Ciliary Rootlets Research Articles

Fine structure of dicyemid mesozoans, with special reference to cell junctions.

The fine structure of the dicyemid mesozoan, Dicyema acuticephalum, from Octopus vulgaris, was studied with special attention to intercellular junctional complexes between various kinds of cells. Two types of intercellular junction, namely, adherens junctions and gap junctions, were found in both vermiform stages and in infusoriform embryos. Adherens junctions were classified into two types. Zonulae adherentes-like junctions were observed between adjacent peripheral cells at vermiform stages, between adjacent external cells of infusoriform embryos, and between members of groups of internal cells that covered the urn in infusoriform embryos. Maculae adherentes-like junctions were seen between a peripheral cell and an axial cell at vermiform stages. In infusoriform embryos, these junctions were observed between various types of cells, excluding urn cells. Gap junctions were found between adjacent peripheral cells at vermiform stages, whereas in infusoriform embryos these junctions were located between various types of cells excluding urn cells. Dicyemids might be the most primitive multicellular animals to possess these basic types of cell junctions. Ciliary rootlet systems at vermiform stages and in infusoriform embryos were unique in structure compared with those of other primitive multicellular animals. J Morphol 231:297-305, 1997. © 1997 Wiley-Liss, Inc.

The surface pattern of Paramecium tetraurelia in interphase: an electron microscopic study of basal body variability, connections with associated ribbons and their epiplasmic environment

Summary In Paramecium tetraurelia, thousands of basal bodies with their three associated rootlets are anchored into the submembranous skeleton, the epiplasm, which is itself segmented into cortical units. In this study, we redescribe more precisely (after Pitelka [31], Hufnagel [19] and Allen [1]) the architecture of these structures after detergent treatment and tannic acid contrast reinforcement. Although this technique leads to loss of membranes and of some labile cytoskeletal elements, it allows to better underline how the three rootlets are linked together and to the basal body. The postciliary microtubules and ciliary rootlets (Kd fibers) are bound together by a “fingered node”, and the transverse microtubules to the previous fibers by dense links. The anterior basal body of the pairs is linked to the ciliary rootlet by a newly identified architecture in this species, the “bone-like node” [1]. A specific ciliary domain can be defined in the epiplasm of each cortical unit: epiplasmic rings encircling the basal bodies are closely apposed on the thicker part of the epiplasm and can include some apertures along the basal bodies towards the cilia. The thickness of the epiplasm seems to vary according to the presumptive elongation direction of the cortical units. Each basal body associated rootlet is specifically linked to the epiplasm itself. Because of their unexpected set of linkage organization, these rootlets may ensure the ciliary cohesiveness in the interphase cell as well as the transmission of the ciliary polarity during division as it was previously discussed [19]. We also show for the first time that there is a variability of the basal body architecture at the cellular level: its height, and the number of cartwheel elements change depending on its location on the cortex and its morphogenetical history; the longest basal bodies with 5 cartwheel elements are found in territories which result from hyperduplication and become invariant, i.e. do not undergo duplication during the next divisions, as is the case along the sutures and in the left oral and cortical areas [22]. These results are discussed in terms of morphogenetical behavior.

Studies on Stenostomum grande Child, 1902 (Platyhelminthes, Catenulida): fine structure of the digestive tract and the endocytotic activity of the gastrodermis

Abstract The digestive tract and its endocytotic activity in the catenulid Stenostomum grande were studied by electron microscopy. The pharynx was typical of the simplex type. At the mouth, between the integumental epithelium and the pharyngeal epithelium proper, was a transition zone. Among the epithelial cells of this transition were monociliated sensory cells and the necks of bucco‐pharyngeal secretory cells of two types. The pharyngeal epithelium proper was densely ciliated, with long ciliary rootlets and mitochondria. It was surrounded by two layers of muscles. The gastrodermis consisted of phagocytes and typical secretory Minotian cells. It was underlain by a delicate basal lamina and muscle fibers. Distinctive of the phagocytes was the presence of differentiated cilia, cup‐shaped mitochondria, and vacuoles with dense inclusions. Morphological differences between pharyngeal and gastrodermal cilia suggest functional differences. Experiments using latex beads as tracers and the identification of acid phosphatase in cytoplasmic vacuoles pointed to a high level of endocytotic and digestive activity in the phagocytes. Our data demonstrate that the basic structure of the digestive tract in S. grande conforms well to that of other free‐living platyhelminths, but it does have ultrastructural peculiarities.

