Anatomy, physiology and pathology of the amniotic and allantoic compartments in the sheep and cow

In both cows and sheep the allantoic fluid is confined to 2 sacs connected by a flattened isthmus, one in the non-pregnant horn and one in the upper part of the pregnant horn. The chorion encloses the amniotic and allantoic compartments, forming the amniochorion and chorioallantois, respectively. In the last third of gestation the compositions of both amniotic and allantoic fluids differ substantially from each other and from those of foetal urine, and maternal and foetal plasma. There is less variation in composition than in volume for a given gestational age. Abnormalities of volume are more common in cows than sheep, and hydrallantois is more common than hydramnois. Data obtained from both physiological experiments and pathological cases suggest that the foetal membranes play an important role in the regulation of composition and volume of foetal fluids. Evidence is presented that the permeability of the membranes to various solutes, as well as their capacity to produce and respond to a number of hormones, can affect the foetal fluid composition and/or volume. Progesterone, oestrogens and prolactin are some of the hormones known to affect foetal fluids. Foetal adrenal insufficiency has been associated with hydramnios implying a lack of hormones from this gland in this disease. The changes in allantoic fluid composition from normal to that closely resembling maternal/foetal extracellular fluid, in hydrallantois, suggests an alteration of membrane function as an aetiology and the continued production of fluid, after removal of the foetus in some cases, favours this hypothesis.     

Amniotic and allantoic fluids from experimentally infected sheep contain immunoglobulin specific for Chlamydophila abortus

Chlamydophila abortus, the aetiological agent of enzootic abortion of ewes (EAE), replicates in trophoblast cells leading to their destruction and dissemination of the bacterium to foetal organs. To further understand the pathogenesis of EAE, amniotic and allantoic fluids were collected from experimentally infected pregnant ewes at 30 (7 samples from each fluid), 35 (8 samples from each fluid), 40 (10 samples from each fluid) and 43 (6 amniotic fluids and 7 allantoic fluids) days post-infection to determine pathogen numbers and other markers of infection. Whilst experimentally infected ewes had characteristic placental lesions, only two amniotic and seven allantoic fluid samples were positive for C. abortus by real-time PCR. In contrast, all amniotic and allantoic fluids were positive for immunoglobulin. Immunoglobulins were generally detected earlier in allantoic fluid than in amniotic fluid and the numbers of samples containing immunoglobulins increased as infection progressed. IgG in amniotic and allantoic fluids was shown to be specific for C. abortus, and reacted with the major outer membrane proteins, polymorphic outer membrane protein and macrophage infectivity potentiator protein. A comparison of two-dimensional immunoblots using purified IgG from the allantoic fluid, amniotic fluid, ewe serum and foetal serum of a C. abortus infected animal at 40 days post infection indicated a pattern of reactivity intermediate between that of the ewe serum and the foetal serum. Results suggest that a maternal source of immunoglobulin is predominant at 30 days post-infection but that foetal derived antibodies may be contributed at a later stage.     

Cytology of bovine foetal fluids.

Cell contents of foetal fluids obtained separately from amniotic and allantoic cavities of live cows by transvaginal amniocentesis and by aspiration from pregnant uteri immediately after slaughter were studied using Papanicolaou's staining procedure. The samples obtained belonged to cows which were between ten to 24 weeks of gestation. Cell scrapings from foetal and membranous surfaces normally in contact with the foetal fluids were also stained simultaneously with foetal fluid smears by Papanicolaou's method to determine the probable origin of the cells detected in the amniotic and allantoic cavities. The cell types were classified according to morphology and tinctorial patterns and their origin was deduced on the basis of similarity to cell scrapings from the surface layers of foetus and foetal cavities. The evaluation of foetal fluid cells revealed that a majority of cells are of foetal origin and that the cytological features of these cells may be useful in assessing the condition of the foetus.     

