Polykristalliner Calcit bei Seeigeln (Echinodermata,Echinoidea)

Zusammenfassung Es wurde erstmals für Echinodermen primär polykristalliner Calcit nachgewiesen, und zwar im Cortex der Primärstacheln der Cidaridae, dem sekundären Zahnskelet von Clypeaster und in den akzessorischen Kalkstrukturen, die im Kauabschnitt die Furche der Diadematiden-Zähne ausfüllen. Es gibt bei anderen Seeigelfamilien keine Bildungen, die dem Cidariden-Cortex oder den akzessorischen Kalkstrukturen der Diadematiden homolog sind. Das polykristalline sekundäre Zahnskelet von Clypeaster ist dagegen dem monokristallinen sekundären Zahnskelet anderer Seeigel homolog.Der Mg-Gehalt des Calcits liegt in den feinkristallinen Zonen (mit Ausnahme des Cortex) im allgemeinen höher; die höchsten Werte finden sich in den Steinteilen der Zähne, gleichgültig ob das sekundäre Zahnskelet mono- oder polykristallin ist.Polykristalline Teile sind im allgemeinen härter als monokristalline Teile. Die Steinteile der Seeigelzähne sind die härtesten Skeletteile von Echinodermen überhaupt; ihre VickersHärte übertrifft weit diejenige von solidem Calcit. Im Steinteil ist das feinkristalline Gefüge von Calcit eng mit organischer Matrix verbunden, und es wird vermutet, daß darauf die besonders hohe Härte der Steinteile beruht.

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Polycristalline calcite in sea urchins

Summary For the first time primary polycrystalline calcite in Echinoderms is shown in the cortex of primary spines of Cidaridae, in the secondary tooth skeleton of Clypeaster and in the accessory calcareous structures filling the crevice fold in the chewing areas of Diadematoidae teeth. Other Echinoid families lack formations homologous to the cortex of Cidaridae and accessory calcareous structures of Diadematoidae. On the other hand the polycrystalline secondary tooth skeleton of Clypeaster is homologous to the monocrystalline one of the other sea urchins.With the exception of cortex the Mg-content in calcite—analyzed by microprobe and X-ray powder method—is generally greater in macrocrystalline parts. The highest Mg-contents are found in the stone parts of teeth irrespective of whether the secondary tooth skeleton is monocrystalline or polycrystalline.Polycrystalline parts are usually harder than monocrystalline ones. The stone parts of Echinoid teeth are the hardest skeleton parts of Echinoderms on the whole; their hardness is much greater than that of solid calcite. It is supposed that the strong interlacing of the microcrystalline calcite and organic matter causes the enormous hardness of the stone part.

Mit Unterstützung durch die Deutsche Forschungsgemeinschaft.     

Potential for paired vestibules in plutei (Echinodermata, Echinoidea)

Abstract. Echinoids are generally thought to develop only a single ectodermal invagination (V1), which expands into a vestibule and contributes to the formation of the juvenile rudiment on the left side of the larval body. However, in 2 echinoid species, Strongylocentrotus purpuratus and Dendraster excentricus, a second invagination (V2) was observed to form on the right side of the larval body exactly opposite V1. Formation of the invaginations was followed and characterized using light microscopy and antibodies directed against the S2amide echinoderm neuropeptide as a morphological tag. V2 consistently formed later than V1, and appeared capable of contributing to the formation of a second rudiment when it received the appropriate mesodermal influence from contact with a hydrocoel. Otherwise, in both species, V2 formed a skeletal element that has not been previously described in this location. Because of the location, shape, and timing of its formation, V2, like V1, appears to have the innate ability of the larval ectoderm to contribute to juvenile rudiment formation.     