Are the most numerous sensilla of terrestrial isopods hygroreceptors? Ultrastructure of the dorsal tricorn sensilla of Porcellio scaber

The ultrastructure of the tricorn sensilla of the woodlouse Porcellio scaber was investigated in cryofixed and freeze-substituted, or chemically fixed specimens. The tricorn sensilla have a foramenized triangular-shaped outer hair and bear a poreless rod-like inner hair. The conical base of the inner hair is connected to the base of the outer hair by a complex cuticular structure. Each sensillum contains three sensory cells. The tip of one of the three dendrites contains a tubular body and is clamped between two bulges of the dendritic sheath. The two other dendrites protrude to the tip of the inner hair, flush against the cuticular wall. The microtubules in the ciliary segments are arranged in nine double tubuli that have neither osmiophilic cores nor arms. The ciliary rootlets are small. The inner segment of the largest dendrite wraps around the two smaller dendrites and one of seven enveloping cells in a mesaxon-like manner. Although this ultrastructure deviates considerably from most crustacean mechanosensitive sensilla, it nevertheless suggests a mechanosensitive function, at least for one of the sensory cells. In many aspects, the tricorn sensilla resemble the thermo-hygrosensilla of insects. However, our results suggest that the structural criteria for thermo-hygro-sensitivity used in insects cannot simply be applied to crustaceans.

Ultrastructure of spermiogenesis and spermatozoa inProrhynchussp. (Platyhelminthes, Lecithoepitheliata, Prorhynchidae)

Summary Mature sperm of Prorhynchus sp. have an elongated nucleus, multiple mitochondria and dense bodies, and two free axonemes which are located in grooves of the main shaft for much of their length. The axonemes are subterminally inserted and have the typical 9+ ‘1’ arrangement unique to Platyhelminthes and synapomorphic for taxa of Trepaxonemata. The testis follicles examined had small numbers of developing spermatids and very few mature sperm were present. During spermiogenesis, spermatids remain joined in clusters by distinctive bridges. In each spermatid two centrioles (with an intercentriolar body between them) give rise to free axonemes which grow out in opposite directions from each other. Indistinct ciliary rootlets are present. The axonemes are carried distally from the main spermatid mass on an elongating process and turn back towards the main spermatid mass. Nucleus, mitochondria and dense bodies move into the shaft, and the spermatid elongates before detaching from others in the cluster. Th...

Cortical development associated with conjugation of Paramecium

ABSTRACT The cortical cytoskeleton of Paramecium is character-ized by a complex, polarized and asymmetrical organi-zation. In order to analyse the cortical development of Paramecium tetraurelia during the sexual process of con-jugation, different antibodies have been used to follow the development of five cortical components: basal bodies, ciliary rootlets, outer lattice, epiplasm and cytospindle. This study demonstrates that the cortex of Paramecium undergoes an elaborate development process following conjugation. Some of the cortical structures that are not renewed at any other moment of the cell cycle, are resorbed after conjugation and replaced by other newly formed ones. The reorganiza-tion of the cortical components occurs according to dif-ferent morphogenetic waves that spread over the cell surface. The oral system and the preoral suture line act as morphogenetic epicentres.

Ultrastructure of the protonephridial system of regenerating Stenostomum sp. (Plathelminthes, Catenulida)

The ultrastructure of the flame bulbs, protonephridial capillaries and duct of fully developed and regenerating Stenostomum sp. is described. Flame bulbs are formed by a single cell whose nucleus is located basally or laterally to the weir. The weir is formed by a single row of transverse ribs connected by a thin ‘membrane’, apparently of extracellular matrix. Internal leptotriches arise from the proximal cytoplasm and extend in a (usually) single row along the weir and into the lumen of the distal cytoplasmic tube. Many or all leptotriches do not fuse with the distal cytoplasm. Two cilia are each anchored in the proximal cytoplasm by a cross-striated vertical and lateral rootlet, the latter bent forward and extending for some distance into one of the two cytoplasmic cords along the weir. Each cord contains the lateral rootlet in its proximal part, as well as many microtubules. The distal cytoplasmic tube contains two (longitudinal) junctions, i.e. lines of contact between cell processes of the same, terminal cell. Occasionally, more than two junctions were seen, apparently due to branches of the terminal cell in contact with each other. Flame bulbs join capillaries lined by several canal cells type I, containing few or no microvilli but lateral flames. Such capillaries join a duct (or ducts?) lined by canal cells type II with many long microvilli. The large protonephridial duct is lined by numerous cells with lateral flames and many long microvilli. In regenerating tissue (10.5 hours after cutting) some flame bulbs were ‘free’, i.e. not connected to capillaries, and some capillaries openly communicated with the surrounding intercellular space. In the presence of a single row of ribs in the weir, of internal leptotriches, and of vertical and lateral ciliary rootlets, the flame bulb of Stenostomum sp. resembles that of other Plathelminthes much more closely than hitherto thought. The species differs from non-catenulid plathelminths mainly in the large number of glandular cells lining the large protonephridial ducts, in the transverse orientation of the ribs in the weir and in the presence of only two cilia in the flame.

