Body structure of marine sponges

Summary Each choanocyte chamber of Petrosia ficiformis is formed by a slightly outpocked choanocyte epithelium and by a ring of three or four uniflagellated cone cells surrounding the apopyle. The apopyle opens into a small aphodus, which leads the water flow to larger excurrent canals. Pinacocytes of the incurrent canal system cover the basal surface of the choanocytes and separate them from the incurrent canals and the mesenchyme. The water flows into the chambers by pores in the pinacocyte cover and then through gaps between adjacent choanocytes. To our knowledge this is the first report of a leuconoid canal system in which choanocyte chambers are covered by a pinacocyte epithelium of the incurrent canal system that isolates the chambers from the mesenchyme. A future comprehensive revision of the types of canal systems in sponges seems to be necessary. Permanent affiliation: Department of Biology and Health Sciences, University of Hartford, West Hartford, CT 06117, USA     

Die Aufnahme partikulärer Nahrung beiReniera sp. (Porifera)

The choanocyte chambers of the marine spongeReniera sp. protrude with their curved outer surface free into the incurrent canals. The water is sucked into the chambers by cavities between the choanocytes. Particles up to 1 µm in diameter may enter the chambers with the water current. These particles are trapped on the outer surface of the choanocyte collars and are ingested by the choanocytes and processes of the pinacocyte epithelium of the incurrent canal system, which project into the chambers. Bigger particles are retained in the incurrent canals mainly on the outer surface of the choanocyte chambers. They are ingested by pinacocytes of the canal wall and transported to cells of the mesenchyme. The present investigation shows the great importance of the pinacocyte epithelium of the incurrent canal system for suspension feeding inReniera sp.     

Body structure of marine sponges

Summary Specimens of Haliclona elegans (Bowerbank, 1866) are covered by a thin, double layered dermal membrane extending over large subdermal spaces. The pores in the dermal membrane are formed by single porocytes with one or sometimes several pores in the center of the cell. The subjacent tissue shows a faintly developed mesenchyme and numerous big choanocyte chambers projecting into lacunar spaces of the incurrent canal system. The outer surface of the chambers is directly covered by the pinacocyte epithelium of the incurrent canal wall, which also separates them completely from the mesenchyme. Water influx into the chambers is guaranteed by prosopylar openings in the pinacocyte cover at the outer chamber surface. The chambers are connected to the excurrent canal system in the eurypylous way by wide apopyles, each of which is surrounded by a small ring of flagellated cone cells. About 15% of the choanocyte chambers in H. elegans contain central cells, which are thought to derive from migrating pinacocytes of the canal systems.     

Formation and construction of asexual buds of the freshwater spongeRadiospongilla cerebellata (Porifera,Spongillidae)

Summary The buds ofRadiospongilla cerebellata are formed asexually. Budding can be induced experimentally by injuring the sponge. The first sign of budding is a slight elevation of some surface areas, which proceed to rise rapidly so that they soon protrude conspicuously from the surface of the sponge. As a bud develops, the broad base joining it to the mother sponge narrows to a stalk, which finally breaks. The free buds drift in the water for 15–20 min and then settle, forming new sessile sponges. The buds, 1.5–2.5 mm in diameter, have an internal organization identical with that of the mother sponge. They are enclosed in a layer of pinacoderm perforated by dermal pores. Under the pinacorderm there is a shallow subdermal space, which is in communication with the incurrent canals leading to the choanocyte chambers. The water sucked into these chambers proceeds into the excurrent canal system and emerges from the sponge through the oscular tube. Spicules projecting radially from the bud bear apical tufts of microscleres. The skeletal spicules of the buds, like their choanocyte chambers, are smaller than those in the mother sponge. The chambers expand to their mature size by choanocyte mitosis. Buds and sponges are colored green by intracellular symbiotic algae of the genusChlorella.     

