Skeletal Ultrastructures in some Cerioporine Cyclostome Bryozoans

Abstract The ultrastructure of the calcareous skeleton is described in nine species of Recent cyclostome bryozoans belonging to the suborder Cerioporina. Two species of Heteropora have interior zooecial walls comprising a granular precursory layer followed by a thick layer of transverse fibres and a subordinate foliated fabric with, in mature proximal walls, a semi-nacreous layer. The remaining seven species have interior walls with no transverse fibres and instead predominantly comprise a distally-imbricated, regularly foliated fabric overlying a granular precursory layer. Older, proximal surfaces often have abundant screw dislocations, but true semi-nacre is absent. Basal walls comprise an outer finely granular precursory fabric and planar spherulitic layer, succeeded by the same ultrastructural succession seen in the interior zooecial walls of the respective groups. Exterior walled diaphragms, peristomes and gonozooids similarly comprise an external fabric of planar spherulitic calcite, lined internally by the predominant fabric seen in the interior walls. Ultrastructurally, therefore, cerioporines may be split into two groups with different fabric suites, the first resembling cinctiporids and many tubuliporines in having interior walls with fabrics of transverse fibres, foliated crystallites and semi-nacre; and the second resembling the rectangulates Lichenopora and Disporella in having interior walls comprising only the foliated fabric. These findings support the close phylogenetic relationship between cerioporines and other cyclostomes but suggest that the cerioporines may constitute either a diphyletic or a paraphyletic group.     

Skeletal Ultrastructure in Some Articulate Cyclostome Bryozoans

The ultrastructure of the calcareous skeleton is described in 11 species of articulate cyclostome bryozoans with elastic joints. Ten species have interior walls comprising semi-nacreous and pseudofoliated fabrics without a precursory granular layer. Exterior walls consist of outer, finely granular and planar spherulitic layers, succeeded by semi-nacreous and pseudofoliated fabrics like those of interior walls. Outer fabrics are calcified as longitudinal strips, each corresponding to a planar sphcrulitic unit. Articulation surfaces comprise ring diaphragms of very fine granular fabric with concentric laminations. The semi-nacre of walls adjacent to ring diaphragms contains minute holes. Crisulipora occidentalis is unique in having interior walls of transverse fibres succeeded by pseudofoliated fabric, articulation surfaces festooned with deep pits but lacking well-differentiated ring diaphragms, and pseudopores containing sieve-like closure plates. The ultrastructure of most articulates resembles tubuliporine cyclostomes with dominantly semi-nacreous walls, although the lack of precursory granular fabric in the interior walls and the presence of subcircular tablets of semi-nacre (without six-fold sectoring) may be peculiar to articulates. In contrast, Crisulipora is more similar to other tubuliporines with transverse fibres. evidence which, together with other skeletal characters, suggests that Crisulipora evolved jointing independently of the rest of the articulate cyclostomes.     

Skeletal Ultrastructure in some Cyclostome Bryozoans of the Family Lichenoporidae

The ultrastructure of the skeleton is described in six species of lichenoporid cyclostome bryozoans using field emission SEM. Both interior walls (vertical, interzooidal walls, and brood chamber roofs and floors) and exterior walls (basal walls) are initially secreted as tiny wedge-shaped crystallites without a strong preferred orientation. These are seeded directly onto pre-existing crystallites in the case of interior walls, but onto the organic cuticle in exterior walls, the bases of the crystallites forming a tightly packed mosaic against the cuticle. With growth the wedges become longer, broader and relatively flatter, developing into platey crystallites. These crystallites grow predominantly distally (i.e. parallel to wall growth direction) and are closely imbricated in a foliated fabric. Local disruptions to this pattern occur, especially in association with crystallite division along “divergent zones”. The pattern also breaks down in old walls where crystallite growing edges become less evident and imbrication is poorly developed. Although conforming to this general model, some differences exist between species of lichenoporids, and in the patterns found in different parts of the skeleton (e.g. apertural spines). Lichenoporid ultrastructure differs from that of both cinctiporid and hornerid cyclostomes: notably, lichenoporids lack the layer of transverse fibres found in cinctiporids, and their predominant distal growth direction of crystallites contrasts with the proximal direction found in hornerids.     