Evolution of a novel muscle design in sea urchins (Echinodermata: Echinoidea)

The sea urchin (Echinodermata: Echinoidea) masticatory apparatus, or Aristotle's lantern, is a complex structure composed of numerous hard and soft components. The lantern is powered by various paired and unpaired muscle groups. We describe how one set of these muscles, the lantern protractor muscles, has evolved a specialized morphology. This morphology is characterized by the formation of adaxially-facing lobes perpendicular to the main orientation of the muscle, giving the protractor a frilled aspect in horizontal section. Histological and ultrastructural analyses show that the microstructure of frilled muscles is largely identical to that of conventional, flat muscles. Measurements of muscle dimensions in equally-sized specimens demonstrate that the frilled muscle design, in comparison to that of the flat muscle type, considerably increases muscle volume as well as the muscle's surface directed towards the interradial cavity, a compartment of the peripharyngeal coelom. Scanning electron microscopical observations reveal that the insertions of frilled and flat protractor muscles result in characteristic muscle scars on the stereom, reflecting the shapes of individual muscles. Our comparative study of 49 derived "regular" echinoid species using magnetic resonance imaging (MRI) shows that frilled protractor muscles are found only in taxa belonging to the families Toxopneustidae, Echinometridae, and Strongylocentrotidae. The onset of lobe formation during ontogenesis varies between species of these three families. Because frilled protractor muscles are best observed in situ, the application of a non-invasive imaging technique was crucial for the unequivocal identification of this morphological character on a large scale. Although it is currently possible only to speculate on the functional advantages which the frilled muscle morphology might confer, our study forms the anatomical and evolutionary framework for future analyses of this unusual muscle design among sea urchins.     

Ultrastructural investigation of matrix-mediated biomineralization in echinoids (Echinodermata,Echinoida)

Summary The formation of echinoderm endoskeletons is studied using echinoid teeth as an example. Echinoid teeth grow rapidly. They consist of many calcareous elements each produced by syncytial odontoblasts. The calcification process in echinoderms needs (1) syncytial sclerocytes or odontoblasts, (2) a spacious vacuolar cavity within this syncytium, (3) an organic matrix coat in the cavity. As long as calcite is deposited, the matrix does not touch the interior face of the syncytium. The cooperation between syncytium, interior cavity and matrix coat during the mineralization process is discussed. The proposed hypothesis applies to the formation of larval skeletons, echinoderm ossicles and echinoid teeth.When calcite deposition ceases the syncytium largely splits up into filiform processes, and the skeleton is partly exposed to the extracellular space. However, the syncytium is able to reform a continuous sheath and an equivalent of the cavity and may renew calcite deposition.The tooth odontoblasts come from an apical cluster of proliferative cells, each possessing a cilium. The cilium is lost when the cell becomes a true odontoblast. This suggests that cilia are primitive features of echinoderm cells. The second step in calcification involves the odontoblasts giving rise to calcareous discs which unite the hitherto single tooth elements. During this process the odontoblasts immure themselves. The structures necessary for calcification are maintained until the end of the process.The mineralizing matrix is EDTA-soluble. The applied method preserves the matrix coating the calcite but more is probably incorporated into the mineral phase and dissolved with the calcite.Abbreviations A adhesive point (LNC) - B adaxial bag - bb basal body (ci) - CA calcareous deposits - cb cytoplasmic bladder (cp) - ce centriole - ci cilium - cp cable-like cell process - cv condensing vacuole - dp distal processes (sh) - E epithelium of the tooth - ex extracellular space - f extracellular fibrils - ga gasket (sh) - ic interior cavity - L lamellae (LNC) - LNC lamellae needle complex - m mitochondrium - mc matrix coat - MF main fold (P) - MI mitosis - mt microtubules - N nucleus - O odontoblast - P primary plate - Ph phagocyte - PR proliferative cell - pr prism - rb reserve body - RER rough endopl. reticulum - rl rootlet (ci) - RY relatively youngest plate - s satellite (bb, ce) - sh synplasmic sheath (O) - SP secondary plate - sv smooth-walled vesicle - TF transversal fold (P) - U umbo (P) - v Golgi vesicle - Y youngest tooth element     