Body structure of marine sponges

Summary The Mediterranean sponges Reniera mucosa, Haliclona mediterranea, Reniera fulva, Dendroxea lenis and Reniera sarai and the Caribbean species Callyspongia sp., Niphates digitalis, Niphates sp. and Amphimedon compressa are the subjects of this study of the arrangement of the choanocyte chambers between the canal systems and their relation to the mesenchymal tissue. The phylogenetic significance of the different organizational features is discussed.Dedicated to Prof. Dr. Norbert Weissenfels on the occasion of his 60th birthday     

The architecture of the canal systems of Petrosia ficiformis and Chondrosia reniformis studied by corrosion casts (Porifera,Demospongiae)

Summary The three-dimensional organization of the canal system in two sponge species, Petrosia ficiformis and Chondrosia reniformis, was studied using corrosion casts. Casts were made of live animals, in situ, and canal replicas were analzyed by scanning electron microscopy (SEM). In P. ficiformis the incurrent system consists of a superficial canal network giving rise to large radial canals, which ramify and anastomosize forming an internal web. Excurrent canals are arranged into modular ramified systems radiating from atrial cavities opening to the exterior. Main excurrent canals run at various depths within the sponge, even through the superficial incurrent network. Both incurrent and excurrent canal replicas show smooth, blind-ending capillaries. Some large incurrent canals merge with excurrent ones, thus bypassing choanocyte chambers. In C. reniformis there is a cortical collagen layer crossed by three-like incurrent canals, the twigs of which communicate with groups of inhalant pores. The stems of tree-like canals penetrate into the sponge medulla where they ramify and anastomosize to form a web. Main excurrent canals arise from large cloacal ducts leading to the oscular openings. They give rise to a sequence of branches intersecting the incurrent web. Both incurrent and excurrent canals have sharp, blind-ending capillaries. Morphometric data functions show that diameter scaling in canal branches is exponential in Petrosia and linear in Chondrosia. Structural differences and homologies between the two species are discussed.     

A New Flow-Regulating Cell Type in the Demosponge Tethya wilhelma – Functional Cellular Anatomy of a Leuconoid Canal System

Demosponges possess a leucon-type canal system which is characterized by a highly complex network of canal segments and choanocyte chambers. As sponges are sessile filter feeders, their aquiferous system plays an essential role in various fundamental physiological processes. Due to the morphological and architectural complexity of the canal system and the strong interdependence between flow conditions and anatomy, our understanding of fluid dynamics throughout leuconoid systems is patchy. This paper provides comprehensive morphometric data on the general architecture of the canal system, flow measurements and detailed cellular anatomical information to help fill in the gaps. We focus on the functional cellular anatomy of the aquiferous system and discuss all relevant cell types in the context of hydrodynamic and evolutionary constraints. Our analysis is based on the canal system of the tropical demosponge Tethya wilhelma, which we studied using scanning electron microscopy. We found a hitherto undescribed cell type, the reticuloapopylocyte, which is involved in flow regulation in the choanocyte chambers. It has a highly fenestrated, grid-like morphology and covers the apopylar opening. The minute opening of the reticuloapopylocyte occurs in an opened, intermediate and closed state. These states permit a gradual regulation of the total apopylar opening area. In this paper the three states are included in a theoretical study into flow conditions which aims to draw a link between functional cellular anatomy, the hydrodynamic situation and the regular body contractions seen in T. wilhelma. This provides a basis for new hypotheses regarding the function of bypass elements and the role of hydrostatic pressure in body contractions. Our study provides insights into the local and global flow conditions in the sponge canal system and thus enhances current understanding of related physiological processes.     

Choanocyte ultrastructure in Halisarca dujardini (Demospongiae,Halisarcida)

