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Laser scanning confocal microscopy is used to reveal the changes that occur in the RFamide-positive nerve net as a free-swimming, solid hydrozoan planula larva is transformed into a sessile, hollow, young polyp. Seven stages of development in Pennaria tiarella are described: planula competent to metamorphose, attaching planula, disc, pawn, crown, developing polyp, and developed primary polyp. The RFamide-positive nervous system undergoes dramatic reorganization during metamorphosis: (1) larval neurons degenerate; (2) new neurons differentiate and reform a nerve net; and (3) the overall distribution pattern of the nervous system changes. This study confirms earlier observations on RFamide-positive neurons of Hydractinia which also show the loss of these cells after the onset of metamorphosis.
The ultrastructure of paraspermatogenesis is examined in the littorinid subfamily Littorininae, with special emphasis on Littoraria (Palustorina) articulata (Philippi 1846). In particular the study focuses on the fate of the nucleus and origin of the rod bodies during parasperm development. Parasperm of the Littorininae are rounded or oblong cells, which undergo an abortive meiosis and eliminate part of the nucleus but often retain a nuclear remnant. The cytoplasm is filled with numerous spherical vesicles in all Littorininae, but in Littoraria (and in certain species of Nodilittorina, Tectarius and Cenchritis)dense ‘rod-bodies’ also occur. Littoraria (Palustorina)are unique in possessing a flagellum-like structure termed the ‘pseudotrich’, which lacks an axoneme but contains microtubules during its development. Paraspermatogonia differ from euspermatogonia in the structure of the nucleus and in the extensive rough endoplasmic reticulum (RER) and swollen cytoplasm. Two types of secretions develop in Littoraria: (1) numerous, spherical granules (composed of putative glycoprotein, also seen in other Littorininae) and (2) rhomboid granules (composition uncertain but reacting positively to RNA stains; these granules arising within RER cisternae close to the nucleus). As the rhomboid granules fuse to form the larger, rod-bodies (polygonal in cross section), the RER membrane enclosing the rod-bodies becomes confluent with the outer nuclear membrane, thereby forming a common compartment. Results of this study clearly show that the rod-bodies are secretions of the RER cisternae and not, as claimed in some light microscopic accounts, the product of fusion of eusperm nuclei which have entered the parasperm cytoplasm (either by active eusperm penetration or by phagocytosis). Developmental characteristics of littorinid parasperm show differences between species and may, in some cases, provide characters diagnostic of subgenera.
The ultrastructural features of the ovary and oogenesis have been described in 6 species of patellid limpets from South Africa. The ovary is a complex organ that is divided radially into numerous compartments or lacunae by plate-like, blind-ended, hollow trabeculae that extend from the outer wall of the ovary to its central lumen. Trabeculae are composed of outer epithelial cells, intermittent smooth muscle bands, and extensive connective tissue. Oocytes arise within the walls of the trabeculae and progressively bulge outward into the ovarian lumen during growth while partially surrounded by squamous follicle cells. During early vitellogenesis, the follicle cells lift from the surface of the underlying oocytes and microvilli appear in the perivitelline space. Follicle cells restrict their contact with the oocytes to digitate foot processes that form desmosomes with the oolamina. When vitellogenesis is initiated, the trabecular epithelial cells hypertrophy and become proteosynthetically active. Yolk synthesis involves the direct incorporation of extraoocytic precursors from the lumen of the trabeculae (hemocoel) into yolk granules via receptor-mediated endocytosis. Lipid droplets arise de novo and Golgi complexes synthesize cortical granules that form a thin band beneath the oolamina. A fibrous jelly coat forms between the vitelline envelope and the overlying follicle cells in all species.