Skeletal Ultrastructure of the Early Astogenetic Stages of Some Cyclostome Bryozoans

The calcific protoecia of cyclostome bryozoans have remarkably uniform skeletal ultrastructure in three suborders (Tubuliporina, Cancellata and Rectangulata). The basal wall or floor has a fine outer granular layer succeeded by planar spherulitic fabric, internally lined by irregular semi-nacre. The roof of the disc comprises an outer granular layer with an inner lining of semi-nacre continuous with that of the floor; planar spherulitic fabric is absent. Growth of the floor of the disc is initiated around the circular outer rim and continues centripetally to the centre; the inner lining has no prevailing growth direction. The gently domed roof also initiates around the outer rim, and grows in strips, which grow toward the centre, then distally toward the distal tube of the ancestrula which has a fully adult ultrastructure. Protoecia] ultrastructure is independent of adult ultrastructure. The uniformity of skeletal ultrastructure in cyclostomes corresponds with the close similarity of larvae and post-settlement metamorphosis in the order. The fabric suite of the protoecium resembles the skeletal ultrastructure of Palaeozoic stenolaemates. The primitive fabric condition is retained by some tubuliporines and cancellates. Complex multilayered fabric suites may have evolved in the Mesozoic by addition of new fabric types.     

Skeletal ultrastructure and phylogeny of cyclostome bryozoans

Recent research on the ultrastructure of the calcareous skeleton in the bryozoan order Cyclostomata is summarized and updated, based on field emission SEM studies of 87 species. Six fundamental ultrastructural fabrics are recognized which differ in the crystallographic orientations, shapes and prevailing growth directions of the constituent crystallites. During the growth of individual walls a succession of fabrics is secreted, defining a fabric suite. Five fabric suites are described in interior walls and four in exterior walls. Nine ultrastructural characters were combined with 37 other skeletal characters in a PAUP analysis of the relationships between 28 post-Palaeozoic cyclostomes chosen to include representatives of all suborders. A single tree of length 142 steps was found. Comparison of tree statistics for three categories of characters showed ultrastructural characters to be more homoplastic than zooidal characters, and the latter more homoplastic than colonial characters. Rooting the tree on the paleotubuliporine Cuffeyella gave four transitions from fixed- to free-walled organization and no reversals. With respect to the five extant suborders of cyclostomes, this first, preliminary analysis implies that Rectangulata and Cancellata are monophyletic groups, whereas Articulata are diphyletic, and both Tubuliporina and Cerioporina paraphyletic.     

Skeletal Ultrastructure in the Cyclostome Bryozoan Hornera

Abstract Scanning electron microscopy of calcified walls in two species of the cyclostome bryozoan Hornera has revealed previously undescribed details of skeletal morphology and growth. The calcitic interior walls of both H. robusta MacGillivray and H. squamosa Hutton have a laminated structure. Walls are extended at distal growing edges where the formation of new crystallites is concentrated and wall fabric is nacreous or semi-nacreous. New crystallites are seeded on the surface of existing crystallites as six-sided rhombs. At the centres of the rhombs in H. robusta there are often three ‘spikes' which point towards alternate sides of the rhomb. Screw dislocations resulting in spiral overgrowths are also common at these distal wall edges. Wall thickening occurs further proximally where walls develop a regularly foliated structure of imbricated laths growing towards the colony base. Although often thought to be ubiquitous in cyclostomes, the division of walls into three layers (an inner, primary layer flanked on both sides by secondary layers) is absent in Hornera. Wall ultrastructure contrasts strongly with the lamellar–fibrous–lamellar structure recently described from cinctiporid cyclostomes. The c-axes of the crystallites are orientated perpendicular to the wall surface in Hornera, unlike cinctiporids in which they are orientated within the plane of the wall. Apparent similarities in ultrastructure suggest that Hornera may provide a good model for wall growth in extinct trepostome bryozoans.     

