The embryonic development of the temnocephalid flatworms Craspedella pedum and Diceratocephala boschmai

We have analyzed the embryonic development of the temnocephalid flatworms Craspedella pedum and Diceratocephala boschmai, using a combination of fuchsin-labeled whole-mount preparation, histology, and transmission electron microscopy. Following the staging system recently introduced for another flatworm species (Mesostoma lingua), we can distinguish eight morphologically defined stages. Temnocephalids produce eggs of the neoophoran type in which a small oocyte is surrounded by a layer of yolk cells. Cleavage takes place in the center of the yolk mass (stages 1-2) and results in an irregular, multilayered disc of mesenchymal cells that moves to the future ventral egg pole (stage 3). Organ primordia, including those of the brain, pharynx, male genital apparatus, sucker, and epidermis "crystallize" within this disc without undergoing gastrulation movements (stage 4). An invagination of the epidermal primordium pushes the embryo back into the center of the yolk ("embryonic invagination"). As a result, organogenesis begins while the embryo is invaginated (stage 5). The brain differentiates into an outer cortex of cell bodies that surround a central neuropile. Precursor cells of the epidermis, pharynx, and protonephridia become organized into epithelia. During stage 6, the embryonic primordium everts back to the surface, where organogenesis and cell differentiation continues. Epidermal cells fuse into a syncytium that expands around the yolk. Myoblasts initially do not spread out in the way epidermal cells do; they remain concentrated in two narrow, longitudinal bands that extend along the sides of the embryo. Three pairs of axon tracts extending posteriorly from the brain follow the bands of myoblasts. Stages 7 and 8 are characterized by the appearance of eye pigmentation, brain condensation, and the formation of tentacles and a sucker that bud out from the epidermis of the anterior and posterior end, respectively. Comparison of morphogenesis in temnocephalids with observations in other flatworm taxa suggests a phylotypic stage for this phylum of invertebrates.     

The embryonic development of the flatworm <Emphasis Type="Italic">Macrostomum</Emphasis> sp.

Macrostomid flatworms represent a group of basal bilaterians with primitive developmental and morphological characteristics. The species Macrostomum sp., raised under laboratory conditions, has a short generation time of about 2–3 weeks and produces a large number of eggs year round. Using live observation, histology, electron microscopy and immunohistochemistry we have carried out a developmental analysis of Macrostomum sp. Cleavage (stages 1–2) of this species follows a modified spiral pattern and results in a solid embryonic primordium surrounded by an external yolk layer. During stage 3, cells at the anterior and lateral periphery of the embryo evolve into the somatic primordium which gives rise to the body wall and nervous system. Cells in the center form the large yolk-rich gut primordium. During stage 4, the brain primordium and the pharynx primordium appear as symmetric densities anterior-ventrally within the somatic primordium. Organ differentiation commences during stage 5 when the neurons of the brain primordium extend axons that form a central neuropile, and the outer cell layer of the somatic primordium turns into a ciliated epidermal epithelium. Cilia also appear in the lumen of the pharynx primordium, in the protonephridial system and, slightly later, in the lumen of the gut. Ultrastructurally, these differentiating cells show the hallmarks of platyhelminth epithelia, with a pronounced apical assembly of microfilaments (terminal web) inserting at the zonula adherens, and a wide band of septate junctions underneath the zonula. Terminal web and zonula adherens are particularly well observed in the epidermis. During stage 6, the somatic primordium extends around the surface dorsally and ventrally to form a complete body wall. Muscle precursors extend myofilaments that are organized into a highly regular orthogonal network of circular, diagonal and longitudinal fibers. Neurons of the brain primordium differentiate a commissural neuropile that extends a single pair of ventro-lateral nerve trunks (the main longitudinal cords) posteriorly. The primordial pharynx lumen fuses with the ventral epidermis anteriorly and the gut posteriorly, thereby generating a continuous digestive tract. The embryo adopts its final shape during stages 7 and 8, characterized by the morphallactic lengthening of the body into a U-shaped form and the condensation of the nervous system.Edited by J. Campos-Ortega     

The embryonic development of the rhabdocoel flatworm Mesostoma lingua (Abildgaard, 1789)