Light and scanning electron microscopy were used to examine protoconch form in eight species of planktotrophic heterobranch larvae, including four nudibranch species with a coiled (type 1) protoconch, two nudibranch species with an inflated (type 2) protoconch, and two cephalaspid species with a coiled protoconch. The coiled protoconchs of the cephalaspids and nudibranchs have a similar form at hatching, and shell growth up to metamorphic competence is hyperstrophic. Shell added to coiled protoconchs during the larval stage overgrows all but the left wall of the initial protoconch that exists at hatching. The entire protoconch of cephalaspids, including overgrown areas, is retained through metamorphosis. However, during later larval development in nudibranchs with a coiled protoconch, overgrown shell is completely removed by dissolution. As a result, regardless of whether nudibranch larvae hatch with an inflated or coiled protoconch type, the protoconch is a large, hollow cup at metamorphic competence. The protoconch of nudibranchs is shed at metamorphosis and absence of a post-metamorphic shell is correlated with absence of visceral coiling in this gastropod group. Internal dissolution of the coiled protoconch in nudibranchs allows the left digestive gland to uncoil prior to metamorphic shell loss. Retention of overgrown protoconch whorls in cephalaspids allows the attachment plaque of the pedal muscle to migrate onto the parietal lip of the post-metamorphic shell. Release from this constraint in nudibranchs, in which the larval pedal muscles and the entire protoconch are lost at metamorphosis, may have permitted internal protoconch dissolution and precocious uncoiling of the visceral mass, as well as evolutionary emergence of the inflated larval shell type.
We describe the genital system of the aeolid nudibranch gastropod Aeolidiella glauca as a basis for our ongoing analysis of the mating system of this hermaphroditic species. In addition we give a short account of its mating behavior. A. glauca has an androdiaulic genital system with a proximally situated semiserial seminal receptacle. There is no bursa copulatrix. After fertilization, eggs pass through six glands, i.e., the capsule gland and the female gland mass which is comprised of five histologically differentiated parts. The prostate is a long, glandular tube. The everted, unarmed penis is very large and bears a series of 3-4 hook-shaped lobes consisting only of a simple, ciliated epithelium on its ventral side. Their function is unknown. After courtship, which involves moving in circles followed by resting in a head-to-head position, reciprocally touching each other with the tentacles, the slugs glide into a position where the everted genital atria are in contact. The huge penes are protruded simultaneously shortly after this contact occurs. Each animal strokes its partner's back with the penis and deposits a spermatophore of undetermined shape onto the partner's notum. Sperm enter the recipient through histolysis. How the sperm find their way to the seminal receptacle is not known.
We have constructed molecular phylogenetic trees for members of the Sphaeriidae in order to test proposed generic level relationships, and to reconstruct the evolutionary pattern of parental care, in this exclusively freshwater family of heterodont bivalves. An -480 nucleotide fragment of the mitochondrial large ribosomal subunit (16s rDNA) was sequenced for 4 corbiculid outgroups in addition to 19 sphaeriid ingroup taxa. Ingroup species were obtained from North and South America, Europe, and Australasia and included representatives of the main sphaeriid genera. Our analyses support four primary conclusions: 1) the Sphaeriinae are robustly monophyletic with respect to Eupera platensis; 2) the genus Pisidium is paraphyletic and P. sterkianum is sister to the 17 other sphaeriine taxa in our dataset; 3) synchronous brooding is the ancestral reproductive pattern in the Sphaeriinae; 4) the sequential brooders form a clade in which Musculium taxa are monophyletic and nested among lineages of Sphueriunz. Our gene trees reveal an evolutionary progression in parental care complexity from the relatively simple pattern in the Euperinae, to the origin of brood sacs and of extraoogonial embryonic nutrition in the common ancestor of the Sphaeriinae, and ultimately to the development of sequential brooding in SphaeriudMusculium taxa.