Phagostimulation of Lytechinus variegatus (Lamarck) (Echinodermata: Echinoidea)

Artificial agarose foods containing either starch, glycogen, carragheenan, alginate, casein, gelatin, albumin, or peptones at a concentration of 1% dry weight were not phagostimulatory. Artificial foods containing sugars, especially galactose, at a concentration of 0.005 M were phagostimulatory. Of the common amino acids, only L‐phenylalanine was phagostimulatory at this concentration. The initiation of feeding postures (movement of the spines, tube feet, peristome, and Aristotle's lantern) of Lytechinus variegatus was little affected by the concentration of plant extracts. Many aspects of this behavior were attributable to physical rather than chemical stimulation. Feeding rates of L. variegatus upon artificial agarose foods containing either Thalassia testudinum or Gracilaria verrucosa were directly dependent upon plant tissue concentration. All artificial agarose food‐items were initially moved to the mouth and fed upon, however the time spent feeding was directly dependent upon plant type and tissue concentration. Retention of the artificial food‐items upon the test after feeding was not influenced by plant type or tissue concentration. Differences in consumption of foods by L. variegatus are determined by the degree of persistence in feeding upon encountered items and not be selection before initiating feeding or by retention of food‐items for subsequent feedings.     

Behavioral ecology of Strongylocentrotus droebachiensis (Muller) (Echinodermata: Echinoidea)

Summary Field observation indicated that extensive aggregation behavior occurs in Strongylocentrotus droebachiensis, an important echinoid herbivore in rocky subtidal zones of northern New England. The relationship of this behavior to the behavioral ecology of this species was investigated. Laboratory studies indicated that urchins are chemically sensitive to the stimuli of food and other urchins. Quantitative field experiments with tagged animals and transects showed the existence of two types of aggregations: feeding and nonfeeding, which appear to be pervasive phenomena in the biology of S. droebachiensis.It is concluded that, depending upon their hunger state, urchins shuttle between feeding aggregations in exposed locations and non-feeding aggregations located in relatively sheltered areas (where they spend most of their time). Individuals of S. droebachiensis aggregate actively through response to other urchins in the form of chemotaxis. The aggregative behavior may be maintained for its selective value in terms of defense from predation and, to lesser extents, facilitation of feeding and breeding.     

Mechanical properties and structure of the spine-joint central ligament of the sea urchin, Diadema setosum (Echinodermata, Echinoidea)

In the regular echinoid Diadema setosum Leske the central ligament that connects the spine to its tubercle is mainly composed of closely-packed collagen fibres which are arranged in the longitudinal axis of the ligament. The mechanical properties of the ligament are quite different from ligament to ligament: the viscosity determined by creep tests ranges from 0·02 to 6 GPa·s. The viscosity of a ligament changes reversibly in response to stimulation. Adrenaline and noradrenaline (10^-6-10^-3 M) decrease the viscosity. Acetylcholine (10^-8-10^-3 M), artificial sea water containing high (100 MM) potassium concentration, and electrical stimulation increase the viscosity. As the ligament contains no muscle cells in it, the viscosity change cannot be attributable to muscle activities. The functional significance of having a central ligament with variable and controllable viscosity is that it binds the spine base onto the articulation surface of the tubercle, so as to reduce the possibility of spine dislocation, while being flexible enough to allow the spine considerable freedom of movement when necessary.     