Understanding poriferan choanocyte ultrastructure is crucial if we are to unravel the steps of a putative evolutionary transition between choanoflagellate protists and early metazoans. Surprisingly, some aspects of choanocyte cytology still remain little investigated. This study of choanocyte ultrastructure in the halisarcid demosponge Halisarca dujardini revealed a combination of minor and major distinctive traits, some of them unknown in Porifera so far. Most significant features were 1) an asymmetrical periflagellar sleeve, 2) a battery of specialized intercellular junctions at the lateral cell surface complemented with an array of lateral interdigitations between adjacent choanocytes that provides a particular sealing system of the choanoderm, and 3) a unique, unexpectedly complex, basal apparatus. The basal apparatus consists of a basal body provided with a small basal foot and an intricate transverse skeleton of microtubules. An accessory centriole, which is not perpendicular to the basal body, is about 45°. In addition, a system of short striated rootlets (periodicity = 50–60 nm) arises from the proximal edge of the basal body and runs longitudinally to contact the nuclear apex. This is the first flagellar rootlet system ever found in a choanocyte. The accessory centriole, the rootlet system, and the nuclear apex are all encircled by a large Golgi apparatus, adding another distinctive feature to the choanocyte cytology. The set of distinct features discovered in the choanocyte of H. dujardini indicates that the ultrastructure of the poriferan choanocyte may vary substantially between sponge groups. It is necessary to improve understanding of such variation, as the cytological features of choanocytes are often coded as characters both for formulation of hypotheses on the origin of animals and inference of phylogenetic relationships at the base of the metazoan tree. J. Morphol., 2009. © 2008 Wiley‐Liss, Inc.     

A new type of central cell in the choanocyte chambers of Pellina fistulosa (Porifera,Demospongiae)

Central cells of a hitherto unknown type, forming a continuous, perforated layer at the level of the distal collar ends in each choanocyte chamber, have been found in the choanocyte chambers of Pellina fistulosa. The collars project through the pores of the perforated central cell layer. The spaces between the collar ends and between the collars and the cone cell ring in the apopyle region are sealed by the central cell cytoplasm. The latter represents an impermeable barrier for particulate material as well as for water and thus enhances the filtration efficiency by preventing a bypass of water and particles between the collar apices.     

Oogenesis and larval development of Scypha ciliata (Porifera,Calcarea)

Summary Scypha ciliata is a syconoid sponge. Its oocytes differentiate from choanocytes located near the apopyle of a flagellated chamber, and initially they remain in that location, in a trophic complex with neighbouring choanocytes. When this first growth phase is completed, the oocyte migrates to the periphery of the sponge. There it undergoes a second growth phase, in which it phagocytizes choanocytes and mesenchyme cells.Fertilization of the mature egg is assisted by a converted choanocyte, the sperm carrier cell. This cell penetrates the oocyte and transfers to it the sperm contained in a carriercell vacuole. No meiotic events have yet been observed.Cleavage is asynchronous, with holoblastic, approximately equal divisions. After the first cleavage steps the blastomeres often contain multiple nuclei. The single-layered blastoderm of the stomoblastula consists of many micromeres with flagella that project into the blastocoel, a few macromeres and four cruciform cells. There is no development of a follicle epithelium.The stomoblastula develops into the amphiblastula by inversion; with the assistance of the maternal choanocyte epithelium, the hollow sphere turns inside out, simultaneously moving out of the mesoderm and into the lumen of the adjacent flagellated chamber. In this process, the blastocoel of the stomoblastula is lost. The flagellated cells that form the wall of the amphiblastula now have their flagella extending outward; the amphiblastula also comprises four cruciform cells, macrogranular and agranular cells. The larval cavity of the amphiblastula is a newly formed structure.Abbreviations AB amphiblastula - AP apopyle - BC blastocoel - aC agranular cell - maC macrogranular cell - miC microgranular cell - CB crystalline body - CC central cavity - Ch choanocyte - fCh flat choanocyte - gCh granulate choanocyte - CM cell membrane - Co collar of choanocyte - CrC cruciform cell - DM dense material - EM electron micrograph - F flagellum - FC flagellated cell - FCm flagellated chamber - FL free larva - FV food vacuole - IR interior region - LC larval cavity - M mesenchyme - Ma macromere - MC mesenchyme cell - Mi micromere - N nucleus - Nu nucleolus - O opening - OC oocyte - P psudopodium - PC pinacocyte - PhM phase-contrast micrograph - Po pore - PP prosopyle - S sperm - SB stomoblastula - SC segmentation cavity - SCC sperm-carrier cell - SV sperm vacuole - lT large trophocyte - sT small trophocyte - V vacuole - VC vesicular cytoplasm - VM vacuole membrane     

