Post-embryonic larval development and metamorphosis of the hydroidEudendrium racemosum (Cavolini) (Hydrozoa,Cnidaria)

The morphology and histology of the planula larva ofEudendrium racemosum (Cavolini) and its metamorphosis into the primary polyp are described from light microscopic observations. The planula hatches as a differentiated gastrula. During the lecithotrophic larval period, large ectodermal mucous cells, embedded between epitheliomuscular cells, secrete a sticky slime. Two granulated cell types occur in the ectoderm that are interpreted as secretory and sensorynervous cells, but might also be representatives of only one cell type with a multiple function. The entoderm consists of yolk-storing gastrodermal cells, digestive gland cells, interstitial cells, cnidoblasts, and premature cnidocytes. The larva starts metamorphosis by affixing its blunt aboral pole to a substratum. While the planula flattens down, the mucous cells penetrate the mesolamella and migrate through the entoderm into the gastral cavity where they are lysed. Subsequently, interstitial cells, cnidoblasts, and premature cnidocytes migrate in the opposite direction, i.e. from entoderm to ectoderm. Then, the polypoid body organization, comprising head (hydranth), stem and foot, all covered by peridermal secretion, becomes recognisable. An oral constriction divides the hypostomal portion of the gastral cavity from the stomachic portion. Within the hypostomal entoderm, cells containing secretory granules differentiate. Following growth and the multiplication of tentacles, the head periderm disappears. A ring of gland cells differentiates at the hydranth's base. The positioning of cnidae in the tentacle ectoderm, penetration of the mouth opening and the multiplication of digestive gland cells enable the polyp to change from lecithotrophic to planktotrophic nutrition.     

Non-neural ectoderm is really neural: evolution of developmental patterning mechanisms in the non-neural ectoderm of chordates and the problem of sensory cell homologies

In chordates, the ectoderm is divided into the neuroectoderm and the so-called non-neural ectoderm. In spite of its name, however, the non-neural ectoderm contains numerous sensory cells. Therefore, the term "non-neural" ectoderm should be replaced by "general ectoderm." At least in amphioxus and tunicates and possibly in vertebrates as well, both the neuroectoderm and the general ectoderm are patterned anterior/posteriorly by mechanisms involving retinoic acid and Hox genes. In amphioxus and tunicates the ectodermal sensory cells, which have a wide range of ciliary and microvillar configurations, are mostly primary neurons sending axons to the CNS, although a minority lack axons. In contrast, vertebrate mechanosensory cells, called hair cells, are all secondary neurons that lack axons and have a characteristic eccentric cilium adjacent to a group of microvilli of graded lengths. It has been highly controversial whether the ectodermal sensory cells in the oral siphons of adult tunicates are homologous to vertebrate hair cells. In some species of tunicates, these cells appear to be secondary neurons, and microvillar and ciliary configurations of some of these cells approach those of vertebrate hair cells. However, none of the tunicate cells has all the characteristics of a hair cell, and there is a high degree of variation among ectodermal sensory cells within and between different species. Thus, similarities between the ectodermal sensory cells of any one species of tunicate and craniate hair cells may well represent convergent evolution rather than homology.     

Scanning electron microscopic observations on the 9-day opossum (Didelphis virginiana) embryo

All three germ layers are present in the opossum embryo by the 9th prenatal day. The embryo proper is part of, and continuous with, the remainder of the chorionic wall. The wall of the yolk sac-chorion away from the embryo consists only of an outer covering of ectoderm and an inner layer of endoderm. Ectodermal cells covering the neural folds have dome-shaped apices and often show large, bleb-like expansions. Microvilli are short and few in number. The apical surfaces of ectodermal cells that overlie the parietal mesoderm are relatively smooth and show scattered, short microvilli that tend to be concentrated at cell junctions. The apices of ectodermal cells that cover the extraembryonic region are more rounded, and the cells balloon from the surface. Each cell shows abundant elongate microvilli and occasional cytoplasmic blebs. Endodermal cells that line the chorion and form the third (innermost) layer of the embryo are similar in their surface morphology.     