The embryonic development of the flatworm Mesostoma lingua was studied using a combination of life observation and histological analysis of wholemount preparations and sections (viewed by both light and electron microscopy.) We introduce a series of stages defined by easily recognizable morphological criteria. These stages are also applicable to other platyhelminth taxa that are currently under investigation in our laboratory. During cleavage (stages 1 and 2), the embryo is located in the center of the egg, surrounded by a layer of yolk cells. After cleavage, the embryo forms a solid, disc-shaped cell cluster. During stage 3, the embryo migrates to the periphery of the egg and acquires bilateral symmetry. The side where it contacts the egg surface corresponds to the future ventral surface of the embryo. Stage 4 is the emergence of the first organ primordia, the brain and pharynx. Gastrulation, as usually defined by the appearance of germ layers, does not exist in Mesos-toma; instead, organ primordia emerge ”in situ” from a mesenchymal mass of cells. Organogenesis takes place during stages 5 and 6. Cells at the ventral surface form the epidermal epithelium; inner cells differentiate into neurons, somatic and pharyngeal muscle cells, as well as the pharyngeal and protonephridial (excretory) epithelium. A junctional complex, consisting initially of small septate junctions, followed later by a more apically located zonula adherens, is formed in all epithelial tissues at stage 6. Beginning towards the end of stage 6 and continuing throughout stages 7 and 8, cytodifferentiation of the different organ systems takes place. Stage 7 is characterized by the appearance of eye pigmentation, brain condensation and spindle-shaped myocytes. Stage 8 describes the fully dorsally closed and differentiated embryo. Muscular contraction moves the body in the egg shell. We discuss Mesostoma embryogenesis in comparison to other animal phyla. Particular attention is given to the apparent absence of gastrulation and the formation of the epithelial junctional complex. Received: 10 February 2000 / Accepted: 10 April 2000     

The embryonic development of the triclad <Emphasis Type="Italic">Schmidtea polychroa</Emphasis>

Triclad flatworms are well studied for their regenerative properties, yet little is known about their embryonic development. We here describe the embryonic development of the triclad Schmidtea polychroa, using histological and immunocytochemical analysis of whole-mount preparations and sections. During early cleavage (stage 1), yolk cells fuse and enclose the zygote into a syncytium. The zygote divides into blastomeres that dissociate and migrate into the syncytium. During stage 2, a subset of blastomeres differentiate into a transient embryonic epidermis that surrounds the yolk syncytium, and an embryonic pharynx. Other blastomeres divide as a scattered population of cells in the syncytium. During stage 3, the embryonic pharynx imbibes external yolk cells and a gastric cavity is formed in the center of the syncytium. The syncytial yolk and the blastomeres contained within it are compressed into a thin peripheral rind. From a location close to the embryonic pharynx, which defines the posterior pole, bilaterally symmetric ventral nerve cord pioneers extend forward. Stage 4 is characterized by massive proliferation of embryonic cells. Large yolk-filled cells lining the syncytium form the gastrodermis. During stage 5 the external syncytial yolk mantle is resorbed and the embryonic cells contained within differentiate into an irregular scaffold of muscle and nerve cells. Epidermal cells differentiate and replace the transient embryonic epidermis. Through stages 6–8, the embryo adopts its worm-like shape, and loosely scattered populations of differentiating cells consolidate into structurally defined organs. Our analysis reveals a picture of S. polychroa embryogenesis that resembles the morphogenetic events underlying regeneration.Edited by D. Tautz     

Freshwater Rhabdocoela (Platyhelminthes) from ephemeral rock pools from Botswana, with the description of four new species and one new genus

Four new species of freshwater rhabdocoel flatworms from ephemeral rock pools in southeastern Botswana are described and discussed. Two of them, Syringoplana kolasai n. gen. n. sp. and Mesostoma thamagai n. sp. belong to the Typhloplanidae Graff, 1905. The unique construction of the excretory system is the main characteristic of S. kolasai. M. thamagai can be separated from other Mesostoma Ehrenberg, 1837 species by the presence of a bundle of eosinophilic glands at the transition from oviduct to seminal receptacle. The other two taxa, Gieysztoria isoldeae n. sp. and G. faubeli n. sp. belong to the Dalyelliidae Graff, 1905. G. isoldeae is characterised by the presence of four separate hollow spines in the male atrium, which are connected to two accessory glandular organs. G. faubeli can be separated from other Gieysztoria Ruebush and Hayes, 1939 species by the detailed construction of the stylet. Apart from these two species the occurrence of an unidentified Microdalyellia Gieysztor, 1938 species is mentioned.     