Detection of Toxoplasma gondii antigens reactive with antibodies from serum, amniotic, and allantoic fluids from experimentally infected pregnant ewes

Toxoplasma gondii, an intracellular protozoan parasite, is one of the major causes of infectious abortion in sheep. To further understand the pathogenesis of toxoplasmosis, serum, amniotic and allantoic fluids and foetal stomach contents were collected from experimentally infected pregnant ewes to determine pathogen numbers and other markers of infection. Fifteen pregnant ewes (90 days of gestation) were each orally inoculated with 3000 sporulated oocysts of T. gondii. Serum samples were collected weekly following challenge. Amniotic and allantoic fluids and foetal stomach contents were collected at 21, 25, 28, 33 and 35 days post-infection. Characteristic placental lesions were detected in 1 of 4 challenged ewes at day 25, 3 of 4 challenged ewes at day 28 and in all challenged ewes at days 33 and 35 post-infection. T. gondii was detected only sporadically in amniotic and allantoic fluids before 35 days of infection, by real-time PCR, and only in ewes with placental lesions. At 35 days post-infection, high numbers of parasite were detected in both amniotic and allantoic fluids. An increase in the number of fluids from challenged animals with IgM and IgG was detected over time, except for IgG in allantoic fluid, which was detected in all samples from day 21 post-infection. IgG in amniotic and allantoic fluids was shown to be specific for T. gondii, and reacted with antigens with an apparent molecular mass of approximately 22 kDa and 30 kDa. Results suggest a maternal source of immunoglobulin in the allantoic fluid and a foetal source of immunoglobulin in the amniotic fluid early in infection but that both sources may contribute immunoglobulin to both fluids at a later stage.     

Biochemical Profile of Amniotic and Allantoic Fluid During Different Gestational Phases in Mares

Fetal fluids have different vital functions that sustain both pregnancy and normal parturition. The biochemical composition of amniotic fluid during gestation is not well established; thus the purpose of the present study was to determine the biochemical profile of both amniotic and allantoic fluids from mares during initial, mid, and latter third phases of pregnancy. Samples were collected after slaughter, using allantocentesis and amniocentesis. Sixty samples of fetal fluids were analyzed. Alkaline phosphatase (AP), glucose, total protein (TP), urea, creatinine, Ca, chloride (Cl), Na, and K concentrations were measured using commercially available kits. The AP concentration in amniotic fluid was higher than that in allantoic fluid during the three gestational phases (P < .05). There were no differences between glucose mean values of allantoic and those of amniotic fluids (P < .05). However, glucose values were higher in the allantoic fluid in the last trimester of pregnancy. TP was higher in the amniotic fluid than in allantoic fluid (P < .05). Urea values varied among the phases; however, there were no differences between the amniotic and allantoic fluid values (P > .05). Creatinine values were higher in allantoic fluid (P < .05). Na and Cl concentrations were higher in amniotic fluid (P < .05). However, Ca and K concentrations were higher in the allantoic fluid.     

The protein composition of sheep foetal fluids

Amniotic fluid contains several proteins including alpha-foetoprotein derived from the foetal plasma, but lacks the higher molecular weight alpha2- and gamma-globulins. The protein composition of the fluid, like that of the serum, remains unchanged after the first month of gestation, and is buffered from the influence of the allantoic fluid by the amnion. Allantoic fluid contains a significantly higher concentration and a greater variety of proteins; it is not a simple transudate of either foetal or maternal plasma, and though containing proteins derived from both sources, it is deficient in albumin but relatively abundant in gamma-globulins. The latter comprises gamma-foetoprotein and mature type IgG. It is believed that maternal IgG may reach the allantoic sac in appreciable amount, but its transmission to the amniotic sac and thence to the foetus may be prevented by the impermeability of the amnion.     

Metabolomic Profile of Allantoic and Amniotic Fluid in Late-term Gestational Mares Characterized by 1H-nuclear Magnetic Resonance Spectroscopy

The amniotic and allantoic fluid compartments in the mare serve essential roles throughout pregnancy and parturition. Although the global metabolomic profile of amniotic fluid in women has been extensively characterized, current data for equine fetal fluids are limited. Therefore, the goal of this study was to characterize the global metabolomic profile of equine allantoic and amniotic fluid through nuclear magnetic resonance spectroscopy. Fetal fluids were collected between 270 and 295 days of gestation from 12 pregnancies through ultrasound-guided transabdominal puncture. A total of 24 samples (n = 10 allantoic fluid; n = 9 amniotic fluid; n = 5 admixed fluid) were analyzed by one-dimensional proton (¹H) and two-dimensional (¹H-¹³ C) nuclear magnetic resonance spectroscopy. Metabolites were integrated and compared between fluid types using a Kruskal–Wallis test at P < .05 significance. A total of 28 distinct metabolites were found in allantoic and admixed fluid, whereas 23 metabolites were identified in amniotic fluid. Allantoic fluid contained significant elevations (P < .05) in the metabolites betaine, creatine, creatinine, citrate, histidine, nitrophenol, tryptophan, π-methylhistidine, and unknown metabolite #1 compared with amniotic fluid, whereas amniotic fluid contained statistically increased concentrations of the metabolite lactate compared with allantoic fluid (P = .003).     