Change of the differentiation pattern of amphibian ectoderm after the increase of the initial cell mass

Summary Early amphibian gastrula ectoderm (Triturus alpestris) has been treated with vegetalizing factor. While normal sandwiches (animal caps of two eggs) differentiated mainly into endoderm derived tissues, giant-sandwiches (a combination of 8 animal caps) formed mesodermal and neural tissues in addition. The results support the interpretation that ectoderm will differentiate into endoderm derived tissues when all or nearly all cells are induced (presumably depending on certain threshold concentrations of the inducer). This is the case in the normal sandwich after treatment with high concentrations of vegetalizing factor for 24 h. However, in a giantsandwich it must be assumed that only the cells in the vicinity of the inducer will be triggered to differentiate into endoderm derived tissues. Mesodermal structures will be formed by secondary interactions between the induced ectoderm (endoderm) and non induced ectodermal cells. The induction of neural structures could be explained as a further interaction between mesodermalized and non induced ectodermal cells. This chain of events is compared with the steps of determination in normogenesis.     

Collar cells in planula and adult tentacle ectoderm of the solitary coral Balanophyllia regia (anthozoa eupsammiidae)

Summary The planula larva of the solitary coral Balanophyllia regia has an ectoderm of flagellate, diplosomal collar cells. The collar of these cells is composed of a ring of microvilli linked with mucus strands. Four types of flagellate gland cells, three types of nematocyst and spirocysts are present in the planula ectoderm. The function of these ectoderm cells is discussed. The mesogloeal muscular and packing tissues of the planula are briefly described. The tentacle of the adult coral, examined for comparison, has an ectoderm of flattened flagellate cells with a shallow collar. Collar cells similar to those of the planula are occasionally found on the tentacle and their function is not known. Independent sensory cells built on a modified collar cell plan with collar of thickened microvilli are common in the tentacle. These are quite separate from the three kinds of tentacular nematocyte. Distended glandular areas occur in the tentacle ectoderm. The flagellate tentacle gastrodermis, muscle and mesogloeal region are briefly described. The evolutionary significance of collar cell ectoderm in a planula is discussed and the occurrence of collar cells throughout the animal kingdom, reviewed.I am most grateful to Paul Tranter of the Plymouth Laboratory for providing material and to Gareth Morgan for assistance with electron microscopy.     

A fine structural analysis of the epidermis and cuticle of the oligochaete aeolosoma Bengalense stephenson

The fine structure of the epidermis and cuticle has been described for the oligochaete Aeolosoma bengalense. The epidermis is a pseudostratified epithelium and consists of the following cell types: ciliated and nonciliated supportive cells, pigment cells and associated satellite cells, mucous cells, basal cells, and ciliated non-supportive columnar cells. Overlying and restricted to the supportive cells is a delicate cuticle composed of: (a) a discontinuous layer of membrane-bounded surface particles; (b) a thin filamentous layer of moderate electron density just under the surface particles; (c) a thicker inner filamentous layer of low electron density. Digestion with pronase effectively removes the cuticle. This, together with the fact that it stains with alcian blue and ruthenium red, indicates that the cuticle contains an acid mucopolysaccharide. Regeneration of the cuticle, following pronase treatment, is marked by the elaboration of numerous microvilli by the supportive cells. Most of the microvilli are transitory and evidence supports a microvillar origin for the cuticular surface particles. The presence of cuticular surface particles may be a characteristic shared in common by all oligochaetes and, perhaps, some polychaetes.     

Ultrastructure of maxillary gland of Antrobathynella stammeri (Syncarida,Malacostraca)

The maxillary gland of the highly adapted stygobiont species, Antrobathynella stammeri (Bathynellacea, Syncarida), consists of an end sac, an excretory tubule, and a terminal duct. No valve was found. The excretory tubule forms a loop extending back into the fourth thoracic segment. The end sac is composed of five typical podocytes. Ultrastructurally and functionally, two cell types characterize, respectively, proximal and distal sections of the excretory tubule. Epithelial cells are covered with extremely broad (up to 0.4 μm) microvilli. A basal labyrinth was not seen. Therefore, it is unlikely that the maxillary gland is able to produce a hypoosmotic urine necessary for freshwater animals. Tubule cells can be surrounded by parenchymal cells that produce numerous vesicles, suggesting possible physiological interactions between tubule cells and parenchyma. The ectodermal terminal duct is lined with cuticle and is differentiated into a pulsatile body consisting of two interconnected ampules. The first functions as a bladder. The second ampule, under muscular control, excretes the urine. Pulsatile body, looping tubule, and broad microvilli appear to be distinctive features of the bathynellacid excretory organ. © 1996 Wiley-Liss, Inc.     