Ultrastructure of the Epidermis and Protonephridium of an Undescribed Species of Luridae (Platyhelminthes, Rhabdocoela)

The epidermis of an undescribed species of Luridae possesses many large cavities filled with a medium-dense material, intraepidermal nuclei, glandular ducts, cilia with vertical and horizontal rootlets, surface microvilli, and a thick basal lamina. Cilia have narrow tips, the peripheral axonemal doublets lose one of the microtubules and, finally, whole doublets are lost near the tip. Flame bulbs have a single row of longitudinal ribs containing microtubules and connected by a 'membrane' apparently of extracellular matrix; cilia of flame bulbs possess cross-striated rootlets, and capillary walls are smooth. The nucleus of the protonephridium was observed near the tip of the flame bulb, along the capillary. It is not clear whether one perikaryon forms more than one flame bulb. The structure of the epidermal ciliary rootlets, the presence of intraepidermal nuclei, and flame bulbs composed of a single row of longitudinal ribs containing microtubules formed by a single cell are typical rhabdocoel characteristics.

Ultrastructure of Sensory Receptors of an Undescribed Species of Luridae (Platyhelminthes, Rhabdocoela)

Nine types of uniciliate sensory receptors are described from an undescribed species of Luridae: a collar receptor with a collar formed by 7-8 rods; a receptor with a long vertical and short horizontal rootlet and a thick ring-like electron-dense collar; a receptor with an approximately 2-mum-long cilium that has a narrow bent tip; a bulbous receptor with a short bulbous cilium; an intraepidermal receptor with a short cilium enclosed in an epidermal cavity; a receptor with a long cilium, short ciliary rootlet and electron-dense rod; pharyngeal receptors with long (at least 4 mum long) and short (0.5 mum long) cilia; and a pharyngeal receptor with a cilium of intermediate length. Centrioles in nerve cells and (in cross-section) butterfly-shaped cilia may also be of a sensory nature. Ecological implications of the large variety of sensory receptors are discussed: in the variety of stimuli to which animals have to respond for survival, interstitial beach habitats are comparable to habitats on or in hosts used by platyhelminth parasites (which have a comparable number of receptor types).

Ovarian ultrastructure and vitellogenesis in ten species of shallow-water and bathyal sea cucumbers (Echinodermata: Holothuroidea)

Comparative ultrastructural features of the ovary and vitellogenesis have been described for six shallow water and four bathyal species of sea cucumbers representing four major holothuroid orders. Ovarian structure is similar in all ten species except for features of the peritoneal cells of the outer layer and the follicular inner epithelial cells surrounding the developing oocytes. The peritoneal cells vary from monociliated squamous or cuboidal cells to large columnar cells. Ultrastructural evidence suggests that these cells might be capable of incorporating materials from the perivisceral coelom. The follicular inner epithelial cells of two deep-sea species resemble podocytes, a feature previously unre-ported in holothuroid ovaries. It is suggested that these cells function to increase nutrient exchange between the genital haemal sinus and the oocyte during vitellogenesis. In all ten species, the oocytes appear to participate in yolk synthesis through the interaction of the Golgi complex and rough endoplasmic reticulum. The similarity in the ultrastructural features of vitellogenesis suggests that the process of yolk synthesis has been highly conserved in holothuroids. Endocytotic activity was detected in seven of ten species but it is uncertain if this is directly related to vitellogenesis. Cilia and intracellular structures resembling striated ciliary rootlets were observed in the oocytes of four of the ten species studied. The significance of this finding is unclear but could indicate that germ cells have a somatic cell origin.

