein B (PSPB) by day 90
intact or ovariectomized pregnant ewes, Prostaglandins and Other Lipid Mediators 58
Weems et al., 1999c Y.S. Weems, P.J. Bridges, B.R. LeaMaster, D.L. Vincent and C.W.
Weems, Effect of trilostane on prostaglandins (PG) E and F23b1; (PGE; PGF 23b1;),
estradiol-173b2; and progesterone secretion and pregnancy of day 90 intact or
ovariectomized pregnant ewes, Prostaglandins and Other Lipid Mediators 58 (1999), pp.
Weems et al., 1999d Y.S. Weems, M. Lammoglia, R.D. Randel, R.G. Sasser and C.W.
Weems, PGE2 induces its own secretion in vitro by bovine 270-day placenta, but, not by
200-day placenta, Prostaglandins and Other Lipid Mediators 57 (1999), pp.
Weems et al., 2001 Y.S. Weems, P.J. Bridges, R.G. Sasser, B.R. LeaMaster, L. Ching
and C.W. Weems, Effect of an aromatase inhibitor, CGS-16949A, on secretion of
estradiol-173b2;, pregnancy specific protein B (PSPB), progesterone, and prostaglandins E
and F23b1; (PGE; PGF23b1;) in 90-day ovariectomized pregnant sheep, Prostaglandins and
Other Lipid Mediators 66 (2001), pp. 772013;87.
Weems et al., 2002a Y.S. Weems, A.W. Lewis, R.D. Randel and C.W. Weems, Effects of
prostaglandins E2 and F23b1; (PGE2; PGF23b1;), trilsotane, mifepristone, palmitic acid (PA),
indomethacin (INDO), ethamoxytriphetol (MER-25), PGE 2 + PA, or PGF23b1; on PGE2,
PGF23b1;, and progesterone by bovine corpora lutea of mid-pregnancy in vitro, Chinese
Journal of Physiology 45 (2002), pp. 1632013;168. Abstract-EMBASE | Abstract-
Weems et al., 2002b Y.S. Weems, P.J. Bridges, R.G. Sasser, B.R. LeaMaster and C.W.
Weems, Effects of mifepristone on pregnancy and secretion of progesterone, estradiol-
173b2;, pregnancy specific protein B (PSPB), and prostaglandins E and F 23b1; (PGE;
PGF23b1;) in 90-day intact pregnant sheep, Prostaglandins and Other Lipid Mediators 70
Weems et al., 2003a C.W. Weems, Y.S. Weems, L. Kim, V. Humphreys and V. Tsuda,
Effect of luteinizing hormone (LH), pregnancy specific protein B (PSPB), or arachidonic
acid (AA) on secretion of progesterone (P4), PGF23b1;, or PGE2 by corpora lutea (CL) of
the estrous cycle or pregnancy, Biology of Reproduction 68 (2003) (Suppl. 1, Abstract),
p. 138.
Weems et al., 2003b Y.S. Weems, L. Kim, V. Humphreys, V. Tsuda and C.W. Weems,
Effect of luteinizing hormone (LH), pregnancy specific protein B (PSPB), or arachidonic
acid (AA) on ovine endometrium of the estrous cycle or placental secretion of
prostaglandins E2 (PGE2) and F23b1; (PGF23b1;), and progesterone in vitro, Prostaglandins
and Other Lipid Mediators 71 (2003), pp. 552013;73. SummaryPlus | Full Text + Links |
PDF (382 K)
Weems et al., 2004a C.W. Weems, Y.S. Weems, S. Tatman, A. Lewis, D.A. Neuendorff
and R.D. Randel, Does estrous synchronisation affect corpus luteum (CL) function?,
Prostaglandins and Other Lipid Mediators 74 (2004), pp. 452013;49.
Weems et al., 2004b C.W. Weems, Y.S. Weems, G. Carstens and R.D. Randel, Do
calcium-mediated cellular signalling pathways, PGE 2, estrogen or progesterone receptor
antagonists, or bacterial toxins affect bovine placental function in vitro?, Prostaglandins
and Other Lipid Mediators 73 (2004), pp. 2652013;278.
Weems et al., 2004c C. Weems, Y. Weems and C. Yin, What regulates ovine placental
(PL) steroidogenesis?, Journal of Animal Science 71 (2004) (Suppl. 1, Abstract), p. 257.
Weems et al., 2004d C.W. Weems, Y.S. Weems, E. Lennon, T. Uchima, A. Ong, A.
Raney, K. Goto and Y. Zaleski, Is nitric oxide luteolytic or antiluteolytic in sheep,
Biology of Reproduction 83 (2004) (Suppl. 1, Abstract), p. 440.
Weipz et al., 1992 G.J. Weipz, M.C. Wiltbank, T. Nett, G.D. Niswender and H.R.
Sawyer, Receptors for prostaglandin F23b1; and E2 in ovine corpora lutea during maternal
recognition of pregnancy, Biology of Reproduction 47 (1992), pp. 9842013;991.
Weipz et al., 1993 G.J. Weipz, M.C. Wiltbank, S.B. Kater, G.D. Niswender and H.R.
Sawyer, PGE2 attenuates PGF23b1;-induced increases in free intracellular calcium in ovine
large luteal cells, Prostaglandins 45 (1993), pp. 1672013;176.
White and Dobson, 1990 A.J. White and H. Dobson, Effect of prostaglandin F23b1; on the
fertility of cows after calving, Veterinary Record 127 (1990), p. 588. Abstract-EMBASE |
Whiteaker et al., 1995 S. Whiteaker, M. Mirando, W. Becker and D. Peters, Relationship