The embryonic development of Schistosoma mansoni eggs: proposal for a new staging system

Schistosomiasis is a water-borne parasitic illness caused by neoophoran trematodes of the genus Schistosoma. Using classical histological techniques and whole-mount preparations, the present work describes the embryonic development of Schistosoma mansoni eggs in the murine host and compares it with eggs maintained under in vitro conditions. Two pre-embryonic stages occur inside the female worm: the prezygotic stage is characterized by the release of mature oocytes from the female ovary until its fertilization. The zygotic stage encompasses the migration of the zygote through the ootype, where the eggshell is formed, to the uterus. Fully formed eggs are laid still undeveloped, without having suffered any cleavage. In the outside environment, eight embryonic stages can be defined: stage 1 refers to early cleavages and the beginning of yolk fusion. Stage 2 represents late cleavage, with the formation of a stereoblastula and the onset of outer envelope differentiation. Stage 3 is defined by the elongation of the embryonic primordium and the onset of inner envelope formation. At stage 4, the first organ primordia arise. During stages 5 to 7, tissue and organ differentiation occurs (neural mass, epidermis, terebratorium, musculature, and miracidial glands). Stage 7 is characterized by the nuclear condensation of neurons of the central neural mass. Stage 8 refers to the fully formed larva, presenting muscular contraction, cilia, and flame-cell beating. This staging system was compared to a previous classification and could underlie further studies on egg histoproteomics (morphological localizome). The differentiation of embryonic structures and their probable roles in granulomatogenesis are discussed herein. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The embryonic development of the bodywall and nervous system of the cestode flatworm Hymenolepis diminuta

Cestodes (tapeworms) are a derived, parasitic clade of the phylum Platyhelminthes (flatworms). The cestode body wall represents an adaptation to its endoparasitic lifestyle. The epidermis forms a non-ciliated syncytium, and both muscular and nervous system are reduced. Morphological differences between cestodes and free-living flatworms become apparent already during early embryogenesis. Cestodes have a complex life cycle that begins with an infectious larva, called the oncosphere. In regard to cell number, cestode oncospheres are among the simplest multicellular organisms, containing in the order of 50-100 cells. As part of our continuing effort to analyze embryonic development in flatworms, we describe here the staining pattern obtained with acTub in embryos and larvae of the cestode Hymenolepis diminuta and, briefly, the monogenean Neoheterocotyle rhinobatidis. In addition, we labeled the embryonic musculature of Hymenolepis with phalloidin. In Hymenolepis embryos, two different cell types that we interpret as neurons and epidermal gland cells express acTub. There exist only two neurons that develop close to the midline at the anterior pole of the embryo. The axons of these two neurons project posteriorly into the center of the oncosphere, where they innervate the complex of muscles that is attached to the hooklets. In addition to neurons, acTub labels a small and invariant set of epidermal gland cells that develop at superficial positions, anteriorly adjacent to the neurons, in the dorsal midline, and around the posteriorly located hooklets. During late stages of embryogenesis they spread and form a complete covering of the embryo. We discuss these data in the broader context of platyhelminth embryology.     

Embryogenesis in the reef-building coral Acropora spp

Embryogenesis in the reef building corals Acropora intermedia, A. solitaryensis, A. hyacinthus, A. digitifera, and A. tenuis was studied in detail at the morphological level, and the relationships among the animal pole, blastopore, and mouth were investigated for the first time in corals. These species showed essentially the same sequence of development. The embryo undergoes spiral-like holoblastic cleavage despite the presence of a dense isolecithal yolk. After the morula stage, the embryo enters the "prawn-chip" stage, which consists of an irregularly shaped cellular bilayer. The embryo begins to roll inward to form the bowl stage; the round shape observed during this stage suggests that it may be the beginning of gastrulation. However, the blastopore closes and the stomodeum (mouth and pharynx) is formed via invagination at a site near the closed blastopore. During the planula stage, a concavity forms in the aboral region in conjunction with numerous spirocysts, suggesting that spirocysts are used to attach to the substrate before the onset of metamorphosis.     

