Reproductive immunosupression and diet. An evolutionary perspective on pregnancy sickness and meat consumption

Pregnancy sickness, a suite of "symptoms" that frequently co-occur during pregnancy, may be an adaptation providing behavioral prophylaxis against infection. Maternal immunosupression, necessary for tolerance of the fetus, results in gestational vulnerability to pathogens. Throughout the period of maximal vulnerability, dietary behavior is significantly altered via changes in nausea susceptibility and olfaction and the development of marked aversions and cravings. Of food types, meat is both the most likely to carry pathogens and the principal target of gestational aversions and pregnancy taboos. Because meat was prominent in ancestral human diets but hygienic procedures that effectively eliminate the risk of meat-borne infection are recent, such pathogens likely constituted a source of selective pressure on pregnant females throughout human history. Both the relatively low protein and energy demands of the first trimester and the existense of nonmeat alternatives would have allowed for the evolution of time-limited gestational meat-avoidance mechanisms.Complementing these mechanisms, gestational cravings target substances that may influence immune functioning and affect the availability of iron in the gastro-intestinal tract, thereby limiting the proliferation of iron-dependent pathogens. Clinical and ethnographic findings are examined in light of these proposals, and directions for future research are outlined.     

Embryo quality: the missing link between pregnancy sickness and pregnancy outcome

Pregnancy sickness is widespread yet its etiology is poorly understood. It is almost certainly endocrine in origin and most likely a product of placental hormones, with human chorionic gonadotropin being the strongest candidate. It has long been known that greater levels of nausea and vomiting during pregnancy are associated with a lower incidence of spontaneous abortion, yet the causal mechanisms remain unclear. One current popular explanation is that nausea and vomiting during pregnancy is fetoprotective, inducing aversions to foods, especially meat, dairy and seafoods, which may carry toxins, pathogens or mutagens. However, most spontaneous abortions arise from genetic or epigenetic defects that are present at or near conception. Moreover, measurements of human chorionic gonadotropin (hCG) at the time of implantation, particularly its hyperglycosylated isoform, accurately predict subsequent spontaneous abortion. Thus the developmental fate of most embryos is fixed before the onset of the symptoms of pregnancy sickness. An alternative explanation for the link between pregnancy sickness and spontaneous abortion is the embryo quality hypothesis: high quality embryos are both more likely to produce the biochemical antecedents of pregnancy sickness and avoid spontaneous abortion. Recent work has shown that the link between pregnancy sickness and spontaneous abortion grows stronger with maternal age, dramatically so in mothers 35 or older. This reflects the parallel rise in the incidence of autosomal aneuploidies with maternal age. The link between pregnancy sickness and spontaneous abortion exists not because nausea and vomiting during pregnancy is fetoprotective, but because nausea and vomiting is an index of a high quality embryo. Pregnancy sickness is not adaptive per se, but the result of an antagonistic pleiotropy over thyroid function, where embryos use hCG to modulate maternal thyroid hormone production during gestation. Embryos benefit from the thyroid hormone production that is key to neurodevelopment, but produce maternal nausea and vomiting as a by-product. Pregnancy sickness, however, may still serve to protect embryo quality but by a different mechanism that posited under the MEPH. Embryo quality is protected by calibrating the dietary intake of a micronutrient – iodine – critical to neuromotor development. For most humans over most of our evolutionary history, iodine has been in short supply, and iodine deficiency is still the most common source of cognitive impairment across the globe. Thus it is of interest that the food aversions most commonly associated with pregnancy sickness, to meat, dairy and seafoods, are also the chief dietary sources of iodine. There is a further intriguing property about iodine: both too little and too much during early pregnancy are damaging to embryo brain development. Given that pregnancy sickness is closely linked to iodine intake and thyroid function (hypothyroidism is associated with lower levels of nausea and vomiting, hyperthyroidism with more), an obvious interpretation emerges. The previously described link between diet and pregnancy sickness – pregnancy sickness is less likely when plants and particularly corn/maize are the sole food staples – arises not because plant food staples are safe, as previously suggested, but because these foods are iodine poor and may, in addition, be goitrogenic. Pregnancy sickness, which reduces the dietary intake of iodine, is clearly maladaptive under conditions of iodine deficiency and hypothyroidism. Conversely, higher levels of pregnancy sickness induced by hyperthyroidism may protect embryos from the inimical effects of excessive dietary iodine during early gestation by reducing the intake of iodine rich foods.     