Ultrastructure and Development of the Ooecial Walls in Some Calloporid Bryozoans (Gymnolaemata: Cheilostomata)

In the brood chambers (ovicells) of six calloporid cheilostomes studied each skeletal wall consists of four calcified layers: (1) a very thin superficial layer of planar spherulitic crystallites, (2) an upper (outer) layer with wall-perpendicular prismatic ultrastructure, (3) an intermediate lamellar layer, and (4) a lower (inner) wall-perpendicular prismatic layer. Comparative studies of both the ovicell wall ultrastructure and early ovicell formation showed a hypothetical opportunity for evolving complex (multilayered) skeletal walls by fusion of the initially separated gymnocystal and cryptocystal calcifications in Cheilostomata. In two species studied, a bilobate pattern in the final stage of the formation of the ooecial roof was encountered in specimens with the cuticle preserved. A possible explanation to this finding is discussed – the bilobate pattern is suggestive of the hypothetical origin of the brood chamber from (1) two flattened spines, or (2) reduction in spine number of an originally multispinous ovicell.     

Electron diffraction studies of the calcareous skeletons of bryozoans

Electron microscopy and electron diffraction were used to investigate mineral crystallites dissociated from the skeletal walls of six species belonging to the Bryozoa, a phylum of predominantly marine colony-forming invertebrate animals. Four cheilostome bryozoans (Flustra foliacea, Membranipora membranacea, Thalamoporella novaehollandiae and Cellarinella foveolata) and two cyclostomes (Fasciculipora ramosa and Hornera robusta) were analysed. In each case, an attempt was made to relate the crystal morphology imaged in situ by scanning electron microscopy with the crystallographic orientation of isolated crystals determined by electron diffraction analysis in the transmission electron microscope. The results showed that the calcitic cheilostome and cyclostome skeletons consisted of closely packed arrays of plate-like Mg-containing calcite crystallites, and that the crystallographic a-axis was preferentially aligned perpendicular to the top and bottom surfaces of the flattened particles. The results suggest that calcite biomineralization occurs under similar crystallographic constraints in the five species studied even though the origins of cheilostomes and cyclostomes are separated by over 300 million years in the fossil record of the bryozoans. Similar studies for the aragonite crystallites in skeletons of M. membranacea indicated that the crystallographic b-axis was preferentially oriented perpendicular to the basal surfaces of irregular plate-like particles.     

Septocea n. g. (Bryozoa cyclostomata), ein neues Kreide-Bryozoen-Genus von Rügen und Maastricht

Septocea septifera n.g. n.sp. (Family Semiceidae Buge) is a very strange free-walled cyclostome bryozoan species of Maastrichtian age with regard to its apertural morphogenesis. The zoarium is erect, eschariform, without heteromorphs, the aperturae lack ringlinke peristoms and are subdivided by vertical septa. The septa are recognized as extern walls of axially budded immature zooecia in the interior of the living chamber. The oval to triangular apertura is subdivided by the septa into one larger and one or two smaller openings. The evidence that axial intrazooecial budding also exists in other, not closely related taxa, proves that these structures are convergent. Their morphogenesis, systematic value and the taxonomic status of the Family Semiceidae are discussed.     

Crystallographic structure of the foliated calcite of bivalves

The foliated layer of bivalves is constituted by platy calcite crystals, or laths, surrounded by an organic layer, and which are arranged into sheets (folia). Therefore, the foliated microstructure can be considered the calcitic analogue to nacre. In this paper, the foliated microstructure has been studied in detail using electron and X-ray diffraction techniques, together with SEM observations on naturally decalcified shells, to investigate the crystallographic organization on different length scales and to resolve among previous contradictory results. This layer is highly organized and displays a coherent crystallographic orientation. The surface of the laths of the foliated layer is constituted by calcite crystals oriented with their c-axis tilted opposite to the growth direction of the laths and one of its {101 4} rhombohedral faces looking in the growth direction. These faces are only expressed as the terminal faces of the laths, whereas the main surfaces of laths coincide with {101 8} rhombohedral faces. This arrangement was consistently found in all specimens studied, which leads us to the provisional conclusion that, unlike previous studies, there is only one possible crystallographic arrangement for the foliated layer. Future studies on other species will help to ascertain this assertion.     