Proteomic analysis of amniotic and allantoic fluid from buffaloes during foetal development

The objective of this study was to describe the dynamic changes in protein composition and protein abundance in amniotic and allantoic fluids from buffaloes during gestation. Amniotic and allantoic fluids were collected during the first, second and third trimesters of gestation. The foetuses were measured and weighed. Fluid samples were centrifuged at 800 g for 10 min and then at 10,000 g for 60 min at 4°C. The supernatant was collected to determine the total protein concentration. Based on total protein concentration, an aliquot (50 μg) was used for in‐solution tryptic digestion, and mass spectrometry analysis (nano‐LC‐MS/MS) was performed. A multivariate statistical analysis of the proteomic data was conducted. Across the different stages of buffalo gestation, fifty‐one proteins were found in the amniotic fluid, and twenty‐one were found in the allantoic fluid. A total of twelve proteins were common among the stages, and four presented significant differences (VIP score α > 1). Fibronectin and alpha‐1‐antiproteinase were more abundant in the amniotic fluid than in the allantoic fluid. Alpha‐2‐macroglobulin and alpha‐2‐HS‐glycoprotein were more abundant in the allantoic fluid than in the amniotic fluid. Alpha‐2‐macroglobulin participates in remodelling and growth of the uterus at beginning of the gestation (first trimester), and these findings indicate that can serve as a potential tool for the early diagnosis of pregnancy in buffaloes.     

Immunoglobulin concentrations and bovine enterovirus inhibitors in fetal bovine fluids.

Fluids from 53 bovine fetuses ranging in age from 90 to 240 days were examined for immunoglobulin G (IgG) and immunoglobulin M (IgM) and neutralizing activity to ten bovine viruses. Non-specific inhibitors to bovine enteroviruses were found in serum, allantoic, and amniotic fluids of most samples tested. In most cases, serum IgG were within normal values. Neither IgG nor IgM was detected in amniotic fluids, whereas 2 samples of allantoic fluid contained traces of IgG.     

鸡胚发育过程中胚胎液中几种游离氨基酸的动态变化

本实验旨在研究鸡胚发育过程中尿囊液和羊水中天冬氨酸、谷氨酸、精氨酸和亮氨酸浓度的动态变化。选用72枚6胚龄的三黄鸡鸡胚,按重量相近原则分为6组,每组12个重复,分别在孵化第7、8、11、13、14、18天采集尿囊液和羊水,并利用高效液相色谱法分析其氨基酸浓度。结果表明:尿囊液中,除8胚龄外,其他各胚龄精氨酸浓度均高于另外3种氨基酸(P<0.05),13胚龄精氨酸浓度显著低于其他胚龄(P<0.05),而18胚龄显则著高于其他胚龄(P<0.05)。羊水中,各胚龄精氨酸浓度均高于其他3种氨基酸;除11胚龄外,13胚龄精氨酸浓度显著高于其他胚龄(P<0.05);7胚龄天门冬氨酸浓度显著高于13、14、18胚龄(P<0.05);但18胚龄谷氨酸及亮氨酸浓度则显著升高(P<0.05)。以上结果说明,在尿囊液及羊水中,所测4种氨基酸浓度变化不尽相同,但总体精氨酸含量最丰富,这可能与鸡胚发育的营养与代谢有关。     

Electrolytes and reproductive hormone concentrations in maternal plasma and fetal fluids of dairy cows with hydrops