between phosphoinositide hydrolysis and prostaglandin F 23b1;secretion in vitro from
endometrium of cyclic pigs on day 15 postestrus, Domestic Animal Endocrinology 12
Wiesak et al., 1992 T. Wiesak, M. Hunter and G. Foxcroft, Effect of prostaglandins on

luteal function during early pregnancy in pigs, Journal of Reproduction and Fertility 95
Wijayagunawardane et al., 1998 M.P. Wijayagunawardane, A. Miyamoto, W.A. Cerbito,

T.J. Acosta, M. Takagi and K. Sato, Local distributions of oviductal estradiol,
progesterone, prostaglandins, oxytocin and endothelin-1 in the cyclic cow,
Theriogenology 49 (1998), pp. 6072013;618. SummaryPlus | Full Text + Links | PDF
(572 K)
Wijayagunawardane et al., 1999 M.P. Wijayagunawardane, A. Miyamoto and K. Sato,

Prostaglandin E2, prostaglandin F23b1;, and endothelin-1 production by cow oviductal
epithelial cell monolayers: effect of progesterone, estradiol-17 beta, oxytocin and
luteinizing hormone, Theriogenology 52 (1999), pp. 7912013;801. Abstract | PDF (465
K)
Wijayagunawardane et al., 2001 M.P. Wijayagunawardane, A. Miyamoto, Y.

Taquahashi, T.J. Acosta, M. Nishimura and K. Sato, Angiotensin and atrial natriuretic
peptide in the cow oviductal contraction in vitro: direct effect and local secretion of
prostaglandins, endothelin-1, and angiotensin II, Biology of Reproduction 65 (2001), pp.
$Order Document
Wilson et al., 1972a L. Wilson, R.J. Cenedella, R.L. Butcher and E.K. Inskeep, Levels of
prostaglandins in the uterine endometrium during the ovine estrous cycle, Journal of
Animal Science 34 (1972), pp. 932013;101.
Wilson et al., 1972b L. Wilson, R.L. Butcher and E.K. Inskeep, Prostaglandin F 23b1; in the
uterus of ewes during early pregnancy, Prostaglandins 1 (1972), pp. 4792013;487.
Wiltbank et al., 1991 M. Wiltbank, M. Diskin and G.D. Niswender, Differential actions
of second messenger systems in the corpus luteum, Journal of Reproduction and Fertility
43 (1991) (Suppl.), pp. 652013;75. Abstract-MEDLINE | $Order Document
Wiltbank et al., 1992 M.C. Wiltbank, G.J. Weipz, J.J. Knickerbocker, C. Belfiore and
G.D. Niswender, Proteins secreted from the early ovine conceptus block the action of
prostaglandin F2 on large luteal cells, Biology of Reproduction 46 (1992), pp.
Wiltbank et al., 1995 M.C. Wiltbank, T.F. Shiao, D.R. Bergfelt and O.J. Ginther,
Prostaglandin F2 alpha receptors in the early bovine corpus luteum, Biology of
Reproduction 52 (1995), pp. 742013;78. Abstract-EMBASE | Abstract-Elsevier
Wray, 1993 S. Wray, Uterine contraction and physiological mechanisms of modulation,