Innate food aversions and culturally transmitted food taboos in pregnant women in rural southwest India: Separate systems to protect the fetus?

Pregnancy increases women's nutritional requirements, yet causes aversions to nutritious foods. Most societies further restrict pregnant women's diet with food taboos. Pregnancy food aversions are theorized to protect mothers and fetuses from teratogens and pathogens or increase dietary diversity in response to resource scarcity. Tests of these hypotheses have had mixed results, perhaps because many studies are in Westernized populations with reliable access to food and low exposure to pathogens. If pregnancy food aversions are adaptations, however, then they likely evolved in environments with uncertain access to food and high exposure to pathogens. Pregnancy food taboos, on the other hand, have been theorized to limit resource consumption, mark social identity, or also protect mothers and fetuses from dangerous foods. There have been few tests of evolutionary theories of culturally transmitted food taboos.We investigated these and other theories of psychophysiological food aversions and culturally transmitted food taboos among two non-Western populations of pregnant women in Mysore, India, that vary in food insecurity and exposure to infectious disease. The first was a mixed caste rural farming population (N = 72), and the second was the Jenu Kurubas, a resettled population of former hunter-gatherers (N = 30). Women rated their aversions to photos of 31 foods and completed structured interviews that assessed aversions and socially learned avoidances of foods, pathogen exposure, food insecurity, sources of culturally acquired dietary advice, and basic sociodemographic information. Aversions to spicy foods were associated with early trimester and nausea and vomiting, supporting a protective role against plant teratogens. Variation in exposure to pathogens did not explain variation in meat aversions or avoidances, however, raising some doubts about the importance of pathogen avoidance. Aversions to staple foods were common, but were not associated with resource stress, providing mixed support for the role of dietary diversification. Avoided foods outnumbered aversive foods, were believed to be abortifacients or otherwise harmful to the fetus, influenced diet throughout pregnancy, and were largely distinct from aversive foods. These results suggest that aversions target foods with cues of toxicity early in pregnancy, and taboos target suspected abortifacients throughout pregnancy.     

The Acute Phase Protein Ceruloplasmin as a Non-Invasive Marker of Pseudopregnancy, Pregnancy, and Pregnancy Loss in the Giant Panda

After ovulation, non-pregnant female giant pandas experience pseudopregnancy. During pseudopregnancy, non-pregnant females exhibit physiological and behavioral changes similar to pregnancy. Monitoring hormonal patterns that are usually different in pregnant mammals are not effective at determining pregnancy status in many animals that undergo pseudopregnancy, including the giant panda. Therefore, a physiological test to distinguish between pregnancy and pseudopregnancy in pandas has eluded scientists for decades. We examined other potential markers of pregnancy and found that activity of the acute phase protein ceruloplasmin increases in urine of giant pandas in response to pregnancy. Results indicate that in term pregnancies, levels of active urinary ceruloplasmin were elevated the first week of pregnancy and remain elevated until 20–24 days prior to parturition, while no increase was observed during the luteal phase in known pseudopregnancies. Active ceruloplasmin also increased during ultrasound-confirmed lost pregnancies; however, the pattern was different compared to term pregnancies, particularly during the late luteal phase. In four out of the five additional reproductive cycles included in the current study where females were bred but no birth occurred, active ceruloplasmin in urine increased during the luteal phase. Similar to the known lost pregnancies, the temporal pattern of change in urinary ceruloplasmin during the luteal phase deviated from the term pregnancies suggesting that these cycles may have also been lost pregnancies. Among giant pandas in captivity, it has been presumed that there is a high rate of pregnancy loss and our results are the first to provide evidence supporting this notion.     

Hormones and social monitoring: Menstrual cycle shifts in progesterone underlie women's sensitivity to social information

During the luteal phase of the menstrual cycle, women's bodies prepare themselves for possible pregnancy and this preparation includes a dramatic increase in progesterone. This increase in progesterone may underlie a variety of functionally relevant psychological changes designed to help women overcome challenges historically encountered during pregnancy (e.g., warding off social threats and recruiting allies). This paper reports data supporting the hypothesis that increases in progesterone during the luteal phase underlie heightened levels of social monitoring—that is, heightened sensitivity to social cues indicating the presence of social opportunity or threat. Increases in progesterone during the luteal phase were associated with increased accuracy in decoding facial expressions (Study 1) and increased attention to social stimuli (Study 2). Findings suggest that increases in progesterone during the luteal phase may be linked functionally with low-level perceptual attunements that help women effectively navigate their social world.     