Electrolytes, metabolites, cortisol and reproductive hormones were measured in maternal plasma taken at least twice daily from three cases of bovine hydrops before, during and after parturition induced by dexamethasone or prostaglandin. Caesarean operations were required for two of the cases. Maternal plasma electrolytes remained within the normal range, but average potassium and creatinine concentrations were higher (9.2 and 0.68 mmol/litre, respectively) than normal (4.7 and 0.42 mmol/litre) in samples of amniotic fluid obtained at calving. Sodium (100 mmol/litre) and chloride (67 mmol/litre) in allantoic fluid were also higher than normal (53 and 20 mmol/litre, respectively). Conversely, creatinine concentrations were lower than normal in allantoic fluid (2.2 vs 13.8 mmol/litre). Oestradiol concentrations were lower than normal in maternal plasma (ranges: less than 20 to 140 pg/ml vs 30 to 440 pg/ml); maximum prostaglandin F metabolite (PGFM) concentrations were slightly elevated (ranges 1.1 to 2.0 ng/ml vs 0.4 to 0.9 ng/ml). Progesterone and cortisol concentrations remained within the normal range; the latter hormone increased markedly in parallel with raised PGFM concentrations. In two cases, the concentrations of reproductive hormones tended to be lower in the amniotic fluid than in the allantoic fluid. For example, progesterone concentrations were 42.8 and 14.9 ng/ml in the amniotic fluids vs 64.2 and 29.8 ng/ml in the allantoic fluids of the two cows; PGFM concentrations were 27.7 and 4.3 ng/ml vs 34.6 and 5.0 ng/ml, and oestradiol concentrations were 1.5 and 3.5 ng/ml vs 1.1 and 6.4 ng/ml in the two fluids, respectively.     

Experimental Infection of Cows with Bovine Viral Diarrhoea Virus in Early Pregnancy – Findings in Serum and Foetal Fluids

Nineteen pregnant cows were experimentally infected with bovine viral diarrhoea virus (BVDV) between day 74 and 81 of pregnancy. All cows became infected and developed serum antibodies. Sixteen of the cows delivered persistently infected (PI) offspring, whereas the remaining three gave birth to calves with detectable serum antibodies and free from BVDV. The 16 cows with PI foetuses developed higher levels of antibodies in serum during pregnancy than did their three peers carrying non‐PI calves. Multivariate analysis showed that the antibody levels in these two groups of cows were significantly different from day 135 of pregnancy. Foetal fluid was successfully collected from 18 of the 19 infected cows and from five uninfected control cows between 10 and 24 days before delivery by use of a percutaneous, blind puncture technique. No negative effects were observed in the cows or their offspring. BVDV was isolated and detected with an immunoperoxidase test in foetal fluid from 13 of the 16 cows carrying PI foetuses, and from 15 of the cows when a quantitative fluorescent polymerase chain reaction (PCR) technique was used. The negative sample in the PCR assay was positive for BVDV antibodies. The number of viral copies per microlitre in foetal fluids varied between 103 and 1080 in the positive samples. All samples taken from the cows carrying non‐PI foetuses were negative for BVDV in both assays. In this experiment, examination of either serum or foetal fluids could identify the cows carrying a PI foetus. Examination of serum for BVDV antibodies was a reliable indicator of a PI foetus if the serum was collected during the last 2 months of pregnancy. For examination of foetal fluids, both viral and serological analyses should be performed. For viral analysis, PCR should be the test of choice. High levels of BVDV antibodies in conjunction with a negative result in the PCR may be indicative of a false‐negative virus result. Further experience with the method of collection of foetal fluids is necessary for evaluation of its safety. Investigation of pregnant cows in order to discover a PI offspring before it is born could be a useful tool in control and eradication of BVDV.     

Experimental infection of cows with bovine viral diarrhoea virus in early pregnancy - findings in serum and foetal fluids