American Journal of Physiology 26 (1993), pp. 12013;16.
Wuttke et al., 1998 W. Wuttke, S. Spiess, I. Knoke, L. Pitzel, S. Leonhard and H. Jarry,
Synergistic effects of prostaglandin F23b1; and tumor necrosis factor to induce luteolysis
in the pig, Biology of Reproduction 58 (1998), pp. 13102013;1315. Abstract-EMBASE |
Yallampalli and Garfield, 1993 C. Yallampalli and R.E. Garfield, Inhibition of nitric
oxide synthesis in rats during pregnancy produces signs similar to those of preeclampsia,
American Journal of Obstetrics and Gynecology 169 (1993), pp. 13162013;1320.
Yallampalli et al., 1998 C. Yallampalli, Y.L. Dong, P.R. Gangula and L. Fang, Role and
regulation of nitric oxide on the uterus during pregnancy and parturition, Journal Society
of Gynecological Investigation 5 (1998), pp. 582013;67. Abstract | PDF (876 K)
Yallampalli and Dong, 2000 C. Yallampalli and Y.L. Dong, Estradiol-173b2; inhibits
nitric oxide synthase (NOS)-II and stimulates NOS-III gene expression in the rat uterus,
Yee and Kennedy, 1991 G. Yee and T.G. Kennedy, Role of cyclic adenosine
32032;,52032; monophosphate in mediating the effect of PGE on decidulization in vitro,
Young et al., 1984 I.M. Young, D.B. Anderson and R.W. Plenderleith, Increased
conception rate in dairy cows after early postpartum administration of prostaglandin
F23b1;, Veterinary Record 115 (1984), pp. 4292013;431. Abstract-EMBASE | Abstract-
Young and Anderson, 1986 I.M. Young and D.B. Anderson, First service conception rate
in dairy cows treated with dinoprost tromethamine early postpartum, Veterinary Record
118 (1986), pp. 2122013;213. Abstract-EMBASE | Abstract-MEDLINE | $Order
Document
Zarco et al., 1988a L. Zarco, G.H. Stabenfeldt, J. Quirke, H. Kindahl and G.E. Bradford,
Release of prostaglandin F23b1; and the timing of events associated with luteolysis in ewes
with estrous cycles of different lengths, Journal of Reproduction and Fertility 83 (1988),
pp. 5172013;526. Abstract-EMBASE | Abstract-MEDLINE | $Order Document
Zarco et al., 1988b L. Zarco, G.H. Stabenfeldt, S. Basu, G.E. Bradford and H. Kindahl,
Modification of prostaglandin F23b1; synthesis and release in the ewe during the initial
establishment of pregnancy, Journal of Reproduction and Fertility 83 (1988), pp.
Zhang and Davis, 1991 Z. Zhang and D.L. Davis, Prostaglandins E and F23b1; secretion by
glandular and stromal cells of the pig endometrium in vitro: effects of estradiol-17 beta,
progesterone, and day of pregnancy, Prostaglandins 42 (1991), pp. 1512013;162.
Zhang et al., 1992 Z. Zhang, M. Krause and D.L. Davis, Epidermal growth factor
receptors in porcine endometrium: binding characteristics and the regulation of
prostaglandin E and F2 production, Biology of Reproduction 46 (1992), pp. 9322013;942.
Zhang et al., 1999 J. Zhang, G.A. Massmann, C.P. Mirabile and J. Figueroa, Nonpregnant
sheep uterine type I and II nitric oxide synthase expression is differentially regulated by
estrogen, Biology of Reproduction 60 (1999), pp. 11982013;1203. Abstract-EMBASE |
Zhang et al., 2000 J. Zhang, Y. Li, A.R. Weiss, I.M. Bird and R.R. Magness, Expression
of endothelial and inducible nitric oxide synthases and nitric oxide production in ovine
placental and uterine tissues during late pregnancy, Placenta 21 (2000), pp. 5162013;524.
Ziecik et al., 1986 A.J. Ziecik, P.D. Stanchev and J.E. Tilton, Evidence for the presence
of luteinizing hormone/human chorionic gonadotropin binding sites in the porcine uterus,
Endocrinology 119 (1986), pp. 11592013;1163. Abstract-EMBASE | Abstract-
Ziecik, 2000 A.J. Ziecik, Importance of endometrial hormone receptors in induction of