Distinct histomorphology for growth arrest and digitate outgrowth in cultivated Haliclona sp. (Porifera: Demospongiae)

The use of sponges in biotechnological processes is limited by the supply problem, and sponge biomass production is becoming a current topic of research. The distinction between characteristics for growth and growth arrest is also important for environmental monitoring. In this study, we analyze the morphology of the digitate outgrowths from the sponge Haliclona sp . The sponge Haliclona sp . was successfully cultivated for 14 months in a closed system. The morphological characterization of growth arrest was performed after submitting explants to starvation‐stress for approximately 2 weeks, to correlate morphology with growth and growth arrest. The digitate outgrowth showed three distinct regions: mature (MR), transition (TR) and immature (IR). Our data suggest a growth developmental program, with collagen fascicles guiding axial growth in IR, followed by progressive development of choanocyte chambers and large aquiferous systems at the more mature proximal region (choanosome). The intercalation of choanocyte chambers and small aquiferous systems inside collagen fascicles previously originated at the IR region can be responsible for thickening expansion and conversion of the collagen fascicles into columnar choanosome in MR. The growth arrest after starvation‐stress assay showed morphological changes in the IR corroborating collagen in the extreme tip of the digitate outgrowth as an important role in guiding of axial growth of Haliclona sp . The identification of distinct morphologies for growth and growth arrest suggest a growth developmental program, and these data could be useful for further investigations addressing sponge biomass gain and environmental monitoring.     

Viviparous development in the Antarctic sponge Stylocordyla borealis Loven, 1868

The complete larval development of the deep-sea sponge Stylocordyla borealis (from eggs to young sponges) was followed in sponges from the Antarctic waters of Terra Nova Bay. S. borealis shows a viviparous strategy which leads to young complete sponges incubated in the mother body, with cortex, spicules and choanocyte chambers. This development can be considered a K-strategy, which is usually employed by deep-sea organisms and cold-water benthic invertebrates.     

The aquiferous systems of three marine demospongiae

The aquiferous systems of three common, coastal, marine Demospongiae, Halichondria panicea (Pallas), Haliclona permollis (Bowerbank) and Microciona Prolifera (Ellis and Solander), are analyzed by measurements of cross-sectional areas of conducting elements. The patterns in demosponges of extremely different organizational morphologies are found to be quantitatively similar. The porocyte nature of the ostia is established for all three species. Choanocyte chamber densities range from 1 to 1.8 × 10⁷ chambers ml⁻¹ with 57 to 95 choanocytes per chamber (means). Cross-sectional area of the intervillar space of the choanocyte collars is calculated to be 12 to 56 times the lateral surface area of the specimen. Velocities of water movement through specific elements of the aquiferous system are calculated from cross-sectional area data and measured oscular flow of Haliclona permollis. The calculated Reynolds numbers lie below the critical value and fluid flow is thus considered laminar throughout the aquiferous systems of these sponges.     

Solenoid: a new aquiferous system to Porifera

The aquiferous system is an essential character of poriferans and supports their monophyly. Within the Calcarea, this system displays its greatest variety and traditionally is classified as: asconoid, syconoid, sylleibid, and leuconoid. Species of Leucascus, however, present a different type of aquiferous system composed of anastomosed (interconnected) choanocyte tubes and have an atrium lacking choanoderm. There is such confusion about the classification of the aquiferous system of Leucascus that, depending on the author, it has been classified as asconoid, syconoid, or leuconoid. Therefore, in the present work, we describe a new type of aquiferous system for Leucascus: the solenoid aquiferous system. This new aquiferous system is defined by the presence of anastomosed tubes internally lined by choanocytes and atrium without choanoderm. Although no deep phylogenetic significance has been attributed to the aquiferous system, the solenoid system raises important evolutionary questions about the variety of systems found among the poriferans.     