Understanding the crystallographic and nanomechanical properties of bryozoans

This study examines how microscale differences in skeletal ultrastructure affect the crystallographic and nanomechanical properties of two related bryozoan species: (i) Hornera currieae, which is found at relatively quiescent depths of c. 1000 m, and (ii) Hornera robusta, which lives at depths of 50–400 m where it is exposed to currents and storm waves. Microstructural and Electron Backscatter Diffraction (EBSD) observations show that in both species the secondary walls are composed of low-Mg calcite crystallites that grow with their c-axes perpendicular to the wall. Branches in H. currieae develop a strong preferred orientation of the calcite c-axes, while in H. robusta the c-axes are more scattered. Microstructural observations suggest that the degree of scattering is controlled by the underlying morphology of the skeletons: in H. currieae the laminated branch walls are smooth and relatively uninterrupted, whereas the wall architecture of H. robusta is modified by numerous deflections, forming pustules and ridges associated with microscopic tubules. Modelling of the Young’s modulus and measurements of nanoindentation hardness indicate that the observed scattering of the crystallite c-axes affects the elastic modulus and nanohardness of the branches, and therefore controls the mechanical properties of the skeletal walls. At relatively high pressure in deep waters, the anisotropic skeletal architecture of H. currieae is aimed at concentrating elasticity normal to the skeleton wall. In comparison, in the relatively shallow and active hydrographic regime of the continental shelf, the elastically isotropic skeleton of H. robusta is designed to increase protection from external predators and stronger omni-directional currents.     

Biomineralization in bryozoans: present,past and future

Many animal phyla have the physiological ability to produce biomineralized skeletons with functional roles that have been shaped by natural selection for more than 500 million years. Among these are bryozoans, a moderately diverse phylum of aquatic invertebrates with a rich fossil record and importance today as bioconstructors in some shallow‐water marine habitats. Biomineralizational patterns and, especially, processes are poorly understood in bryozoans but are conventionally believed to be similar to those of the related lophotrochozoan phyla Brachiopoda and Mollusca. However, bryozoan skeletons are more intricate than those of these two phyla. Calcareous skeletons have been acquired independently in two bryozoan clades – Stenolaemata in the Ordovician and Cheilostomata in the Jurassic – providing an evolutionary replicate. This review aims to highlight the importance of biomineralization in bryozoans and focuses on their skeletal ultrastructures, mineralogy and chemistry, the roles of organic components, the evolutionary history of bimineralization in bryozoans with respect to changes in seawater chemistry, and the impact of contemporary global changes, especially ocean acidification, on bryozoan skeletons. Bryozoan skeletons are constructed from three different wall types (exterior, interior and compound) differing in the presence/absence and location of organic cuticular layers. Skeletal ultrastructures can be classified into wall‐parallel (i.e. laminated) and wall‐perpendicular (i.e. prismatic) fabrics, the latter apparently found in only one of the two biomineralizing clades (Cheilostomata), which is also the only clade to biomineralize aragonite. A plethora of ultrastructural fabrics can be recognized and most occur in combination with other fabrics to constitute a fabric suite. The proportion of aragonitic and bimineralic bryozoans, as well as the Mg content of bryozoan skeletons, show a latitudinal increase into the warmer waters of the tropics. Responses of bryozoan mineralogy and skeletal thickness to oscillations between calcite and aragonite seas through geological time are equivocal. Field and laboratory studies of living bryozoans have shown that predicted future changes in pH (ocean acidification) combined with global warming are likely to have detrimental effects on calcification, growth rate and production of polymorphic zooids for defence and reproduction, although some species exhibit reasonable levels of resilience. Some key questions about bryozoan biomineralization that need to be addressed are identified.     