luteolysis and maternal recognition of pregnancy in the pig, Reproduction Domestic
Animals 35 (2000), pp. 1902013;192. Abstract-Elsevier BIOBASE | $Order Document |
Ziecik et al., 2001 A.J. Ziecik, G. Bodek, A. Stepien, R. Ciereszko and G. Kotwica,
Involvement of gonadotropins in induction of luteolysis in pigs, Reproductive Biology 1
Ziecik, 2002 A.J. Ziecik, Old, new and the newest concepts of inhibition of luteolysis
during early pregnancy in pig, Domestic Animal Endocrinology 23 (2002), pp.
Zor et al., 1970 A. Zor, T. Kaneko, H.P.G. Schneider, S.M. McCann and J.B. Field,
Further studies on stimulation of anterior pituitary cyclic adrenosine-3,52032;-
monophosphate formation by the hypothalamic extract and prostaglandins, Journal of
Biological Chemistry 245 (1970), pp. 28832013;2885.

Prostaglandins and Reproduction Infemale Animals

Uploaded by

Document Information

Original Description:

Original Title

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Prostaglandins and Reproduction Infemale Animals

Uploaded by

Copyright:

Available Formats

Prostaglandins and reproduction in female farm

animals The Veterinary Journal172:206-228, 2006.

Accepted 20 November 2004. Available online 26 January 2005.

Keywords: Prostaglandins; Reproduction; Corpus luteum; Pregnancy; Placenta;

Pharmaceutical agents influence PG metabolism. Aspirin or indomethacin inhibits COX-

3. Follicular growth and ovulation

Indomethacin inhibition of ovulation can be overcome by PGE 2 or PGF23b1; (Murdoch,

Nordihydroguaratic acid, a lipoxygenase inhibitor, decreases LT and leukocyte

Understanding luteolysis requires an understanding of the anatomical relationships

The source of the luteolysin is the endometrium. Destruction of the endometrium or

An intrauterine device (IUD) adjacent or opposite to the side of ovulation on days

Indomethacin or destruction of follicles, the source of oestrogen, at midcycle delays

Although ovine endometrial progesterone receptors decline by day 12 post-oestrous,

In sows and cows, monocyte chemoattractant protein-1 causes macrophages and

6. Maternal recognition and maintenance of pregnancy

At the first postpartum ovulation in cows, an embryo-toxic factor is secreted by the

Establishment and maintenance of pregnancy is dependent on changes in a plethora of

Concentrations of PGF23b1; in uterine venous, or ovarian arterial blood, or in luteal tissue

In cows, endometrial prostaglandin E-synthase mRNA is expressed from days 18 to 50

Progesterone is required throughout pregnancy (Weems et al., 1995). Fetal growth

Circulating oestradiol-173b2;, PGE2, and PSPB increase as pregnancy progresses and

Interaction between the uterus, the hypothalamic2013;pituitary2013;ovarian axis, and

Controlled internal drug-releasing devices (CIDR) to deliver progesterone have been

Bauer et al., 2001 M. Bauer, I. Reibiger and K. Spanel-Borowski, Leucocyte proliferation

Bekana et al., 1996a M. Bekana, P. Jonsson and H. Kindahl, Intrauterine bacterial

Bekana et al., 1996b M. Bekana, K. Odensvik and H. Kindahl, Prostaglandin F 2 alpha

Breeveld-Dwarkasing et al., 2003 V.N.A. Breeveld-Dwarkasing, J.M. Koppele, R.A.