Ultrastructure of the Flame Bulbs and Protonephridial Capillaries of Artioposthia sp. (Platyhelminthes, Tricladida, Geoplanidae)

AbstractThe protonephridial terminal complex of Artioposthia is formed by one or two terminal cells, each with a nucleus located in the lateral wall of the flame bulb, and probably two proximal canal cells forming the wall of the protonephridial capillary. The weir is restricted to the proximal parts of the flame bulbs and consists of convoluted slits separated by thick cytoplasmic columns. Cross‐striated ciliary rootlets running parallel with and obliquely or transversely to the longitudinal axis of the flame bulbs strengthen the walls of the flame bulbs and, to a lesser degree, that of the capillary. Numerous cristate mitochondria are present in the terminal and proximal canal cells. Cytoplasmic processes extend from the terminal cells into the adjacent tissue, and narrow internal leptotriches extend from the cytoplasm of the terminal cells into the lumen of the flame bulbs. The wall of the capillary contains many interconnected, liquid filled spaces that communicate with the lumen of the capillary, and two septate junctions. Phylogenetic implications of the findings are discussed.

Development and differentiation of the eye in Platynereis dumerilii (Annelida, Polychaeta).

The nereid polychaete, Platynereis dumerilii, possess two pairs of post-trochophoral eyes with one vitreous body each. The development of these eyes has first been observed in 2-day-old larvae. Whether the eye anlagen arise from stem cells or from undifferentiated ectodermal tissue was not determined. At first, the anlagen of the anterior and the posterior eyes adjoin each other. They separate in late 3-day-old larvae. The first separated eye complexes consist each of two supporting and two sensory cells. The supporting cells synthesize two different kinds of granules, the pigment granules of the pigment cup and the prospective tubules of the vitreous body. These tubules accumulate in the distal process of the supporting cell. The vitreous body is formed by compartments of the supporting cells filled with the osmiophilic vitreous body tubules. The short, bulbar photosensory processes bear microvilli that emerge into the ocular cavity. At the apex of each sensory cell process, a single cilium (or occasionally two) arises. The sensory cells contain a different kind of pigment granule within their necks at the level of the pigment cup. The rate of eye development and differentiation varies. New supporting cells are added to the rim of the eye cup. They contribute to the periphery of the vitreous body like onion skins, and sensory cells move between supporting cells. The older the individual compartments of the vitreous body are, the more densely packed is their content of vitreous body tubules. Elongation of the sensory and supporting cell processes of the older cells increases the volume of the eye. The eyespots of the trochophore are briefly described as of the two-celled rhabdomeric type with a single basal body with ciliary rootlet.

Sense receptors of larval Lobatostoma manteri (Trematoda, Aspidogastrea)

Seven types of uniciliate and two types of non-ciliate receptors were found in the larva of Lobatostoma manteri. Uniciliate receptors differ in the length of the cilium, the shape of the cilium, presence or absence of a ciliary rootlet, and the number of electrondense collars. Non-ciliate receptors differ in the shape of the ciliary rootlet. The variety of receptors is smaller than that of larval Multicotyle purvisi, possibly due to the different modes of infection: ingestion of the egg by a snail, hatching of a non-ciliate larva in the stomach and migration into the digestive gland in Lobatostoma, hatching of a ciliate larva in fresh water, inhalation by a snail and migration into the kidney in Multicotyle.

Cytoskeletal elements in arthorpod sensilla and mammalian photoreceptors

Ciliary receptor cells, typified by cilia or modified cilia, are very common in the animal kingdom. In addition to the cytoskeleton of their ciliary processes these receptors possess other specific prominent cytoskeletal elements. Two representative systems are presented: i) scolopidia, mechanosensitive sensilla of various arthropod species; and ii) photoreceptor cells of the retina of the bovine eye. Two cytoskeletal structures are characteristic for arthropod scolopidia: a scolopale typifies the innermost auxiliary cell, and long ciliary rootlets are extending well into the sensory cells. The latter element is also charactristics for the inner segment of the photoreceptor cells in bovine. The scolopale of scolopidia is mainly composed of actin filaments. In the absence of myosin, the uniform polarity of the actin filaments and their association with tropomyosin all indicate a stabilizing role of the filament bundles within the scolopale. This function and a certina elasticity of actin filament bundles may be important during stimulation of the sensilla. The ciliary rootlets of both systems originate at the basal bodies at the ciliary base of the sensory cells and project proximally. These rootlets are composed of longitudinally oriented, fine filaments forming a characteristic regular cross-striation. An α-actinin immunoreactivity was detected within the ciliary rootlets of scolopidia. In addition, antibodies to centrin react with the rootlets of both types of receptors. Since centrin is largely responsible for the contraction of the flagellar rootlets in green algae, contraction may also occur in the ciliary rootlets of insect sensilla and vertebrate photoreceptors. In both system, contraction or relaxation of the ciliary rootlets could serve in sensory transduction or adaptation.