Evaluation of Succession in an Estuarine Macrobenthic Soft-Bottom Community near Tampa,Florida

A subtidal macrobenthic infaunal community was quantitatively sampled monthly from February 1975 to July 1978 (42 months). During that period, complete defaunation (presumably due to hypoxia) occurred three times at approximately annual intervals. The recovery of the community was examined for successional patterns by quantitative and qualitative normal and inverse classification analyses and by rank-order analysis of the dominant species. There was no consistent pattern of succession from recovery to recovery. Samples taken just after defaunation were not similar to each other and no consistent suites of species were detected. Classical succession in which suites of species are successively replaced by other suites until a persisting suite of species occurs (faciliation model) was not found. Other models of succession are discussed.     

A new early cyclostome bryozoan from the Lower Ordovician (Volkhov Stage) of Russia

Goryunovia hemiseptata n.gen. n.sp. is an unusual Lower Ordovician cyclostome bryozoan, represented by a single colony from the Middle Volkhov Stage (Arenig) of the Leningrad Oblast. The encrusting uniserial colony consists of large, slender zooids, many with closely-spaced partial diaphragms (hemisepta), and has a distinctive trichotomous branching pattern. Skeletal organization is of the sagenellid grade: calcified interior walls appear to be present, and the exterior frontal walls lack pseudopores. As details of microstructure and early astogeny are unknown, it is difficult to interpret the phylogenetic relationships ofGoryunovia until further material is forthcoming. However, there is some resemblance with the enigmatic hederellids, which appear in the Upper Silurian and possibly range into the Triassic.     

How did the carrier shell Xenophora crispa (König, 1825) build its shell? Evidence from the Recent and fossil record

The genus Xenophora comprises species of marine gastropods (Cretaceous-Recent) able to add fragments of various origins to their shell surface. Agglutination potentials vary, from species lacking attachments to species completely covered by agglutinated materials, as in the Mediterranean species Xenophora crispa. Here, we analyse Recent and fossil specimens of Xenophora crispa from the Mediterranean area using SEM and XRD, to better understand their biomineralization patterns and the mechanisms leading to the agglutination of shells, bioclasts and lithoclasts, and their evolution in time. We also provide new data on poorly studied gastropod shell microstructures. We conclude that: (1) most of the Xenophora crispa shell consists of an aragonitic crossed lamellar fabric, but fibrous to spherulitic prismatic fabrics, seemingly of calcite, have been found in the columella and peripheral edge (the thickest parts of the shell); (2) attachment of objects is mediated by a prismatic microstructure, indicating that this may be the most functional fabric in attachment areas in molluscs; and (3) the functional purpose of the agglutination in Xenophora crispa may be related to a snowshoe strategy to successfully colonize muddy substrates, coupled with tactile and olfactory camouflage. Indeed, this species secretes in the columella and peripheral edge a less dense and a more organic rich calcitic fabric, possibly to lighten the shell thickest parts in order not to sink in soft sediments and to facilitate the shell raising from the substrate to create a protected feeding area. This behaviour seems to have been maintained by X. crispa over 2 My time span.     

Amino silicones finished fabrics for lipase immobilization: Fabrics finishing and catalytic performance of immobilized lipase

Finishing of silk fabric was achieved by using amino-functional polydimethylsiloxane (PDMS) and lipase from Candida sp. 99-125 was immobilized on the treated silk fabrics. Hydrophobic fabrics were obtained by dipping the native fabric in 0.125–0.25% (w/v) PDMS solution and dried at 70 °C. The direct adsorption on PDMS-treated fabric was verified to be a better strategy for lipase immobilization than that by covalent binding. Compared to unfinished fabrics, the hydrolytic activity of immobilized enzyme on the finished fabric was improved by 1.6 times. Moreover, the activity of immobilized enzymes on hydrophobic fabrics was significantly improved in different concentrations of strong polar solvents such as methanol and ethanol, and in common organic solvents with different octanol–water partition coefficients (Log P). Enzymatic activity and stability in 15% water content system (added water accounted for the total reaction mixtures, v/v) showed more than 30% improvement in each batch. The amino–silicone finished fabric surface was investigated by scanning electron microscopy and X-ray photoelectron spectroscopy. The hydrophobic fabric immobilized enzyme could be recycled for more than 80 times with no significant decrease in esterification activity. PDMS-treated woven silk fabrics could be a potential support for lipase immobilization in catalytic esterification processes.     