Nineteen pregnant cows were experimentally infected with bovine viral diarrhoea virus (BVDV) between day 74 and 81 of pregnancy. All cows became infected and developed serum antibodies. Sixteen of the cows delivered persistently infected (PI) offspring, whereas the remaining three gave birth to calves with detectable serum antibodies and free from BVDV. The 16 cows with PI foetuses developed higher levels of antibodies in serum during pregnancy than did their three peers carrying non-PI calves. Multivariate analysis showed that the antibody levels in these two groups of cows were significantly different from day 135 of pregnancy. Foetal fluid was successfully collected from 18 of the 19 infected cows and from five uninfected control cows between 10 and 24 days before delivery by use of a percutaneous, blind puncture technique. No negative effects were observed in the cows or their offspring. BVDV was isolated and detected with an immunoperoxidase test in foetal fluid from 13 of the 16 cows carrying PI foetuses, and from 15 of the cows when a quantitative fluorescent polymerase chain reaction (PCR) technique was used. The negative sample in the PCR assay was positive for BVDV antibodies. The number of viral copies per microlitre in foetal fluids varied between 103 and 1080 in the positive samples. All samples taken from the cows carrying non-PI foetuses were negative for BVDV in both assays. In this experiment, examination of either serum or foetal fluids could identify the cows carrying a PI foetus. Examination of serum for BVDV antibodies was a reliable indicator of a PI foetus if the serum was collected during the last 2 months of pregnancy. For examination of foetal fluids, both viral and serological analyses should be performed. For viral analysis, PCR should be the test of choice. High levels of BVDV antibodies in conjunction with a negative result in the PCR may be indicative of a false-negative virus result. Further experience with the method of collection of foetal fluids is necessary for evaluation of its safety. Investigation of pregnant cows in order to discover a PI offspring before it is born could be a useful tool in control and eradication of BVDV.     

Clinical evaluation of ketone bodies in body fluids of cows in late pregancy and their fetuses

Ketone body concentrations were determined in six body fluids of cows in late pregnancy and their fetuses (blood plasma and urine of cows, amniotic and allantoic fetal fluids, and fetal blood serum and urine) using the head-space gas chromatographic method. Physiological ratios 0/T% of oxidized (acetone and acetoacetic acid) to total (acetone, isopropanol, acetoacetic and 3-hydroxybutyric acids) ketone bodies were below 10% in cow blood plasma and urine, fetal serum and urine, and allantoic fluid, and below 20% in amniotic fluid. A scheme for the evaluation of the degree and duration of ketosis based on the 0/T% ratio is proposed. The existence of a renal regulatory mechanism for excretion of ketone bodies is suggested by analogy with the renal acidobasic regulation.     

Clinical evaluation of ketone bodies in body fluids of cows in late pregnancy and their fetuses

Ketone body concentrations were determined in six body fluids of cows in late pregnancy and their fetuses (blood plasma and urine of cows, amniotic and allantoic fetal fluids, and fetal blood serum and urine) using the head-space gas chromatographic method. Physiological ratios O/T% of oxidized (acetone and acetoacetic acid) to total (acetone, isopropanol, acetoacetic and 3-hydroxybutyric acid) ketone bodies were below 10% in cow blood plasma and urine, fetal serum and urine, and allantoic fluid, and below 20% in amniotic fluid. A scheme for the evaluation of the degree and duration of ketosis based on the O/T% ratio is proposed. The existence of a renal regulatory mechanism for excretion of ketone bodies is suggested by analogy with the renal acidobasic regulation.     

Predicting the energy and protein requirements of the pregnant grasscutter (Thryonomys swinderianus, Temminck) using the changes in weight and composition of the foetus and associated tissues of pregnancy

Eighteen (18) gravid uteri collected from hunters’ kills during February and March 2008 were used to investigate growth and accretion of energy and protein in the gravid uterus during pregnancy in the grasscutter. Each gravid uterus was separated into foetal, placental, empty uterus and foetal fluids (amniotic and allantoic fluids combined) components. Individual tissues were weighed and analyzed for dry matter (DM), ash, crude protein (CP) and crude fat. Regression equations were fitted to the weights of each component to describe the effects of gestational age. The DM, protein and energy contents of individual uterine tissues, with the exception of the DM of the foetal fluids, increased significantly with advancing gestation age so that the DM of the gravid uterus increased ~20-fold from days 45 to 152 (3.85 v. 76.24 g). However, the average elemental N in the foetus was 8.3 g?N/100 g. Percentage DM did not change with advancing gestation age. Fat content of the foetal grasscutter was 2.79 %. Changes in foetal protein and energy fitted a multi-phasic regression that consisted of two linear equations. Similarly, changes in foetal fat and water contents fitted a multi-phasic regression that consisted of two linear equations, indicating that protein and energy growth accelerated after days 95 and 97 of gestation, respectively, with increases in fat and water accelerating after days 127 and 96 of gestation, respectively. Foetal protein and energy accretions were 0.04 g?day^?1 and 0.001 MJ?day^?1 before days 95 and 97 of gestation, respectively, and increased to 0.31 g?day^?1 and 0.009 MJ?day^?1 after days 95 and 97 of gestation, respectively. Fat and water accretions were also 0.012 g?day^?1 and 0.15 ml?day^?1 before days 127 and 96 of gestation, respectively, and increased to 0.104 g?day^?1 and 1.29 ml?day^?1 after the respective days of gestation. Protein needs for tissue protein gain increased ~8-fold after day 95 of gestation. Results of this study indicate that the growth of the foetus and foetal tissues occur at different rates during gestation and points to a two-phase feeding strategy before and after day 95 of gestation for pregnant grasscutters.     