Key steps in the morphogenesis of a cranial placode in an invertebrate chordate, the tunicate Ciona savignyi

Tunicates and vertebrates share a common ancestor that possessed cranial neurogenic placodes, thickenings in embryonic head epidermis giving rise to sensory structures. Though orthology assignments between vertebrate and tunicate placodes are not entirely resolved, vertebrate otic placodes and tunicate atrial siphon primordia are thought to be homologous based on morphology and position, gene expression, and a common signaling requirement during induction. Here, we probe key points in the morphogenesis of the tunicate atrial siphon. We show that the siphon primordium arises within a non-dividing field of lateral-dorsal epidermis. The initial steps of atrial primordium invagination are similar to otic placode invagination, but a placode-derived vesicle is never observed as for the otic vesicle of vertebrates. Rather, confocal imaging reveals an atrial opening through juvenile stages and beyond. We inject a photoactivatable lineage tracer to show that the early atrial siphon of the metamorphic juvenile, including its aperture and lining, derives from cells of the atrial placode itself. Finally, we perturb the routing of the gut to the left atrium by laser ablation and pharmacology to show that this adaptation to a sessile lifestyle depends on left-right patterning mechanisms present in the free-swimming chordate ancestor.     

半滑舌鳎胚胎发育中后期卵黄囊超微结构观察

在胚胎发育中期,半滑舌鳎胚胎由胚体、卵黄囊和卵周液构成.对半滑舌鳎胚胎发育中后期的卵黄囊进行超微结构观察.结果表明,卵黄囊是由卵黄囊膜和包裹其内的卵黄物质组成.在半滑舌鳎胚胎发育过程中,卵黄囊内的卵黄物质逐渐消耗产生低分子量的卵黄磷蛋白分裂小泡.分裂小泡转移到卵黄囊内部消黄细胞中,在消黄细胞的作用下分裂小泡转化成卵黄颗粒.随后卵黄颗粒在卵黄囊内表面聚集成囊状结构并转移运输到卵黄囊膜内部,最后把卵黄物质从卵黄囊膜转移并释放到卵周液中,为胚胎发育提供营养.     

Temporal dissociation of developmental events in the chick eye under low temperature conditions

The chick embryonic eye is an excellent model for the study of vertebrate organogenesis. Key events in eye development involve thickening, invagination and cytodifferentiation of the lens primordium. While these events occur successively at different developmental stages, the extent to which these events are temporally related is largely unknown. Here we show that the lens invagination is highly sensitive to temperature. Lowering of incubation temperature to 29°C at embryonic day 2 delayed the onset of invagination of the lens, but not thickening and cytodifferentiation, leading to abnormal protrusion of the eye. The temperature shift also delayed the inward bending of the underlying retinal primordium, even in the absence of the lens. Taken together, our results suggest that lens invagination is initiated independently of thickening and cytodifferentiation, possibly by mechanisms associated with morphogenesis of the primordial retina.     

尖唇散白蚁胚胎发育激光共聚焦扫描显微镜观察

【目的】本研究旨在探究尖唇散白蚁Reticulitermes aculabialis胚胎在不同发育阶段的变化特征。【方法】每日收集尖唇散白蚁的卵,并固定其胚胎发育状态,采用DAPI染剂对白蚁胚胎进行染色,通过激光共聚焦扫描显微镜观察记录尖唇散白蚁胚胎在不同发育阶段的形态特征。【结果】在25℃下尖唇散白蚁胚胎发育过程历经25~30 d,按照发育特征将其划分为12个阶段。胚胎发育早期,卵黄细胞均匀分布在卵内部,卵内细胞核向卵的中间浓缩,在细胞到达卵的后表面时形成浓缩的囊胚细胞作为胚盘;胚胎发育中期,胚胎开始进行“反转型”的囊胚运动,头部和前后轴从后极到前极反转,胚带出现明显的“双弯”结构。胚胎发育中后期,胚胎变宽,内部器官逐渐开始发育,出现明显的伸长与分节;胚胎发育后期,附肢发育明显,内部器官发育成熟。【结论】尖唇散白蚁胚胎发育过程历经12个阶段,属于短胚带型,胚带出现“双弯”结构,发育中期经历两次囊胚反转。本研究为真社会性昆虫白蚁的胚胎发育过程提供了形态学和生物学依据。     

Early development and axis specification in the sea anemone Nematostella vectensis