Communication compartments in the ectoderm of embryos of Patella vulgata

Summary Patterns of gap junctional communication in the ectoderm of embryos of Patella vulgata have been studied by intracellular injection of the fluorescent dye Lucifer Yellow, and by analysis of its subsequent spread to adjacent cells (dye-coupling). We found that dye-coupling became progressively restricted to different domains of the ectoderm, forming communication compartments. These communication compartments are characterized by their high coupling abilities within the compartment, and reduction of coupling across their boundaries. During development, the pretrochal (anterior) ectoderm becomes subdivided into two communication compartments, the apical organ and the anlage of the head ectoderm. The posttrochal (posterior) ectoderm becomes subdivided into different communication compartments in two successive phases. Firstly, in the 15-h embryo the dorsal and ventral domains of the ectoderm form separate communication compartments. A dorso-ventral communication boundary restricts the passage of dye between the two domains. Secondly, in the 24-h embryo dye-coupling becomes further compartmentalized in both the dorsal and ventral domains. These compartments correspond to the anlagen of different ectodermal structures. In order to study whether any level of coupling persists between the ectodermal compartments we injected currents through a microelectrode inserted into one cell of one compartment and monitored its spread by means of a second microelectrode inserted into one cell of another compartment (electrical coupling). Despite the absence of dye-coupling, electrical coupling between the ectodermal dye-coupling compartments was detected, which suggests that some level of communication is maintained between compartments. Our results demonstrate that within the ectoderm layer of Patella vulgata the transfer of dyes becomes progressively restricted to communication compartments and, concomitantly with the specification of the different ectodermal anlagen, these compartments become subdivided into smaller communication compartments.     

From larval bodies to adult body plans: patterning the development of the presumptive adult ectoderm in the sea urchin larva

Echinoderms are unique among bilaterians for their derived, nonbilateral adult body plan. Their radial symmetry emerges from the bilateral larval body plan by the establishment of a new axis, the adult oral–aboral axis, involving local mesoderm–ectoderm interactions. We examine the mechanisms underlying this transition in the direct-developing sea urchin Heliocidaris erythrogramma. Adult ectoderm arises from vestibular ectoderm in the left vegetal quadrant. Inductive signals from the left coelom are required for adult ectodermal development but not for initial vestibule formation. We surgically removed gastrula archenteron, making whole-ectoderm explants, left-, right-, and animal-half ectoderm explants, and recombinants of these explants with left coelom. Vestibule formation was analyzed morphologically and with radioactive in situ hybridization with HeET-1, an ectodermal marker. Whole ectodermal explants in the absence of coelom developed vestibules on the left side or ventrally but not on the right side, indicating that left–right polarity is ectoderm autonomous by the gastrula stage. However, right-half ectodermal explants robustly formed vestibules that went on to form adult structures when recombined with the left coelom, indicating that the right side retains vestibule-forming potential that is normally suppressed by signals from the left-side ectoderm. Animal-half explants formed vestibules only about half the time, demonstrating that animal–vegetal axis determination occurs earlier. However, when combined with the left coelom, animal-half ectoderm always formed a vestibule, indicating that the left coelom can induce vestibule formation. This suggests that although coelomic signals are not required for vestibule formation, they may play a role in coordinating the coelom–vestibule interaction that establishes the adult oral–aboral axis.     