Rates of nausea and vomiting in pregnancy and dietary characteristics across populations

Nausea and vomiting in pregnancy (NVP) is a pervasive and debilitating phenomenon in humans. Several adaptive explanations for NVP occurrence have been recently proposed, the two most prominent of which predict associations with nutritional intake or specific dietary components. Here we extend previous cross-cultural analyses by analysing associations between NVP prevalence in 56 studies (21 countries) and quantitative estimates of per capita intake across major dietary categories, measured for the year of study by the Food and Agriculture Organisation (FAO). Rates of nausea and vomiting in pregnancy were correlated with high intake of macronutrients (kilocalories, protein, fat, carbohydrate), as well as sugars, stimulants, meat, milk and eggs, and with low intake of cereals and pulses. Restricting analyses to studies from North America and Europe caused relationships between macronutrient intake and NVP to disappear, suggesting that they might be influenced by non-dietary confounds associated with geographical region of study. However, factor analysis of dietary components revealed one factor significantly associated with NVP rate, which was characterized by low cereal consumption and high intake of sugars, oilcrops, alcohol and meat. The results provide further evidence for an association between diet and NVP prevalence across populations, and support for the idea that NVP serves an adaptive prophylactic function against potentially harmful foodstuffs.     

Women's attractiveness judgments of self-resembling faces change across the menstrual cycle

Two lines of reasoning predict that women's preferences for people exhibiting cues to kinship will be lower in the follicular phase than in the luteal phase of the menstrual cycle. Women may avoid kinship cues during the follicular phase when they are most fertile due to the costs of inbreeding. Alternatively, women may seek kinship cues during the luteal phase as a byproduct of the benefits of associating with kin during pregnancy, which is also characterized by high progesterone. We find that preferences for facial resemblance, a putative kinship cue, follow this predicted pattern and are positively correlated with estimated progesterone levels based on cycle day. Neither estimated estrogen levels nor conception risk predicted preferences for self-resemblance, and the cyclic shift was stronger for preferences for female faces than male faces. These findings lead to the possibility that this cyclic change in preference for self-resemblance may be a byproduct of a hormonal mechanism for increasing affiliative behavior toward kin during pregnancy rather than a mechanism for preventing inbreeding during fertile periods.     

Neural activation in the orbitofrontal cortex in response to male faces increases during the follicular phase

Women's sexual interest changes with hormonal fluctuations across the menstrual cycle. It is unclear how hormones modify women's sexual behavior and desire, but one possibility is that they alter women's positive appraisals of stimuli and thus their sexual interest. Using 3 T fMRI, we measured neural activation in women at two time points in their menstrual cycle (late follicular, luteal) while they evaluated photos of men presented as potential sexual partners. Participants were ten heterosexual women aged 23-28 none of who was using hormonal contraceptives or in a committed relationship. In an event-related design, the women were presented with as series of photos of male faces and asked questions to assess their degree of sexual interest in the men depicted. Results demonstrate an overall effect of menstrual cycle phase on neural activation. During their follicular versus luteal phase, women demonstrated increased activation in the right medial orbitofrontal cortex (OFC), suggesting increased positive appraisal. Activation in the OFC was positively correlated with women's estradiol to progesterone ratios. There were no areas that demonstrated increased activation during the luteal versus follicular phase. The observed increase in activation in the OFC during the follicular phase may reflect a hormonally mediated increase in appetitive motivation and may prime women towards increased sexual interest and behavior around ovulation.     