Cortical morphogenesis in Paramecium: A transcellular wave of protein phosphorylation involved in ciliary rootlet disassembly

In Paramecium, the morphogenesis of the cortex at cell division, which assures reconstruction of shape and surface pattern, has been shown to involve transcellular signals which spread across the cortex like a wave, originating principally from the oral apparatus. One of the events these signals control is the reorganization of the ciliary rootlets through a cycle of regression and regrowth. The ciliary rootlets are nucleated on the ciliary basal bodies and form a scaffold extending over the entire cell surface that is important in aligning the basal bodies and the unit territories organized around them in longitudinal rows. We present evidence that the mechanism underlying their reorganization is cell-cycle-dependent phosphorylation of the structural proteins which compose the ciliary rootlets. We have isolated the rootlets and prepared a polyclonal antibody against them. In situ immunofluorescence of dividing cells with the anti rootlet antibody, and with the monoclonal antibody MPM-2 specific for phosphoproteins shows that a wave of phosphorylation of the ciliary rootlets spreads across the cell at division and just precedes their regression. Two-dimensional Western blot analysis of cytoskeleton and isolated rootlets along with alkaline phosphatase treatment demonstrates that the rootlets are composed of phosphoproteins, while experiments with interphase and dividing cells provide direct evidence that hyperphosphorylation of these proteins at division brings about disassembly of the structure.

Centrin- and ?-actinin-like immunoreactivity in the ciliary rootlets of insect sensilla

Long ciliary rootlets are a characteristic feature of the dendritic inner segments of the sensory cells in insect sensilla. These rootlets are composed of highly ordered filaments and are regularly cross-striated. Collagenase digestion and immunohistochemistry reveal that the rootlets are probably not composed of collagen fibers. However, double-labeling experiments with phalloidin and anti-α-actinins show that antibodies to α-actinin react with the ciliary rootlets of the sensilla, but do not stain the scolopale, which is composed of actin filaments as visualized by phalloidin. Antibodies to centrin, a contractile protein isolated from flagellar rootlets of green algae, also stain the ciliary rootlets. Within the ciliary rootlets of insect sensilla, α-actinin may be associated with filaments other than actin filaments. The immunohistochemical localization of a centrin-like protein suggests that contractions probably occur within the rootlets. The centrin-like protein may play a role during the mechanical transduction or adaptation of the sensilla.

Sensory and secretory cells ofOphryotrocha puerilis (Polychaeta)

As revealed by glyoxylic acid induced fluorescence, the protandric polychaeteOphryotrocha puerilis possesses different types of catecholaminergic primary bipolar sensory cells, the perikarya of which are located beneath the epidermis. About 20 of such receptors are situated in each segment but they are mostly found on antennae, palps, urites and parapodial cirri. The dendrites of these sensory neurones run to the cuticle and dilate to form receptive endings. Three different types of dendritic endings could be distinguished: (1) multiciliary receptors with 4–8 cilia and ciliary rootlets, (2) monociliary receptors with microvilli arranged like a funnel and electron-dense cuffs and (3) monociliary receptors of the collar-type with, constantly, ten microvilli surrounding one single central cilium. The latter type is also characterized by rootlet fragments. Dendrites and dilated receptive endings of all three types contain clear (putative secretory) vesicles, multivesicular bodies and mitochondria. Pharmacological treatment (dopamine, reserpine) does not affect the number of secretory vesicles of the receptor neurones. Extra vesicular storage of catecholamines is discussed. Secretory cells of unknown function containing large numbers of electron-dense vesicles are usually found in close association with sensory cells.