We investigated the early development of the sea anemone Nematostella vectensis, an emerging model system of the Cnidaria. Early cleavage stages are characterized by substantial variability from embryo to embryo, yet invariably lead to the formation of a coeloblastula. The coeloblastula undergoes a series of unusual broad invaginations-evaginations which can be blocked by cell cycle inhibitors suggesting a causal link of the invagination cycles to the synchronized cell divisions. Blastula invagination cycles stop as cell divisions become asynchronous. Marking experiments show a clear correspondence of the animal-vegetal axis of the egg to the oral-aboral axis of the embryo. The animal pole gives rise to the concave side of the blastula and later to the blastopore of the gastrula, and hence the oral pole of the future polyp. Asymmetric distribution of granules in the unfertilized egg suggest an animal-vegetal asymmetry in the egg in addition to the localized position of the pronucleus. To determine whether this asymmetry reflects asymmetrically distributed determinants along the animal-vegetal axis, we carried out blastomere isolations and embryonic divisions at various stages. Our data strongly indicate that normal primary polyps develop only if cellular material from the animal hemisphere is included, whereas the vegetal hemisphere alone is incapable to differentiate an oral pole. Molecular marker analysis suggests that also the correct patterning of the aboral pole depends on signals from the oral half. This suggests that in Nematostella embryos the animal hemisphere contains organizing activity to form a normal polyp.     

The embryonic development of the polyclad flatworm Imogine mcgrathi

In this paper we describe the embryonic development of the polyclad flatworm Imogine mcgrathi. Imogine is an indirect developer that hatches as a planctonic Goette’s larva after an embryonic period of approximately 7 days. Light and electron microscopic analyses of sections of staged embryos were combined with antibody stainings of wholemounted embryos to reconstruct the origin and movement of the primordia of the various organ systems, with particular emphasis on the nervous system. We introduce a system of morphologically defined stages aimed at facilitating future studies and cross-species comparisons among flatworm embryos. Imogine embryos undergo typical spiral cleavage. Micromere quartets 1–3 form an irregular double layer of mesenchymal cells that during gastrulation expands over micromere quartet 4. Micromere 4d divides into several large mesendodermal precursors whose position defines the ventral pole of the embryo. These cells, along with the animal micromeres that obtained a sub-surface position during cleavage, form a deep layer of cells that gives rise to all internal structures, including the nervous system, musculature, nephridia, and gut. Micromeres 4a–c are large yolky cells that are incorporated into the lumen of the gut, but do not themselves contribute to the gut epithelium. Shortly after gastrulation, cell differentiation sets in. Cells located at the surface adopt epithelial characteristics and form cilia that result in continuous movement of the post-gastrula stage embryo. Deep cells at the lateral margins of the embryo become organized into a protonephridial tube. A cluster of approximately 50 deep cells at the anterior pole forms the brain, in which we have identified sets of founder neurons of the brain commissure and the dorsal and ventral connectives. The early differentiating neurons, along with other cells forming stabilized microtubules (ciliated cells of the epidermis, gut and protonephridia; apical gland cells) could be analyzed in detail because of their labeling with an antibody against acetylated α-tubulin. Our findings indicate that, despite significant differences in the cleavage pattern and arrangement of blastomeres in the early embryo, morphogenesis and organ formation of a polyclad embryo follows a pattern that is very similar to the pattern observed by us and others in phylogenetically more evolved rhabdocoel flatworms. Received: 10 February 2000 / Accepted: 10 April 2000     

Embryonic development of the jumping bristletail Pedetonutus unimaculatus Machida,with special reference to embryonic membranes (Hexapoda: Microcoryphia,Machilidae)

In the machilid Pedetonutus unimaculatus, a germ disc is formed by the aggregation and proliferation of cells within a broadly defined embryonic area. Cells adjacent to the embryonic area form the serosal fold that grows beneath the embryo. Then the embryonic margin is extended to form a cell layer or amnion that lies between the embryo and serosal fold. Thus, an amnioserosal fold is formed by the addition of the amnion to the serosal fold. Serosal cells cover the entire surface of the egg and begin to secrete a serosal cuticle. Soon the amnioserosal fold is withdrawn, and the embryo is exposed to the egg surface. The spreading amnion replaces the serosal cells that finally degenerate through the formation of a secondary dorsal organ. In the areas of amnion anterior and lateral to the embryo, yolk folds form and encompass the embryo. The amnion is a provisional dorsal closure and never participates in the formation of the definitive one. The amnioserosal fold of the Microcoryphia appears to have the functional role of secreting a serosal cuticle beneath the embryo. This fold of the Microcoryphia may be regarded as an ancestral form of the amnioserosal folds of the Thysanura-Pterygota. the yolk folds may appear to be passive transformation of the yolk mass linked to positioning of the growing embryo within the egg. There is no evidence that the yolk folds and the cavity appearing between them in the Microcoryphia are homologous to the amnioserosal fold and amniotic cavity in the Thysanura-Pterygota. The yolk folds appear to be one of the embryological autapomorphies in the Microcoryphia. © 1994 Wiley-Liss, Inc.     