The aquiferous system of two <Emphasis Type="Italic">Oceanapia</Emphasis> species (Porifera,Demospongiae) studied by corrosion casts

The aquiferous systems of two Indopacific Oceanapia species (Oceanapiidae) were studied by corrosion casts: O. amboinensis living in shallow lagoons and O. fistulosa living at the base of the reef slope. Both species show a massive, entirely buried body, emerging from the sediment only by long, completely close fistules. Particularly in O. fistulosa the corrosion casts revealed a complex, grape-like structure of the choanosome organised in anatomical and functional units composed by an incurrent web whose anastomosed meshes are crossed by a central excurrent canal. A system of thin canals connects the two systems giving rise to an area of choanocyte chambers. The corrosion casts revealed that in both species incurrent water penetrates into the sponge body by the fistules and that it is expelled through specialised structures buried in the sediment. This observation is in accordance with field experiments performed on O. fistulosa. In some specimens of this species, a solution of china ink injected into plastic bags enveloping the external fistules was observed, after waiting for a while, to flow through the buried structures.     

Ultrastructure of the lycophora larva of Gyrocotyle urna (Cestoda,Gyrocotylidea)

Summary The ultrastructure of the protonephridial system of the lycophore larva of Gyrocotyle urna Grube and Wagener, 1852, is described. It consists of six terminal cells, at least two proximal canal cells, two distal canal cells and two nephridiopore cells. The terminal cells and the proximal canal cell build up the filtration weir with its two circles of weir rods. The proximal canal cell constitutes a solid, hollow cylinder without a cell gap and desmosome. The distal canal cell is characterized by a strong reduction of the canal lumen by irregularly shaped microvilli. The nephridiopore region is formed by a nephridiopore cell; its cell body is located at some distance proximally within the larva. The connection among different canal cells is brought about by septate desmosomes. Morphological, evolutionary and functional aspects of the protonephridial system within Platyhelminthes are discussed. The structure of the proximal canal cells without a desmosome is considered an autapomorphy of Cestoda.Abbreviations ci cilia of the terminal cell - Co distal canal cell - col lumen of the distal canal cell - Ep epidermis - er outer rods of the filtration weir - il inner leptotriches - ir inner rods of the filtration weir - ld lipid droplets - mt microtubule - mv microvilli - Nc nephridiopore cell - Ne neodermis anlage cells - nu nucleus - pC proximal canal cell - ro ciliary rootlets - sd septate desmosome - Tc terminal cell     

Lateral line morphology and cranial osteology of the Rubynose Brotula,Cataetyx rubrirostris

Cranial osteology, canal neuromast distribution, superficial neuromast distribution and innervation, and cephalic pore structure were studied in cleared and stained specimens of the deep sea brotulid Cataetyx rubrirostris. The cranial bone structure of C. rubrirostris is similar to other brotulids (Dicrolene sp.) and zoarcids (Zoarces sp.), except for an unusual amount of overlapping of the bones surrounding the cranial vault. The superficial neuromasts are innervated by the anterodorsal, anteroventral, middle and posterior lateral line nerves and are organized similarly to those of the blind ophidioid cave fish Typhliasina pearsei. The cephalic pores open into a widened lateral line canal system. The canal is compartmentalized into a series of neuromast‐containing chambers that probably amplify signals received by the system. J. Morphol. 241:265–274, 1999. © 1999 Wiley‐Liss, Inc.     

Three‐dimensional virtual histology of silurian osteostracan scales revealed by synchrotron radiation microtomography

We used propagation phase contrast X‐ray synchrotron microtomography to study the three‐dimensional (3D) histology of scales of two osteostracans, Tremataspis and Oeselaspis, members of a jawless vertebrate group often cited as the sister group of jawed vertebrates. 3D‐models of the canal systems and other internal structures are assembled based on the virtual thin section datasets and compared with previous models based on real thin sections. The primary homology framework of the canal systems in the two taxa is revised and new histological details are revealed based on the results of this work. There is no separation of vascular canals and lower mesh canals in the Tremataspis scale, contrary to previous results. The secondary upper mesh canals have a limited distribution to the anterior region of the Tremataspis scale. The upper and lower mesh canal systems of Tremataspis have different geometries, inferred to reflect different developmental origins: we interpret the upper system as a probable epithelial invagination, the lower system as entirely vascular. Oeselaspis has no equivalent of the upper mesh canal system. The upper mesh canal system of Tremataspis may have been sensory in function. In Oeselaspis, numerous polyp‐shaped structures opening from the canal system onto the surface of the scale resemble the innervation tracts for neuromast organs. The growth of the Oeselaspis scale proceeds by addition of small odontodes containing unmineralized lacunae, which may further mineralize and become more compact. Our results highlight that 3D‐histological investigation on scales and other dermal skeletons of osteostracans is necessary to fully appreciate the diversity of skeletal histologies in the group. Traditional 3D‐models based on thin sections alone are not reliable and should no longer be used as the basis for homology assessments or functional hypotheses. J. Morphol. 276:873–888, 2015. © 2015 Wiley Periodicals, Inc.     