The Protonephridia of the Arctic Kinorhynch Echinoderes aquilonius (Cyclorhagida,Echinoderidae)

The cyclorhagid kinorhynch Echinoderes aquilonius Higgins & Kristensen, 1988 possesses a single pair of protonephridia located in segments 10 and 11. The protonephridia consist of: (1) three terminal cells T-1, T-2. T-3, each with two cilia; (2) a single non-ciliated canal cell; (3) a nephridiopore cell with many microvilli and a cuticular sieve plate. The protonephridia of Echinoderes are presumed to develop from the ectoderm near the area of the sieve plate on the eleventh segment, and are suspended in the dorso-lateral pseudocoelomic cavity where they are surrounded by a basal lamina. One of the terminal cells (T-1) secondarily penetrates the basal lamina of the tenth segment and a part of the cell attaches to the cuticle. The kinorhynch protonephridia are compared with the excretory organs of other Bilateria. expecially the ‘aschelminths’, and apomorphic characters of the kinorhynch protonephridia are defined.     

Embryonic development and metamorphosis of the scyphozoan <Emphasis Type="Italic">Aurelia</Emphasis>

We investigated the development of Aurelia (Cnidaria, Scyphozoa) during embryogenesis and metamorphosis into a polyp, using antibody markers combined with confocal and transmission electron microscopy. Early embryos form actively proliferating coeloblastulae. Invagination is observed during gastrulation. In the planula, (1) the ectoderm is pseudostratified with densely packed nuclei arranged in a superficial and a deep stratum, (2) the aboral pole consists of elongated ectodermal cells with basally located nuclei forming an apical organ, which is previously only known from anthozoan planulae, (3) endodermal cells are large and highly vacuolated, and (4) FMRFamide-immunoreactive nerve cells are found exclusively in the ectoderm of the aboral region. During metamorphosis into a polyp, cells in the planula endoderm, but not in the ectoderm, become strongly caspase 3 immunoreactive, suggesting that the planula endoderm, in part or in its entirety, undergoes apoptosis during metamorphosis. The polyp endoderm seems to be derived from the planula ectoderm in Aurelia, implicating the occurrence of “secondary” gastrulation during early metamorphosis.     

A quantitative and morphological study of ectodermal microvilli in ten areas in the control and experimental prenatal rat.

This study provides a baseline of mainly quantitative morphologic information in relation to the ectodermal microvilli in ten areas in the control and experimental prenatal rat with special reference to the period of neural tube closure. Embryos from six litters were used and ten areas were examined mainly with the scanning electron microscope. Statistical analysis showed no significant difference between litters nor between the ten areas examined. In the 376 hr (day 15.6) fetus only the ectodermal cells of the nostril region demonstrated a rich population of microvilli, a fact possibly associated with late maturation of that region. Some evidence is provided to show that there is an increase in the population of microvilli in the 276 hr (day 11.5) embryo following experimentally induced zinc deficiency and introduction of nicotine in the culture medium. The possible mechanisms underlying the increase in ectodermal microvilli are i) an attempt by ectodermal cells to absorb nutrients, ii) a reflection of cells under stress, iii) failure of early embryonic ectodermal cells to shed microvilli normally associated with developmental changes, and iv) a generalized developmental delay rather than some cellular response to a trace element nutritional deficiency and a teratogen.     

Regionalisation of the mouse embryonic ectoderm: allocation of prospective ectodermal tissues during gastrulation

The regionalisation of cell fate in the embryonic ectoderm was studied by analyzing the distribution of graft-derived cells in the chimaeric embryo following grafting of wheat germ agglutinin--gold-labelled cells and culturing primitive-streak-stage mouse embryos. Embryonic ectoderm in the anterior region of the egg cylinder contributes to the neuroectoderm of the prosencephalon and mesencephalon. Cells in the distal lateral region give rise to the neuroectoderm of the rhombencephalon and the spinal cord. Embryonic ectoderm at the archenteron and adjacent to the middle region of the primitive streak contributes to the neuroepithelium of the spinal cord. The proximal-lateral ectoderm and the ectodermal cells adjacent to the posterior region of the primitive streak produce the surface ectoderm, the epidermal placodes and the cranial neural crest cells. Some labelled cells grafted to the anterior midline are found in the oral ectodermal lining, whereas cells from the archenteron are found in the notochord. With respect to mesodermal tissues, ectoderm at the archenteron and the distal-lateral region of the egg cylinder gives rise to rhombencephalic somitomeres, and the embryonic ectoderm adjacent to the primitive streak contributes to the somitic mesoderm and the lateral mesoderm. Based upon results of this and other grafting studies, a map of prospective ectodermal tissues in the embryonic ectoderm of the full-streak-stage mouse embryo is constructed.     