Experimental Neospora Caninum Infection in Pregnant Dairy Heifers Raises Concentrations of Pregnancy‐Associated Glycoproteins 1 and 2 in Foetal Fluids

Plasma concentrations of PAG‐1 are used for pregnancy diagnosis and as a marker of placental/foetal well‐being, while those of PAG‐2 may be an indicator of abortion risk in Neospora caninum‐infected cows. Studies have shown that N. caninum infection modifies PAG‐1 and PAG‐2 patterns in maternal blood plasma. However, no prior work has examined the effects of N. caninum infection on concentrations of PAGs in foetal fluids. In this study, PAG‐1, PAG‐2 and pH levels were determined in the amniotic and allantoic fluids of foetuses collected at 152 days of gestation from control uninfected dams and from dams experimentally infected with N. caninum on Day 110 of gestation. Foetal fluids from infected foetuses had significantly higher PAG‐2 concentrations (p = 0.026) and pH values (p = 0.02) than fluids from non‐infected foetuses. In infected foetuses, significantly higher concentrations of PAG‐1 (p < 0.001) and PAG‐2 (p < 0.001) were detected in fluid samples showing antibodies against N. caninum than those without antibodies. Moreover, pH values were significantly higher (p = 0.011) in foetal fluid samples with antibodies than in samples from non‐infected foetuses. In conclusion, this is the first report on the effect of N. caninum infection on PAG levels in foetal fluids. Our results indicate that following the experimental infection of dams with N. caninum on Day 110 of gestation, foetal fluids collected from the infected foetuses of these dams featured higher PAG‐1 and PAG‐2 levels and pH values than fluids from non‐infected controls, provided that the samples tested showed the presence of antibodies. The clinical implications of these findings are that following infection with N. caninum, most cows will experience some level of placental damage and that this injury correlates with foetal fluid PAG levels and pH.     

EQUINE HERPESVIRUSES: Type 3 as an Abortigenic Agent

The inoculation of equine herpesvirus type 3 (EHV3) strain 65/61 into the amniotic cavity of a mare 6-7 months pregnant resulted in abortion 11 days later. Following abortion typical lesions of coital exanthema were not observed in the genital tract of the mare, nor was EHV3 isolated from her. Serological evidence, however, indicated that the mare was infected with EHV3 following inoculation. Grossly the foetal disease was characterised by placentitis, focal ulcerative dermatitis, focal necrosis of the lungs and a striking diptheritic gastritis. Histological findings were interstitial pneumonia, diffuse hepatitis, generalised myositis, extensive vascular necrosis and degeneration of a range of epithelial cells. EHV3 was isolated from the placenta and placental fluids, stomach fluid, pooled thoracic and abdominal fluid, skin, lung, spleen and small intestine of the foetus.     

Placental transfer and renal clearance of sulphamethoxazole and its metabolite N4-acetylsulphamethoxazole in a pregnant ewe

Sulphamethoxazole and its metabolite, N₄-acetylsulphamethoxazole, were shown to cross the placenta of a pregnant ewe. The plasma concentration of N₄-acetylsulphamethoxazole increased in the foetus due to the limited elimination properties of the immature kidney. The amniotic fluid concentration of sulphamethoxazole and N₄-acetylsulphamethoxazole were low, amounting to 2.0 and 0.46μg/ml respectively. Protein binding of the sulphonamides was slightly higher in the foetus than in the ewe.     

胚胎发育过程中羊水和尿囊液量及pH变化的研究

通过孵化试验，研究了孵化过程中羊水及尿囊液量及pH的变化。结果表明，羊水量在16胚龄达到最大，pH一直维持在7.0左右。尿囊液量在12胚龄达到最大，pH一直偏碱性，其中12胚龄的pH最大。羊水和尿囊液的pH具有一定的互补性。