A common epitope is shared by ciliary rootlets and cell-cell adherens junctions in ciliated ependymal cells

Using immunoblot, light and electron immunocytochemistry, we investigated the presence and the localization of polypeptides cross-reacting with the monoclonal antibody CC.310 (mAb CC.310), which is mainly directed against a 175K (K = 10(3) Mr) ciliary rootlet protein. In hypothalamic ependymal cultures, the unique antigen recognized by mAb CC.310 was associated with the Triton X-100-insoluble fraction in these cultures and electrophoretically migrated to these cultures and electrophoretically migrated to 94K. mAb CC.310, which appears to be a very suitable marker for ciliated ependymocytes, allowed us to observe ciliogenesis during the growth of the ependymal cultures, from a single spot in each undifferentiated ependymal cell to a massive labeling in ciliated ependymal cells. In fully differentiated ciliated ependymocytes, mAb CC.310 strongly reacted with fibrous structures corresponding to ciliary rootlets, as confirmed by ultrastructural observations. In addition, a weaker immunostaining was also found along the intercellular junctions, and showed that proteins sharing a common epitope are located in ependymal ciliary rootlets and near adherens-type junctional complexes. Immunofluorescence studies confirmed the presence of positive labeling at the level of junctional complexes between cells in two epithelial lines, HeLa and PtK2, in which mAb CC.310 mainly reacted with one polypeptide of 85K.

Ultrastructure of the flame bulbs and protonephridial capillaries of Prorhynchus (Lecithoepitheliata, Prorhynchidae, Turbellaria)

The flame bulb of Prorhynchus is formed by a single cell. Its nucleus is not located in the cytoplasm at the base of the flame. Cilia of the flame have cross‐striated hollow ciliary rootlets converging towards their tips. The maximum number of cilia counted was 13. The weir consists of a single row of longitudinal ribs that contain longitudinal filaments and possess regularly arranged protrusions along their surface appearing as transverse bands in horizontal section. A ‘membrane’ of extracellular material extends between the ribs. and loose material fills the places between the ribs, with a denser layer midway between adjacent ribs. Distally, the ribs fuse to form a continuous tube without a junction. Small protonephridial capillaries lack junctions, larger ducts have lateral flames and patches of long microvilli. Large excretory ducts open into a ciliated and lamellated cavity which is connected by a narrow canal to the excretory pore. The terminal part of the canal close to the pore possesses many cilia and microvilli. Phylogenetic implications of the findings are discussed.

The fine structure of the growth stage oocytes of a lancelet (= amphioxus),Branchiostoma lanceolatum

Summary Oocytes of the European lancelet (Branchiostoma lanceolatum) were fixed for transmission electron microscopy at Banyuls (French Mediterranean) in mid spring, which is just before the spawning season. Special attention is given to the growth stage (= diplotene) oocytes and to their relations with non-germinal cells of the ovary. At the time of year studied, the ovaries contain both relatively small (7–57 μm) and relatively large (97–127 μm) oocytes, but none of medium-size. In addition to the usual cell organelles (like free ribosomes, mitochondria, and endoplasmic reticulum), the oocytes include a number of more specialized structures. Nuage is present throughout the growth stage, both adjacent to the nucleus and scattered elsewhere in the cytoplasm. Other specialized structures do not begin to appear until the oocytes attain a certain diameter. The following structures are first detected at the following oocyte diameters: cortical granules, always in close association with Golgi complexes (20 μm); vacuoles containing presumed precursors of the vitelline layer (30 μm); yolk granules (35 μm); and a central vacuole within the nucleolus (50 μm). The cytoplasm of the largest oocytes includes a few annulate lamellae and some very unusual striated fibers resembling ciliary rootlets. These fibers, which are banded with a 70-nm periodicity (but are evidently not associated with centrioles), occur only at the animal pole of the oocyte, and their functional significance is unknown. The smaller oocytes have relatively smooth plasma membranes, except where endocytotic pits are abundant, whereas the larger oocytes bear abundant microvilli and are covered by a vitelline layer of dense, granular material. During oocyte growth, areas of contact between the oocytes and neighboring non-germinal cells shrink progressively. In the large oocytes, such contacts are limited to the animal pole, where a few processes from non-germinal cells pass through the vitelline layer and terminate on the oocyte plasma membrane at adhaerens type junctions (perhaps mixed with gap junctions). The possible functions and phylogenetic significance of such junctions are discussed.