A transmission electron microscope study of fresh and cryopreserved Nereis virens larvae at the three chaetiger stage is described with special emphasis on examining the structure of the photoreceptors and surface ciliation of the head, the midgut epithelium, and muscle cells. Complex ectodermal structures such as the developing rhabdomeric adult eyes were unaffected by the cryopreservation procedure. Some loss of surface cilia on the prostomium was observed but is not life-threatening though it may restrict ciliary swimming in the recovered larvae. Loss of pigment from the prostomium is caused by osmotic stress. Structural damage was observed in the digestive tissues of the larvae cryopreserved before or after the optimum stage of development. This damage is potentially more serious and may account for the relatively short time period during development where cryopreservation can be successfully applied.
Attachment and gas exchange are design problems for the spheroidal egg masses that annelids and molluscs tether in sediments. At an intertidal site in Bodega Harbor, a high proportion of masses of arenicolid worms became detached, but they were not stranded higher than attached masses, and the embryos developed to advanced stages. The large masses, weighing 100-200 g, challenge predictions on limiting size for oxygen supply by diffusion. The estimated concentrations of embryos and thickness of the embryo layer exceed the limits predicted by a simple model for oxygen supply by diffusion from the surrounding water, but several features may enhance oxygen supply to embryos. (1) There is an internal cavity. Distal tears in the gel layer admit pulses of water into the center, thus adding an inner surface for gas exchange. (2) The motile embryos aggregate near the outer and sometimes inner surfaces of the gel layer. Embryos migrate in the gel toward regions of greater oxygen concentration. (3) Pennate diatoms colonize the masses, coating the surfaces and penetrating into the gelatinous matrix. Photosynthesis can exceed respiration when a mass is in the light. The pH in gel changes diurnally from a low of ∼7 to as much as 9.5, indicating an excess of respiration over photosynthesis at night and at least sometimes an excess of photosynthesis over respiration by day.
The terrestrial polychaete Hrabeiella periglandulata has many features in common with the Clitellata and the polychaete taxon Parergodrilidae. An ultrastructural investigation of the central nervous system and the sense organs of H. periglandulata individuals was undertaken to look for structural similarities with these taxa as well as to elucidate whether these structures might exhibit adaptive characters typical of terrestrial annelids in general. The central nervous system of H. periglandulata is subepidermal and consists of a brain situated in the first achaetigerous segment. The circumoesophageal connectives are without dorsal and ventral roots, and the ventral nerve cord has closely associated connectives and ill-defined ganglia. In contrast to clitellates and the terrestrial parergodrilid Parergodrilus heideri, nuchal organs are present. They are internal and highly modified compared with those of marine polychaetes but are similar to those of the intertidal parergodrilid Stygocapitella subterranea. A pair of ciliary sense organs is present inside the brain, resembling similar structures in many microdrile oligochaetes. These observations indicate that there are, in fact, structural similarities between the nervous system and the sense organs of clitellates, parergodrilids, and Hrabeiella individuals. These similarities may very likely be the result of convergent evolution in adaptation to the terrestrial environment.
The apex of the larval antenna of the crane fly Nephrotoma suturalis has 6 cuticular sensilla that stained intensely black with silver nitrate, which indicates their porosity. The large sensory cone is innervated by 14 neurons and the 3 small, smooth surfaced, conical pegs have 4 neurons each. The small and large cylindrical sensilla with their smooth walls and pleated apices are innervated by 4 and 6 nerve cells, respectively. The 15 sensilla on the apex of the maxillary palp are all stained by silver nitrate. These sensilla are of five types: 7 type A sensilla with a smooth surface, a distinct apical pore, and 3 or 4 neurons; 2 type B sensilla with a smooth surface, many pores, and 5 neurons; 1 type C sensillum with a grooved surface, a large apical pore, smaller pores in the grooves, and 6 neurons; 3 type D sensilla with a smooth surface, a grooved apex that is elongated into a projection, and 4 neurons; 2 type E sensilla with many pores covering the surface, leaf-like appearance, and 4 neurons. The number and types of sensilla are similar to those in other nematocerous larvae, but in the many different forms of sensilla and the structure of the sensory cone, these tipulid larvae differ greatly from other larvae of lower Diptera.
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