Morphology of the Mechanosensory Lateral Line System in Elasmobranch Fishes: Ecological and Behavioral Considerations

The relationship between morphology of the mechanosensory lateral line system and behavior is essentially unknown in elasmobranch fishes. Gross anatomy and spatial distribution of different peripheral lateral line components were examined in several batoids (Raja eglanteria, Narcine brasiliensis, Gymnura micrura, and Dasyatis sabina) and a bonnethead shark, Sphyrna tiburo, and are interpreted to infer possible behavioral functions for superficial neuromasts, canals, and vesicles of Savi in these species. Narcine brasiliensis has canals on the dorsal surface with 1 pore per tubule branch, lacks a ventral canal system, and has 8–10 vesicles of Savi in bilateral rows on the dorsal rostrum and numerous vesicles ( = 65 ± 6 SD per side) on the ventral rostrum. Raja eglanteria has superficial neuromasts in bilateral rows along the dorsal body midline and tail, a pair anterior to each endolymphatic pore, and a row of 5–6 between the infraorbital canal and eye. Raja eglanteria also has dorsal canals with 1 pore per tubule branch, pored and non-pored canals on the ventral surface, and lacks a ventral subpleural loop. Gymnura micrura has a pored dorsal canal system with extensive branch patterns, a pored ventral hyomandibular canal, and non-pored canal sections around the mouth. Dasyatis sabina has more canal pores on the dorsal body surface, but more canal neuromasts and greater diameter canals on the ventral surface. Sphyrna tiburo has primarily pored canals on both the dorsal and ventral surfaces of the head, as well as the posterior lateral line canal along the lateral body surface. Based upon these morphological data, pored canals on the dorsal body and tail of elasmobranchs are best positioned to detect water movements across the body surface generated by currents, predators, conspecifics, or distortions in the animal's flow field while swimming. In addition, pored canals on the ventral surface likely also detect water movements generated by prey. Superficial neuromasts are protected from stimulation caused by forward swimming motion by their position at the base of papillar grooves, and may detect water flow produced by currents, prey, predators, or conspecifics. Ventral non-pored canals and vesicles of Savi, which are found in benthic batoids, likely function as tactile or vibration receptors that encode displacements of the skin surface caused by prey, the substrate, or conspecifics. This mechanotactile mechanism is supported by the presence of compliant canal walls, neuromasts that are enclosed in wide diameter canals, and the presence of hair cells in neuromasts that are polarized both parallel to and nearly perpendicular to the canal axis in D. sabina. The mechanotactile, schooling, and mechanosensory parallel processing hypotheses are proposed as future directions to address the relationships between morphology and physiology of the mechanosensory lateral line system and behavior in elasmobranch fishes.     

Cephalic lateral line canal system of the golden venus chub, <Emphasis Type="Italic">Hemigrammocypris rasborella</Emphasis> (Teleostei: Cypriniformes)

We investigated the cephalic lateral line canal system of the golden venus chub, Hemigrammocypris rasborella. The cephalic lateral line canal system consists of the infraorbital canal (IOC), the preopercular canal (POC), the mandibular canal (MC), the supraorbital canal (SOC), the temporal canal (TC), and the supratemporal canal (STC), and is characterized by the following pedomorphic features: disjunction of IOC and SOC, of TC and POC, and of POC and MC. We also discuss the phylogenetic significance of the cephalic lateral line canal system of H. rasborella.