Prolonged lactational infertility in adolescent rhesus monkeys

The present study examined the effects of first pregnancy and nursing behavior on postpartum infertility in seasonally breeding rhesus monkeys to assess whether prolonged lactational infertility observed in adolescent mothers is due to a particular pattern of nursing or to decrements in body growth rates. After a successful first pregnancy, a significant percentage of the lactating adolescent mothers (57.1%; n = 8) failed to exhibit an ovulation with normal luteal phase during the subsequent breeding season. In contrast, the remaining lactating adolescents (42.9%, n = 6) and all of the adult mothers (100%, n = 6) exhibited ovulations with a normal luteal phase. Age alone was not the critical variable, since all nonlactating adolescents exhibited ovulations with normal luteal phase parameters in the subsequent breeding season. The luteal phase abnormalities exhibited by the subset of lactating adolescent females were characterized by an inadequate luteal phase (ILP) and by significantly lower serum levels of progesterone, estradiol, and bioactive luteinizing hormone. The occurrence of these ILP ovulations was associated with more frequent nursing bouts prior to ovulation and during the subsequent luteal phase. In contrast, nursing patterns for adult females who had ovulations with normal luteal phases were more similar to those of the infertile lactating adolescents exhibiting significantly longer and more frequent nursing bouts, suggesting that fully adult females may be less sensitive to the inhibitory aspects of a suckling stimulus. Differences in luteal phase function among lactating adolescents were not related to differential rates of ponderal or skeletal growth. A still-developing neuroendocrine system may thus render a significant proportion of adolescent females more sensitive to suckling-induced suppression of gonadotropin secretion.     

Influence of progesterone on arterial blood and CSF acid-base balance in women

To study the mechanism of the action of progesterone on pulmonary ventilation during pregnancy, arterial and cerebrospinal fluid (CSF) acid-base parameters were measured in 59 pregnant and 36 nonpregnant women at the periods of follicular phase, luteal phase, early pregnancy, late pregnancy, and puerperium. Marked respiratory alkalosis in both arterial blood and CSF was observed in pregnancy and puerperium. The degree of hypocapnia observed in the luteal phase and during pregnancy was closely related to the progesterone level in arterial blood. In conclusion, it is unlikely that the observed hyperventilation results from stimulation at the central chemosensitive areas or peripheral chemoreceptors.     

Food cravings, mood, and the menstrual cycle

The primary objective of the present investigation was to document positive changes in food cravings, food consumption, and mood changes over the menstrual cycle and to explore the relation between these factors and dieting habits. A total of 32 female undergraduates completed daily self-reports of food cravings, foods eaten, and mood, for 5 or more weeks. Comparisons of these measures in the 10 days preceding (luteal phase) and 10 days following menstruation onset (follicular phase) revealed that both food cravings and amount eaten were greater in the luteal phase than in the follicular phase. Although women also experienced less positive affect in the luteal phase, there was no correlation between mood and either cravings or amount eaten. Women rated as chronic dieters craved less than those rated an nondieters. The findings on mood and eating confirm previous research, whereas those on cravings extend such research by suggesting that carbohydrate cravings may be augmented by the luteal phase in healthy women.     

Food Aversions and Cravings during Pregnancy on Yasawa Island,Fiji

Women often experience novel food aversions and cravings during pregnancy. These appetite changes have been hypothesized to work alongside cultural strategies as adaptive responses to the challenges posed by pregnancy (e.g., maternal immune suppression). Here, we report a study that assessed whether data from an indigenous population in Fiji are consistent with the predictions of this hypothesis. We found that aversions focus predominantly on foods expected to exacerbate the challenges of pregnancy. Cravings focus on foods that provide calories and micronutrients while posing few threats to mothers and fetuses. We also found that women who experience aversions to specific foods are more likely to crave foods that meet nutritional needs similar to those provided by the aversive foods. These findings are in line with the predictions of the hypothesis. This adds further weight to the argument that appetite changes may function in parallel with cultural mechanisms to solve pregnancy challenges.     

The role of the adenylyl cyclase system in the regulation of corpus luteum function in the human and in nonhuman primates

We have reviewed the properties of luteinizing hormone/human chorionic gonadotropic (LH/hCG)-sensitive adenylyl cyclase (AC) of human corpus luteum (CL) and its regulation by several hormones and nonhormonal activators. We have also described the changes in enzyme activity in membrane preparations of human and cynomolgus monkey CL obtained at various stages of the menstrual cycle and pregnancy. The data have been analyzed with respect to the functional status of the luteal tissue and to the species differences among primate CL. In the menstrual cycle, luteal AC responsiveness to LH/hCG was detectable during the midluteal phase, but not during the late luteal phase or in the follicular phase of the following cycle. In addition, nonhormonal stimulation was high in CL obtained during the midluteal and late luteal phases, but declined drastically by the follicular phase of the next cycle. In early pregnancy, the enzyme was unresponsive to LH/hCG stimulation, yet its sensitivity to nonhormonal stimulation was similar, if not identical, to that of midluteal phase CL. Functional activity was also evident at the end of pregnancy. These results demonstrate that expression of AC activity in primate luteal membrane changes significantly with varying hormonal status under physiologic conditions. It is concluded that the AC system in luteal membranes is an effective model to study the mechanisms that regulate function and life span of the human and nonhuman primate CL.     