Domes,arches and urchins: The skeletal architecture of echinoids (Echinodermata)

Summary A combination of simple membrane theory and statical analysis has been used to determine how stresses are carried in echinoid skeletons. Sutures oriented circumferentially are subject principally to compression. Those forming radial zig-zags are subject to compression near the apex and tension near the ambitus. Radial and circumferential sutures in Eucidaris are equally bound with collagen fibers but in Diadema, Tripneustes, Psammechinus, Arbacia and other regular echinoids, most radial sutures are more heavily bound, and thus stronger in tension. Psammechinus, Tripneustes and several other echinoids have radial sutures thickened by ribs which increase the area of interlocking trabeculae. Ribs also increase flexural stiffness and carry a greater proportion of the stress. Further, ribs effectively draw stress from weaker areas pierced by podial pores, and increase the total load which can be sustained.Allometry indicates that regular echinoids become relatively higher at the apex as size increases, thus reducing ambital stresses. Some spatangoids with very high domes (eg Agassizia) maintain isometry, but others (eg Meoma) become flatter with size. Both holectypoids (Echinoneus) and cassiduloids (Apatopygus) maintain a constant height to diameter relationship. Flattening, and consequently ambital tensile stress, is greatest in the clypeasteroids. In this group the formation of internal buttresses which preferentially carry stress, reaches maximum development. A notable exception, however, is the high domed Clypeaster rosaceus.In this analysis it was assumed that local buckling or bending does not occur. The test of some echinoids (e.g. Diadematoida) have relatively wide sutures swathed in collagen, which allows local deformation. Others (e.g. Arbacia) have rigid sutures with reduced collagen. In Psammechinus and other members of the Order Echinoida, in addition to rib formation, inner and outer surface trabeculae are thickened so that the individual plates are stiffened. Some spatangoids (Meoma, Paleopneustes) have extensive sutural collagen, but the cassiduloid Apatopygus has collagen confined to junctions of sutures, and elsewhere the joints are strengthened and stiffened by fusion of trabeculae. Fusion of surface trabeculae is almost complete in the holectypoid, Echinoneus, and the sutures are obscured.     

Structure and function of clypeasteroid miliary spines (Echinodermata,Echinoides)

Summary Histological and ultrastructural techniques have been used to describe the functional morphology of clypeasteroid miliary spines, with special reference to their supposed mucus-secreting role. Mucus cells were not found in the miliary spines of any members of the Arachnoididae, Fibulariidae, Laganidae, Echinarachniidae, Dendrasteridae, Astriclypeidae, or Mellitidae examined in this study. Only members of the Clypeasteridae have mucus-secreting cells in these spines. Characteristics of the skeleton, ultrastructure of the nervous system, and histology of the musculature and epithelia of the base, shaft and tip are also discussed. Miliary spines have two bands of cilia running along the entire length of opposite sides of the shaft. The geometric packing of cilium-bearing cells in these bands is described for the first time, as is the remarkable form of the sacs found at the tips of dendrasterid, astriclypeid, and mellitid miliary spines. These sacs are definitely not mucous sacs, as previously described, but are balloons of single-celled epithelium internally tethered to the skeletal tip by copious quantities of collagenous connective tissue. Miliary spines prevent obstruction of aboral nutritive and ventilatory ciliary currents caused by substrate particles falling to the test surface during burrowing. They do this in two ways: (1) they help generate ciliary currents that sweep finer material off the test, and (2) they contribute to the formation of a spine canopy that mechanically blocks larger particles from falling between the spines. Members of the Clypeasteridae secrete an interspine mucous tent that traps potentially clogging material. The miliary spine sacs of sand dollars are deformable space-fillers that plug holes between primary spines in the aboral canopy, even as the spines rock on their tubercles to push sand backwards over the test. Allometry of spines from Echinarachnius parma suggests that aboral military spines and club-shaped spines exhibit co-ordinated growth that maintains the aboral canopy throughout post-metamorphic ontogeny, and that aboral spins have an overall lower growth rate than spines on the oral surface.     