Light and scanning microscopic studies of integument differentiation in the grass snake Natrix natrix L. (Lepidosauria,Serpentes) during embryogenesis

We analysed the differentiation of body cover in the grass snake (Natrix natrix L.) over the full length of the embryo's body at each developmental stage. Based on investigations using both light and scanning electron microscopes, we divided the embryonic development of the grass snake integument into four phases. The shape of the epidermal cells changes first on the caudal and ventral parts of the embryo, then gradually towards the rostral and dorsal areas. In stage V on the ventral side of the embryo the gastrosteges are formed from single primordia, but on the dorsal side the epidermis forms the scale primordia in stage VII. This indicates that scalation begins on the ventral body surface, and spreads dorsally. The appearance of melanocytes between the cells of the stratum germinativum in stage VII coincides with changes in embryo colouration. The first dermal melanocytes were detected in stage XI so in this stage the definitive skin pattern is formed. In the same stage the epidermis forms the first embryonic shedding complex and the periderm layer begins to detach in small, individual flakes. This process coincides with rapid growth of the embryos.     

Asymmetric distribution of Echinoid defines the epidermal leading edge during Drosophila dorsal closure

During Drosophila melanogaster dorsal closure, lateral sheets of embryonic epidermis assemble an actomyosin cable at their leading edge and migrate dorsally over the amnioserosa, converging at the dorsal midline. We show that disappearance of the homophilic cell adhesion molecule Echinoid (Ed) from the amnioserosa just before dorsal closure eliminates homophilic interactions with the adjacent dorsal-most epidermal (DME) cells, which comprise the leading edge. The resulting planar polarized distribution of Ed in the DME cells is essential for the localized accumulation of actin regulators and for actomyosin cable formation at the leading edge and for the polarized localization of the scaffolding protein Bazooka/PAR-3. DME cells with uniform Ed fail to assemble a cable and protrude dorsally, suggesting that the cable restricts dorsal migration. The planar polarized distribution of Ed in the DME cells thus provides a spatial cue that polarizes the DME cell actin cytoskeleton, defining the epidermal leading edge and establishing its contractile properties.     

The follicular placenta of the viviparous fish,Heterandria formosa. I. Ultrastructure and development of the embryonic absorptive surface

Embryos of the poeciliid Heterandria formosa develop to term in the ovarian follicle in which they establish a placental association with the follicle wall (follicular placenta) and undergo a 3,900% increase in embryonic dry weight. This study does not confirm the belief that the embryonic component of the follicular placenta is formed only by the surfaces of the pericardial and yolk sacs; early in development the entire embryonic surface functions in absorption. The pericardial sac expands to form a hood-like structure that covers the head of the embryo and together with the yolk sac is extensively vascularized by a portal plexus derived from the vitelline circulation. The hood-like pericardial sac is considered to be a pericardial amnion-serosa. Scanning and transmission electron microscopy reveal that during the early and middle phases of development (Tavolga's stages 10–18 for Xiphophorus maculatus) the entire embryo is covered by a bilaminar epithelium whose apical surface is characterized by numerous, elongate microvilli and coated pits and vesicles. Electron-lucent vesicles in the apical cytoplasm appear to be endosomes while a heterogeneous group of dense-staining vesicles display many features characteristic of lysosomes. As in the larvae of other teleosts, cells resembling chloride cells are also present in the surface epithelium. Endothelial cells of the portal plexus lie directly beneath the surface epithelium of the pericardial and yolk sacs and possess numerous transcytotic vesicles. The microvillous surface epithelium becomes restricted to the pericardial and yolk sacs late in development when elsewhere on the embryo the non-absorptive epidermis differentiates. We postulate that before the definitive epidermis differentiates, the entire embryonic surface constitutes the embryonic component of the follicular placenta. The absorptive surface epithelium appears to be the principle embryonic adaptation for maternal-embryonic nutrient uptake in H. formosa, suggesting that a change in the normal differentiation of the surface epithelium was of primary importance to the acquisition of matrotrophy in this species. In other species of viviparous poeciliid fishes in which there is little or no transfer of maternal nutrients, the embryonic surface epithelium is of the non-absorptive type.