Interaction of induced and uninduced cells during primary induction

Summary UsingTriturus pyrrhogaster embryos, the effects of uninduced cells on the differentiation of induced cells were investigated. The inducing stimulus was given to the presumptive ectoderm of early gastrulae by treatment with protein sooution from guinea pig bone-marrow. Mesodermal induction was evoked in the ectodermal explants. After the treatment, some of the ectodermal explants were cut into pieces 1/8 of their original size and combined with untreated presumptive ectoderm. Mesodermal tissues were differentiated in the combined explants too, but the mesodermal tissues evoked in these combined ectodermal explants were different in their regional characters from these in uncombined explants; dorsal structures, such as notochrod and muscle, were observed predominatly in the latter, whereas the dominant structures observed in the former were ventral ones, such as mesothelium and mesenchyme. The shifting of the regional characters in the combined explants was regarded as the result of an unknown effect from the uninduced cells.     

Neural ectoderm,neural crest,and placodes: Contribution of the otic placode to the ectodermal lining of the embryonic opercular cavity in Atlantic cod (Teleostei)

Development of neural ectoderm, neural crest, and otic placode with special reference to a new placodal derivative, the ectodermal lining of the opercular cavity, is described in a teleost fish, the Atlantic cod Gadus morhua, from a stage-by-stage examination of embryonic development. The ectodermal lining of the opercular cavity forms by invagination of the otic placode. The neural plate “infolds” by a wave of cellular rearrangement that transforms the neural plate into a neural rod. This transformation creates a distinct dorsal ectodermal cell layer. When the neural rod is arranged as monostratified columnar cells in the forebrain and midbrain, dorsal ectoderm at the midbrain level thickens lateral to the neural rod to form a cell cluster—the presumptive neural crest and placode. Upon migration of the neural crest from the postoptic midbrain, the dorsolateral area of the dorsal ectoderm thickens and segregates from the neural crest as a placode that is continuous with the presumptive lens placode. As the neural crest migrates from the hindbrain, this placode extends along the hindbrain as a single continuous cluster of cells. At the onset of formation of the lens placode, this continuous placode becomes the placode in the postoptic area of the midbrain and separates into the otic placode at the hindbrain. The otic placode gives rise to the otic neuromast and probably the otic lateral line nerves rostrally and to the ectodermal cell lining of the opercular cavity and otic vesicles caudally. The opercular cavity forms by invagination of the otic placode, creating an internal lumen lined by ectoderm that becomes continuous with evaginated endodermal pharyngeal cells. Free neuromasts are observed along the trailing edge of the external opening of the opercular cavity, which lies horizontally, ventral to the otic vesicles. As embryos develop to hatching, the opening rotates and takes up a vertical position. The adult opercular apparatus, including associated bones and muscles, forms during larval stages. The otic neuromast may be a remnant of neuromasts in the spiracle organ. The spiracle opening lies between the mandibular and hyoid arches, whereas the opercular cavity opens between the hyoid and the first branchial arches. The spiracle opening is, therefore, not homologous with the external opening of the opercular cavity, although the cell lining of the spiracle opening may be of placodal origin. J Morphol 231:231–252, 1997. © 1997 Wiley-Liss, Inc.     

A novel guanine exchange factor increases the competence of early ectoderm to respond to neural induction.

Inductive interactions between different cell layers have an extremely important role in early embryogenesis. One of the most intensively studied and best characterised of these is the induction of neural tissue from ectodermal cells by the dorsal mesoderm. The competence of ectodermal cells to respond to neural induction varies according to dorsal-ventral position; with dorsal ectoderm (much of which forms the neural plate) having a far higher competence. Here we show that overexpression of the nucleotide exchange factor lfc increases ectodermal competence for neural induction as well as the amount of neural tissue in the whole embryo. Lfc is expressed pan ectodermally soon after gastrulation and may respond to an early determinant of dorsal ectoderm.     