Cavicchio et al., 2002 V. Cavicchio, J. Pru, B. Davis, J. Davis, B. Rueda and D.

Chwalisz et al., 1999 K. Chwalisz, E. Winterhager, T. Thienel and R.E. Garfield,

De La Llosa-Hermier et al., 1991 M.P. De La Llosa-Hermier, J. Martal, A. Ricour and C.

Freidman et al., 1995 S. Freidman, M. Gurevich and M. Shemesh, Bovine cyclic

Geisert et al., 1994 R. Geisert, R. Short and G. Morgan, Establishment of pregnancy in

Gleicher et al., 1992 N. Gleicher, L. Harlow and M. Zilberstein, Regulatory effect of

Grusenmeyer and Pate, 1992 D. Grusenmeyer and J. Pate, Localization of prostaglandin

Gurevich et al., 1993 M. Gurevich, E. Harel-Markowitz, S. Marcus, L.S. Shore and M.

Jaroszewki and Hansel, 2000 J. Jaroszewki and W. Hansel, Intraluteal administration of a

Kankover and Wiercinski, 1999 M. Kankover and J. Wiercinski, Prostaglandin E 2-9-keto

Kobayashi et al., 2001 S. Kobayashi, B. Berisha, W.M. Amselgruber, D. Schams and A.

Kotwica et al., 1999 G. Kotwica, A. Franczak, S. Okrasa and J. Kotwica, Effect of an

Lefevre et al., 1998 F. Lefevre, Martinat-Botte, A. Locatelli, P. de Niu, M. Terqui and C.

Lukaszewska and Hansel, 1980 J. Lukaszewska and W. Hansel, Corpus luteum

Mirando et al., 1993 M. Mirando, W. Becker and S. Whiteaker, Relationship among

Miyamoto et al., 2002 Y. Miyamoto, R. Sakumoto, Y. Sakabe, M. Miyake, A. Okano and

Murdoch, 1985W.J. Murdoch, Follicular determinants of ovulation in the ewe, Domestic

Murdoch, 1986 W.J. Murdoch, Accumulation of thromboxane B 2 within periovulatory

Murdoch, 1988b W.J. Murdoch, Effect of a steroidal (prednisolone) and nonsteroidal

Murdoch, 1990 W.J. Murdoch, Localization and hormonal regulation of ovarian

Murdoch, 1992 W.J. Murdoch, Comparative morphometry and steroidogenic function of

Murdoch, 1994b W.J. Murdoch, Immunoregulation of mammalian fertility, Life Science

Murdoch, 1996a W.J. Murdoch, Microtubular dynamics in granulosa cells of

Narumiya, 1995 S. Narumiya, Structures, properties and distributions of prostanoid

Nett, 1987 T.M. Nett, Function of the hypothalamic2013;hypophyseal axis during

Novaro et al., 1996 V. Novaro, V. Rettori, E.T. Gonzalez, A. Jawerbaum, A. Faletti, G.

Okuda et al., 2003 K. Okuda, M.M. Bah, K.M. Deptula, I. Woclawek-Potocka, A.

Penny, 2000 L. Penny, Monocyte chemoattractant protein 1 in luteolysis, Reviews

Peters, 1989 A.R. Peters, Effect of prostaglandin F23b1; on hormone concentrations in

Platz-Christensen et al., 1997 J.J. Platz-Christensen, P. Pernevi and H. Bokstrom,

Sakamoto et al., 1995 K. Sakamoto, K. Miwa, T. Ezashi, E. Okuda-Ashitaka, T. Houtani,

Sakumoto et al., 2000 R. Sakumoto, B. Berisha, N. Kawate, D. Schams and K. Okuda,

Sander et al., 1982 H. Sander, T. Brunn, W. Elger and F. Ellendorff, Prostaglandin E

Sasser et al., 1989 R. Sasser, J. Crock and C. Ruder-Montegomery, Characteristics of

Schallenberger et al., 1985 E. Schallenberger, J. Ramp and D.L. Walters, Gonadotropins