Luteal function during the periimplantation period and requirement for estrogen for implantation and pregnancy maintenance in the non-human primate

An attempt has been made in this paper to review our present understanding of luteal function during the periimplantation period and in particular hormonal requirement for implantation and maintenance of early pregnancy in the non-human primate.In a fertile cycle thecorpus luteum is apparently rescued from luteolysis by chorionic gonadotropin secreted by the implanted blastocyst, In the bonnet monkey the serum progesterone titers during the luteal phase of a fertile cycle seems higher compared to that of nonmated cycling monkeys. This suggested that thecorpus luteum is receiving some stimulatory signal from the blastocyst even prior to implantation. The recent demonstration that human blastocyst in culture secretes into the medium human chorionic gonadotropin essentially support the above assumption. However, attempts to extend the luteal phase of cycling unmated monkeys with exogenous human chorionic gonadotropin injection has hitherto not met with complete success suggesting that there could be other than chorionic gonadotropin, additional luteal stimulatory factors the unimplanted blastocyst is secreting.Corpus luteum is the principle source of both progesterone and estrogen produced during the periimplantation period and dysruption of luteal function, brought about by either lutectomy or ovariectomy or luteinizing hormone antiserum treatment, followed by progesterone supplementation leads to maintenance of pregnancy. This has lead to questioning the need for estrogen in the maintenance of early pregnancy. Recent work using Zuclomiphene, an antiestrogen during days 5–11 of cycle in rhesus monkeys mated between day 9–14, has however, suggested that estrogen may be required for implantation. Further work is needed to arrive at an unequivocal decision regarding the need for estrogen in maintenance of early pregnancy in the primate.     

Female body odour is a potential cue to ovulation

Human body odours have been reported to influence female mate choice. Women prefer the odours of immunocompatible men and, during their fertile period, judge the body odours of men with symmetrical bodies--which is indicative of genetic quality--as sexy and pleasant. The reproductive success of men largely depends on mating with fertile women, but it is not known whether men can detect a woman's fertile period. We asked women who had regular menstrual cycles and who were not using hormonal contraceptives to wear a T-shirt for three consecutive nights during their late follicular (ovulatory) phase and another T-shirt during the luteal (non-ovulatory) phase of their menstrual cycle. Male raters judged the odours of T-shirts worn during the follicular phase as more pleasant and sexy than odours from T-shirts worn during the luteal phase. The odour differences between the follicular and luteal phases did not dissipate quickly over time as male raters were able to detect and judge follicular phase body odours as more pleasant and sexy than the odours from the luteal phase even after the T-shirts were kept at room temperature for one week. These findings suggest that ovulation may not be concealed and that men could use ovulation-linked odours in their mate selection.     

Evidence for a switch in the site of relaxin production from small theca-derived cells to large luteal cells during early pregnancy in the pig

The presence of immunoreactive relaxin was studied in corpora lutea of sows during the oestrous cycle and early pregnancy by immunohistochemistry and radioimmunoassay using three different anti-relaxin sera. Sections were immunostained using the peroxidase-anti-peroxidase or the immunogold-silver technique. Before Day 14, staining in corpora lutea from non-pregnant and pregnant animals was indistinguishable. With all antisera, no immunostaining was seen on Day 3, but was detected on Days 5-7 in cells from the theca interna. In non-pregnant animals, this immunostaining decreased and by Day 15 only an occasional large cell in the centre of the corpus luteum was stained. No staining was seen by Day 22. The relaxin content of corpora lutea measured by radioimmunoassay remained low throughout the luteal phase. In contrast, the amount of immunoreactive relaxin in corpora lutea rose dramatically (140-fold) between Days 11 and 14 of pregnancy and by Day 14 of pregnancy immunostaining was seen in the majority of large luteal cells. By Day 20 of pregnancy the concentrations of immunoreactive relaxin had further increased. Histochemical staining for alkaline phosphatase suggested that, while the relaxin-immunoreactive cells seen in the early luteal phase may be theca-derived, those during early pregnancy may be derived from the granulosa. The results are compatible with the suggestion that relaxin is produced by theca-derived cells during the early luteal phase and that between Days 11 and 14 there is a switch in the site of relaxin synthesis from theca-derived cells to granulosa-derived large luteal cells. In the absence of luteolysis, as during pregnancy, this switch is accompanied by a dramatic increase in relaxin synthesis.     