Echinoids (Echinodermata: Echinoidea) from the Gulf of Mexico

The echinoid fauna of the Gulf of Mexico collected during three research cruises (20-1260 m depth) was surveyed from samples were taken at 43 stations. A total of 190 individuals were identified (eight orders, 11 families, 15 genera and 18 species). Six species are new records for the Gulf of Mexico: Stylocidaris lineata, Phormosoma placenta placenta, Plesiodiadema antillarum, Plethotaenia spatangoides, Brissopsis atlantica and Hypselaster limicolus. This adds to the little information available on the echinoid fauna of Tamaulipas, Veracruz, Tabasco, Campeche and Yucatan states in Mexico.     

Thyroid hormones determine developmental mode in sand dollars (Echinodermata: Echinoidea)

Evolutionary transitions in larval nutritional mode have occurred on numerous occasions independently in many marine invertebrate phyla. Although the evolutionary transition from feeding to nonfeeding development has received considerable attention through both experimental and theoretical studies, mechanisms underlying the change in life history remain poorly understood. Facultative feeding larvae (larvae that can feed but will complete metamorphosis without food) presumably represent an intermediate developmental mode between obligate feeding and nonfeeding. Here we show that an obligatorily feeding larva can be transformed into a facultative feeding larva when exposed to the thyroid hormone thyroxine. We report that larvae of the subtropical sand dollar Leodia sexiesperforata (Echinodermata: Echinoidea) completed metamorphosis without exogenous food when treated with thyroxine, whereas the starved controls (no thyroxine added) did not. Leodia sexiesperforata juveniles from the thyroxine treatment were viable after metamorphosis but were significantly smaller and contained less energy than sibling juveniles reared with exogenous food. In a second starvation experiment, using an L. sexiesperforata female whose eggs were substantially larger than in the first experiment (202+/-5 vs. 187+/-5 microm), a small percentage of starved L. sexiesperforata larvae completed metamorphosis in the absence of food. Still, thyroxine-treated larvae in this experiment completed metamorphosis faster and in much higher numbers than in the starved controls. Furthermore, starved larvae of the sand dollar Mellita tenuis, which developed from much smaller eggs (100+/-2 microm), did not complete metamorphosis either with or without excess thyroxine. Based on these data, and from recent experiments with other echinoids, we hypothesize that thyroxine plays a major role in echinoderm metamorphosis and the evolution of life history transitions in this group. We discuss our results in the context of current life history models for marine invertebrates, emphasizing the role of egg size, juvenile size, and endogenous hormone production for the evolution of nonfeeding larval development.     

Abundance of Diadema antillarum (Echinodermata: Echinoidea) in the coasts of Venezuela

Diadema antillarum is a shallow-water sea-urchin from the tropical Atlantic whose populations almost disappeared in 1983-84 because of widespread mortalities which reached 87-100 %. In Venezuela, urchin population densities before the mortality event were comparable to those of other Caribbean regions; however, later abundancies remain unknown. The objectives of this study were to evaluate the recent densities of certain D. antillarum populations along the Venezuelan coasts and compare the densities at the Parque Nacional Mochima before and after the mortality. At each location urchin densities were determined by means of transects using 1 m2-quadrats as sampling units. The highest mean densities were observed at the sites on the central coast: Ensenada de Oricao, 0.28 ind/m2 (2002) and 1.05 ind/m2 (2003), and Chichiriviche de la Costa, 0.84 ind/m2 (2002) and 0.74 ind/m2 (2003). In Mochima, the mean density before the mortality for D. antillarum oscillated between 0.28 and 4 ind/m2, after the mortality event the mean density varied between 0.15 ind/m2 (2000) and 0.47 ind/m2 (2000). The populations of D. antillarum studied at Parque Nacional Morrocoy and Refugio de Fauna Silvestre Cuare showed highest densities at Playuela (0.43 ind/m2) and Cayo Sur (0.95 ind/m2) respectively, whereas other sites showed densities below 0.1 ind/m2. The density registered at Playuela in 2003 is lower than that reported before the mortality event (0.58-3.64 ind/m2). The density for Parque Nacional Archipiélago de Los Roques, specifically for the Arrecife de Herradura remained constant between 2002 and 2003 with values between 0.22-0.23 ind/m2 respectively. To conclude, the sea urchin abundancies observed at most of the Venezuelan coastal sites that we studied were higher than those reported for other areas of the northern Caribbean, even though the values have not yet returned to those preceding the 1984 mass-mortality event, due to the slow recovery of the populations.     

