To elucidate the morphology of the initial stages of epitheliochorial placentation in the pig, material from 10 sows of the Danish Landrace and from one Göttinger minipig gilt from day 13 to day 26 of gestation was processed for scanning and transmission electron microscopy. The observed foetomaternal interaction from day 19 1/2 minipig placenta corresponded well to the observations on the Danish Landrace placenta. From the results and the discussion it was concluded that the following structures were implicated in the initial phases of placentation in the pig: Protruding epithelial proliferations of the uterine epithelium enclosed by chorionic caps serving to immobilize the blastocyst (days 13 and 14). A thick glycocalyx on the maternal and a thin one on the foetal epithelium before contact. Close apposition between the apical plasma membranes from trophoblastic and uterine epithelium (day 14). Development of interdigitating microvilli (days 15-16). Formation of apical domes on the uterine epithelium closely related to the trophoblast provided with long cytoplasmic extensions into a luminal space between the apical domes, apparently representing a transition from histiotropic to haemotrophic nutrition (days 15-20). Placentation, development of interdigitating microvilli between foetal and maternal epithelium, was extended but not terminated in the peripheral zone at day 26.
The microvascular architecture of the pig placenta was studied by serial semithin histological sections for light microscopy, which were compared with scanning electron microscopy of artificially exposed materno-fetal contact surfaces as well as of vessel casts prepared from the maternal, fetal, and combined maternal and fetal sides. The superficial reliefs from the exposed surfaces as well as from the casts are almost identical with the complementary maternal and fetal sides. In order to meet the physiological needs of materno-fetal exchange for the rapidly growing fetuses, these reliefs develop from a simple to a more complex system during pregnancy and can be described as follows: (1) The degree of interlocking increases between the fetal ridges or bulbous protrusions and maternal ridges of different orders separated by maternal troughs of variable depth, most clearly seen on vessel casts. It creates a three-dimensional notch-arrangement, giving strength to the materno-fetal contact area. (2) The structure of precapillary vessels as well as of the meshwork, and the diameter of capillaries of the maternal and fetal sides, adapt during gestation giving a good distribution of oxygenated blood into the maternal capillaries; these, with the development of large prevenous connecting capillaries on the fetal side, favour a high arterio-venous difference of fetal blood O2 pressure. (3) The vascular architecture of endometrial and fetal ridges and troughs develop into a crosscurrent to countercurrent materno-fetal blood interrelationship. Our demonstration of the materno-fetal capillary inter-relationship in the porcine placenta thus shows that the latter is a much more efficient organ for exchange than hitherto assumed.
The arrangement of the enteric ganglia and nerve fibre plexuses was examined in the submucous and mucous layers and around Peyer's patches of the porcine small intestine to clarify their organisation. Immunohistochemistry of vasoactive intestinal peptide (VIP) and neurofilament proteins in wholemounts, chopped or paraffin sections was used to locate the neural elements. The ganglia of the internal and external submucous plexuses were situated at 2 different topographic locations, being clearly demarcated by the submucosal vascular arcades and differing in neuronal composition. The internal submucous plexus was the only contributor to the plexus surrounding the follicles of Peyer's patches as a continuous mesh of 3 ganglionated nerve subplexuses. VIP-immunoreactive fibres from this mesh innervated the dome. The mucosal plexus, which was subdivided into 4 subunits-the outer proprial, inner proprial, pericryptal and villous plexuses-contained a few solitary neuronal perikarya. Labelling for neurofilament proteins revealed Dogiel types II, IV and VI neurons. The observations reveal several new features in the enteric nervous system of the pig and clarify its nomenclature.
The architecture of the fetal villous tree and its vasculature in the bovine placentome were studied in the second half of gestation using both conventional histology and histology of ink-filled blood vessels. These were compared with corrosion casts of plastic fillings of the vasculature, prepared for scanning electron microscopy. This combination of morphological methods allows perception of the villous tree throughout gestation from broad-conical to tall-conical form where branch ramification occurs mainly at right angles to the stem. The stem villus typically contains a single central artery and several peripheral veins arranged in parallel. The proximal branches to the stem, the intermediate villi, contain a central arteriole and accompanying venules. The distal branches, the terminal villi, enclose capillary convolutions which consist of an afferent arterial capillary limb, capillary loops and efferent venous capillary limbs. Vascular interconnections exist within the terminal villi, as capillaries or venules between the capillary convolutions, serially bridging them in up to 5 places, and as capillary anastomoses between the capillary loops. Coiling and sinusoidal dilatations of these loops develop near the end of gestation. The intraplacentomal rearrangement of villous trees with progressive gestation and their morphological vascular adaptations are discussed in relation to placental function, including the ever increasing need for transplacental substance exchange. This adaptation allows the blood to traverse the shortest possible arterioarteriolar route to the periphery of the trees where exchange takes place. The need for an increasing blood flow stimulates capillary growth and at the same time optimises the blood flow reaching the placental barrier represented by the vessel cast surface. The capillaries also carry the blood back into the very voluminous system of venules and veins where back diffusion may occur. The total volume of terminal villi of bovine placentome, the ' working part ' of villous trees, hence distinctly increases with respect to the stem and intermediate villi, the ' supplying part ' of the villous tree. In morphological terms the efficiency of the bovine transplacental diffusional exchange is higher than in the closely related ' co-ruminants ' sheep and goats and distinctly higher when compared with the human placenta.
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