Pathogen disgust,but not moral disgust,changes across the menstrual cycle

The Compensatory Prophylaxis Hypothesis (CPH) proposes that during periods of increased susceptibility to infections, e.g., during the luteal phase of the menstrual cycle when progesterone suppresses immune function, women should feel more disgust toward pathogen cues and behave prophylactically. We investigate differences in disgust sensitivity and contamination sensitivity during different phases of the menstrual cycle in regularly cycling, healthy 93 rural and urban Polish women using the within-subject design. Disgust sensitivity was measured during two different phases of a menstrual cycle: 1) the follicular phase (the 5th or 6th day of the cycle) and 2) the luteal phase (on the 5th day after a positive ovulatory test or on 20th day of a cycle if the result of the ovulatory test was not positive). In the luteal phase, women scored higher on the Pathogen Disgust of the Three-Domain Disgust Scale, the Contamination Obsessions and Washing Compulsions Subscale of Padua Inventory, and on ratings of photographs showing sources of potential infections than in the follicular phase. Moral Disgust of the Three-Domain Disgust Scale did not differ between cycle phases. Hence, results suggest that women feel more disgusted toward cues to pathogens during the luteal phase, when susceptibility to infection is greater. We suggest that it is necessary to incorporate ovulatory testing as well as to conduct repeated measurements of disgust sensitivity in future tests of the CPH. Moreover, we believe that understanding how the feeling of pathogen disgust varies across the menstrual cycle and in relation to progesterone levels could be useful in designing effective infectious diseases prevention strategies for women.     

Dimorphic foraging behaviors and the evolution of hominid hunting.

In contemporary foraging societies men typically hunt more than women. This observation has played an important role in many reconstructions of hominid evolution. The gender difference in human hunting, likely a product of both ecological and cultural factors, is mirrored by a similar sex difference among nonhuman primates. Existing explanations of such primate behavioral dimorphism are augmented by the recognition of an additional factor that may contribute to differences between males and females in the value of meat. Episodic female immunosuppression is a normal part of reproduction. Because meat is a source of pathogens, females can be expected to exhibit less constant attraction to meat. Sexual dimorphism in the attraction to meat may then contribute to dimorphic foraging specializations, a divergence that is likely augmented by the differential value of insectivory across the sexes. With the rise of cultural transmission of foraging knowledge, dimorphic foraging behaviors would have been reinforced, creating a more comprehensive gender-based division of labor.     

Expression of prostaglandin synthesis pathway enzymes in the porcine corpus luteum during the oestrous cycle and early pregnancy

Prostaglandins (PGs) of luteal origin may have paracrine and/or autocrine actions on the functions of the corpus luteum (CL). Previously, we have shown that enzymes of PG synthesis pathway such as prostaglandin E synthase (mPGES-1), prostaglandin F synthase (PGFS) and prostaglandin 9-ketoreductase (CBR1) are important in regulation of PG production in the conceptuses and endometrium of cyclic and pregnant pigs. Therefore, localization and expression patterns of these enzymes were determinated in porcine CL. The PGFS protein content was lower in metestrus and higher around luteolysis, and then decreased in late regressing CL. PGFS protein levels were lower on days 5-8 of pregnancy and did not differ between days 10 and 25. Elevated expression of mPGES-1 mRNA was found in early luteal phase. The mPGES-1 protein content, similarly to PGFS, was higher during luteolysis. mPGES-1 mRNA and protein levels were constant between days 5 and 25 of pregnancy. PGFS and mPGES-1 expression was down-regulated on days 16-17 of the oestrous cycle when compared to the corresponding days of pregnancy. Enhanced mPGES-1/PGFS ratio occurred during early luteal phase and days 5-8 of pregnancy. Expression of CBR1 mRNA and protein was constant during the cycle and pregnancy. Our studies revealed higher mPGES-1/PGFS ratios in the CL during early luteal phase and corresponding days of pregnancy that could favor PGE(2) synthesis and may be important in the control of luteal development. However, PG synthesis in the endometrium/conceptus rather than in the CL could be involved in luteolysis and maternal recognition of pregnancy in pigs.