Etude ultrastructurale de l'évolution du tégument de la Sangsue Hirudo medicinalis L. (Annélide,Hirudinée) au cours d'un cycle de mue / Developmental changes in the integument of the leech Hirudo medicinalis L. (Annelida,Hirudinea) during a molting cycle. An ultrastructural study

Summary

The integument of the leech Hirudo medicinalis is mainly composed of a single layer of cuticle-secreting epidermal cells. The cuticle is made up of collagen fibers which support a layer of membrane-bound epicuticular projections.

Shedding of the old cuticle is preceded by the formation of a new cuticle. The epicuticular projections are the first to develop: they originate from the tips of numerous microvilli of the epidermal cells. As soon as it appears, the newly-formed collagen layer is firmly attached to the epidermal cells by numerous hemidesmo-somes, whereas the old cuticle is no longer connected with the epidermal surface. The epidermal cells exhibit marked characteristics of secretory activity during the laying down of the new cuticle.

The observations are discussed in connexion with recent findings of high ecdysteroid levels in leeches at the beginning of the molting cycle.     

Sperm storage by spermatodoses in the spermatheca of Trioza alacris (Flor, 1861) hemiptera,psylloidea, triozidae: A structural and ultrastructural study

Female insects generally store sperm received during mating in specific organs of their reproductive tract, i.e., the spermathecae, which keep the sperm alive for a long time until fertilization occurs. We investigated spermatheca morphology and ultrastructure in the psylloidean insect Trioza alacris (Flor, 1861 ) in which spheroidal sperm packets that we refer to as ‘spermatodoses’ are found after mating. The ectoderm‐derived epithelium of the sac‐shaped spermatheca that has a proximal neck, consists of large secretory and flat cuticle‐forming cells. Secretory cells are characterized by a wide extracellular cavity, bordered by microvilli, in which electron‐dense secretion accumulates before discharge into the spermathecal lumen. The cuticle‐forming cells produce the cuticular intima of the organ and a peculiar specialized apical structure, through which secretion flows into the lumen. At mating, the male transfers bundles of sperm cells embedded in seminal fluid into the spermathecal neck. Sperm cells proceed towards the spermathecal sac lumen, where they are progressively compacted and surrounded with an envelope that also encloses secretions of both male and female origin. We describe the formation of these sperm containing structures and document the contribution of the female secretion to spermatodose or female‐determined spermatophore construction. We also discuss the choice of the term ‘spermatodose’ for T. alacris and suggest it be used to refer to sperm masses constructed in the female reproductive organs, at least when they involve the contribution of female secretion. © 2011 Wiley Periodicals, Inc.     

Neurobiology of the gorgonian coelenterates,Murcea californca and Lophogorgia chilensis

Summary Diffuse and synaptic nerve nets are present in the coenenchymal mesoglea and ectoderm of Muricea and Lophogorgia colonies. The nerve nets extend into the polyp column and tentacles maintaining a subectodermalmesogleal position. The density of nerve elements is low in comparison with similar nerve nets found in pennatulids.In the column of the polyp anthocodium, and throughout the oral disk region, neurons cross the mesoglea and enter the polyp endoderm. These neurons presumably connect with the endodermal nerve net which innervates the septal musculature. The trans-mesogleal neurons probably represent the connection between colonial and polyp nervous systems.In the tentacles, longitudinal ectodermal musculature is present with an overlying nerve plexus. These muscles and nerves, as well as tentacular sensory cells, are well represented in the oral side of the tentacles only.Presumed sensory cells form ciliary cone complexes in which one cell possesses an apical cilium. The other cells as well as the centrally located nematocyte contribute microvilli to the cone. The basal portion of the sensory cells is drawn into one or more neurite-like processes which enter the ectodermal nerve plexus. Similar processes form synapses with longitudinal muscle cells and nematocytes. The sensory cells of the ciliary cones presumably include chemoreceptors which can activate or modify nematocyst discharge, local muscle twitches, and tentacle bending.This work was supported by Office of Naval Research Contract N00014-75-C-0242, NSF Grant BMS 74-23242 and General Research Funds of the University of California, Santa Barbara. We wish to thank Dr. Steven K. Fisher for the use of facilities in his lab. This paper is part of a thesis to be submitted by R.A.S. to the Department of Biological Sciences, University of California, Santa Barbara in partial fulfillment of the requirements for the Ph. D.