Thyme essential oil for antimicrobial protection of natural textiles

The study was aimed at increasing the resistance of lignocellulosic textiles to bacteria and mould action using a biocide of plant origin. The biocide used in the study was thyme essential oil. This kind of oil is characterized by low toxicity for humans and the environment. The antimicrobial efficiency of thyme essential oil applied to linen–cotton blended fabric and linen fabric was evaluated by determining bacterial growth, degree of mould growth, and their impact on fabric strength. Thyme essential oil applied as 8% concentration in methanol to linen–cotton blended fabric showed very high antibacterial and antifungal activity – no mould growth and no significant loss of breaking force were observed. Microscopic evaluation of the tested fabrics was also performed by Scanning Electron Microscopy. Applying the eco-friendly biocide to fabrics containing natural fibres in the finishing process produces antimicrobial barrier properties.     

The interstitial crystal-nucleating sheet in molluscan Haliotis rufescens shell: A bio-polymeric composite

The interstitial green sheets in abalone shell nacre are shown to be bifacially differentiated trilaminate polymeric complexes, with glycoprotein layers sandwiching a central core containing chitin. They share some common feature with the organic matrix layers between the aragonite tablets in the nacre and the periostracum, and show similarities to the myostracum. Thus, although the green sheet is reported to be unique to the abalone shell, it represents an interesting model for the study of molluscan shell biomineralization processes. Indeed, during shell formation, prismatic and spherulitic aragonite precedes and follows the deposition of the interstitial green polymeric composite sheets, and there is evidence to suggest that these sheets demark the interruption of nacre synthesis and serve to nucleate the resumption of calcium carbonate crystal growth. The green polymeric interstitial sheet purified from the abalone shell was investigated by spectroscopic and imaging techniques: FTIR, confocal microscopy, scanning and transmission electron microscopy, and by pyrolysis combined with GC–MS. Structural and compositional differences are observed between the surfaces of the two sides of the interstitial polymeric composite sheets. Moreover, comparative crystallization experiments on the green sheet sides also reveal asymmetry with respect to the nucleation of calcium carbonate. These findings suggest that these bifacially differentiated interstitial composites may play an active role in the mineral assembly processes, with one of the surfaces acting as a crystal nucleator.     

Crocodilian Nest in a Late Cretaceous Sauropod Hatchery from the Type Lameta Ghat Locality,Jabalpur, India

The well-known Late Cretaceous Lameta Ghat locality (Jabalpur, India) provides a window of opportunity to study a large stable, near shore sandy beach, which was widely used by sauropod dinosaurs as a hatchery. In this paper, we revisit the eggs and eggshell fragments previously assigned to lizards from this locality and reassign them to crocodylomorphs. Several features point to a crocodilian affinity, including a subspherical to ellipsoidal shape, smooth, uneven external surface, discrete trapezoid shaped shell units with wide top and narrow base, basal knobs and wedge shaped crystallites showing typical inverted triangular extinction under crossed nicols. The crocodylomorph eggshell material presented in this paper adds to the skeletal data of these most probably Cretaceous-Eocene dryosaurid crocodiles.     

蝗虫消化道形态结构研究的一种新方法

采用扫描仪和扫描电镜相结合的方法对中华稻蝗Oxyachinensis(Thunberg)消化道内部的超微结构进行定位研究。结果表明直接将蝗虫消化道的内壁翻出(不染色)铺展于扫描仪的玻璃板上进行扫描,就能得到蝗虫消化道在自然状态下的完整图像,将扫描仪扫描后的样品用于扫描电子显微镜的样品制备,并对照上述研究结果,即可定位蝗虫消化道在扫描电子显微镜下的超微结构。为蝗虫消化道的形态学研究提供一个简便有效的方法,同时也为蝗虫消化道超微结构的定位研究提供新的手段。