Histology of the globiferous pedicellariae of Psammechinus miliaris (Echinodermata: Echinoidea)

The general morphology, function and histology of the globiferous pedicallariae of the regular sea-urchin, Psammechinus miliaris are described. There is a ganglion under the sensory epidermis on the inner surface of each jaw. This is the first discovery of a ganglion in any echinoid.
The regeneration of globiferous pedicellariae in laboratory conditions takes 25 to 40 days.     

Siphons and siphonal grooves in the digestive systems of the Echinoidea (Echinodermata)

Here we use histological sections and, to a limited extent, scanning electron microscopy to ascertain whether siphons or siphonal grooves parallel the stomach of five sea urchin species representing five higher taxa in the Echinoidea. We find a siphonal groove in Centrostephanus coronatus of the Diadematidae and in Caenopedina diomedeae of the Pedinidae; we find a siphon in Tromikosoma panamense of the Echinothurioida and in Strongylocentrotus purpuratus of the Camarodonta; and we find a hemisiphon in Aspidodiadema hawaiiense of the Aspidodiadematidae (it is currently unsettled whether this last group belongs in the Pedinoida or the Diadematoida). Several recent accounts of echinoid gut anatomy have claimed that species in the Echinothurioida have a siphonal groove instead of a siphon and that species in the Diadematidae and Pedinidae have siphons instead of siphonal grooves. The present work shows that these claims are mistakes and confirms that Holland and Ghiselin (1970) correctly described the distribution of siphons and siphonal grooves in the Echinoidea.     

Pourtalesiid sea urchins (Echinodermata: Echinoidea) of the northern Mid-Atlantic Ridge

Five species of deep-sea pourtalesiid echinoids were collected by the RV G.O. Sars MAR-ECO expedition to the northern part of the Mid-Atlantic Ridge: Solenocystis imitans new genus, new species, Echinosigra phiale (Thomson, 1872), Echinosigra (Echinogutta) fabrefacta Mironov, 1974, and two unidentified species of Pourtalesia. Three types of plastron plating are distinguished within the family Pourtalesiidae; in Solenocystis new genus the plating is similar to that in Spatagocystis, a genus known only from the deep-sea Antarctic. Each of three groups with distinctive plastron plating shows a continuous morphological gradation from a less elongated test without subanal rostrum to an extraordinarily elongated test with large rostrum and wide subanal fasciole; the ratio width/length of test varies from 0.86 to 0.12. Some morphological characters of the pourtalesiids are regarded as adaptations to burrowing and appear to be correlated with test elongation. The family is represented in the northern Atlantic only by 'advanced' genera (Echinosigra, Pourtalesia and Solenocystis) with very elongated tests.     

Phylogeny of Early Cretaceous spatangoids (Echinodermata: Echinoidea) and taxonomic implications

A phylogenetic analysis of 36 species provides a test for the taxonomy and the history of Early Cretaceous spatangoids. Most taxonomic units from genera to suborders are consistent with the proposed phylogenetic framework. We retain Hemiasterina, Micrasterina, Hemiasteridae, Schizasteridae, Hemiaster , Heteraster , Mecaster , and Periaster as original monophyletic groups. However, all of these clades originate without the classical apomorphies normally ascribed to them. We suggest a revision of their diagnoses and of the generic attributions of basal species. Some ill-defined, 'primitive', and paraphyletic taxa are recognised: Toxaster , Epiaster , Palhemiaster , and Toxasteridae. Even if they do not have phylogenetic meaning, they are retained here, pending a more complete revision.