Where's the glass? Biomarkers,molecular clocks,and microRNAs suggest a 200‐Myr missing Precambrian fossil record of siliceous sponge spicules

The earliest evidence for animal life comes from the fossil record of 24-isopropylcholestane, a sterane found in Cryogenian deposits, and whose precursors are found in modern demosponges, but not choanoflagellates, calcareans, hexactinellids, or eumetazoans. However, many modern demosponges are also characterized by the presence of siliceous spicules, and there are no convincing demosponge spicules in strata older than the Cambrian. This temporal disparity highlights a problem with our understanding of the Precambrian fossil record – either these supposed demosponge-specific biomarkers were derived from the sterols of some other organism and are simply retained in modern demosponges, or spicules do not primitively characterize crown-group demosponges. Resolving this issue requires resolving the phylogenetic placement of another group of sponges, the hexactinellids, which not only make a spicule thought to be homologous to the spicules of demosponges, but also make their first appearance near the Precambrian/Cambrian boundary. Using two independent analytical approaches and data sets – traditional molecular phylogenetic analyses and the presence or absence of specific microRNA genes – we show that demosponges are monophyletic, and that hexactinellids are their sister group (together forming the Silicea). Thus, spicules must have evolved before the last common ancestor of all living siliceans, suggesting the presence of a significant gap in the silicean spicule fossil record. Molecular divergence estimates date the origin of this last common ancestor well within the Cryogenian, consistent with the biomarker record, and strongly suggests that siliceous spicules were present during the Precambrian but were not preserved.     

Early sponge evolution: A review and phylogenetic framework

Sponges are one of the critical groups in understanding the early evolution of animals. Traditional views of these relationships are currently being challenged by molecular data, but the debate has so far made little use of recent palaeontological advances that provide an independent perspective on deep sponge evolution. This review summarises the available information, particularly where the fossil record reveals extinct character combinations that directly impinge on our understanding of high-level relationships and evolutionary origins. An evolutionary outline is proposed that includes the major early fossil groups, combining the fossil record with molecular phylogenetics. The key points are as follows. (1) Crown-group sponge classes are difficult to recognise in the fossil record, with the exception of demosponges, the origins of which are now becoming clear. (2) Hexactine spicules were present in the stem lineages of Hexactinellida, Demospongiae, Silicea and probably also Calcarea and Porifera; this spicule type is not diagnostic of hexactinellids in the fossil record. (3) Reticulosans form the stem lineage of Silicea, and probably also Porifera. (4) At least some early-branching groups possessed biminerallic spicules of silica (with axial filament) combined with an outer layer of calcite secreted within an organic sheath. (5) Spicules are homologous within Silicea, but also between Silicea and Calcarea, and perhaps with Homoscleromorpha. (6) The last common ancestor of extant sponges was probably a thin-walled, hexactine-bearing sponge with biminerallic spicules. (7) The stem group of sponges included tetraradially-symmetric taxa that grade morphologically into Cambrian fossils described as ctenophores. (8) The protomonaxonid sponges are an early-branching group, probably derived from the poriferan stem lineage, and include the problematic chancelloriids as derived members of the piraniid lineage. (9) There are no definite records of Precambrian sponges: isolated hexactine-like spicules may instead be derived from radiolarians. Early sponges had mineralised skeletons and thus should have a good preservation potential: the lack of sponge fossils in Precambrian strata may be due to genuine absence of sponges. (10) In contrast to molecular clock and biomarker evidence, the fossil record indicates a basal Cambrian diversification of the main sponge lineages, and a clear relationship to ctenophore-like ancestors. Overall, the early sponge fossil record reveals a diverse suite of extinct and surprising character combinations that illustrate the origins of the major lineages; however, there are still unanswered questions that require further detailed studies of the morphology, mineralogy and structure of early sponges.     

Evidence for spicule homology in calcareous and siliceous sponges: biminerallic spicules in Lenica sp. from the Early Cambrian of South China

Botting, J.P., Muir, L.A., Xiao, S., Li, X. & Lin, J.‐P. 2012: Evidence for spicule homology in calcareous and siliceous sponges: biminerallic spicules in Lenica sp. from the Early Cambrian of South China. Lethaia, Vol. 45, pp. 463–475. The relationships of the extant sponge classes, and the nature of the last common ancestor of all sponges, are currently unclear. Early sponges preserved in the fossil record differ greatly from extant taxa, and therefore information from the fossil record is critical for testing hypotheses of sponge phylogenetic relationships that are based on modern taxa. New specimens of the enigmatic sponge Lenica sp., from the Early Cambrian Hetang Biota of South China, exhibit an unusual spicule structure. Each spicule consists of a siliceous core with an axial canal, an organic outer layer and a middle layer interpreted to have been originally calcium carbonate. This finding confirms previous work suggesting the existence of biminerallic spicules in early sponges. Combined with data from other early sponges, the new findings imply that the two fundamental spicule structures of modern sponges were derived from a compound, biminerallic precursor. Spicules are therefore homologous structures in Calcarea and Silicea, and if sponges are paraphyletic with respect to Eumetazoa, then spicules may also have been a primitive feature of Metazoa. □Calcarea, Early Cambrian, Hetang Biota, phylogeny, Silicea, taphonomy.     

‘Cambrian’ demosponges in the Ordovician of Morocco: Insights into the early evolutionary history of sponges

Cambrian spicular sponge faunas are dominated by a distinctive assemblage of demosponges and hexactinellids that are known from Burgess shale-type faunas worldwide. Most of these are previously unknown outside the Lower-Middle Cambrian (and perhaps Tremadoc) and have no obvious close relatives in later sequences. This paper describes examples of Choia sp., Pirania auraeum sp. nov. and Hamptonia christi sp. nov. from the Arenig of Morocco, associated with isolated hexactinellid spicules. A summary of the stratigraphic ranges of the major Cambrian sponge lineages is provided. These indicate an environmental contrast in the Lower Palaeozoic evolution of hexactinellids and non-lithistid demosponges, with demosponges probably undergoing cryptic diversification in nearshore environments during the Upper Ordovician.     

Bio-sintering processes in hexactinellid sponges: Fusion of bio-silica in giant basal spicules from Monorhaphis chuni

The two sponge classes, Hexactinellida and Demospongiae, comprise a skeleton that is composed of siliceous skeletal elements (spicules). Spicule growth proceeds by appositional layering of lamellae that consist of silica nanoparticles, which are synthesized via the sponge-specific enzyme silicatein. While in demosponges during maturation the lamellae consolidate to a solid rod, the lamellar organization of hexactinellid spicules largely persists. However, the innermost lamellae, near the spicule core, can also fuse to a solid axial cylinder. Similar to the fusion of siliceous nanoparticles and lamella, in several hexactinellid species individual spicules unify during sintering-like processes. Here, we study the different stages of a process that we termed bio-sintering, within the giant basal spicule (GBS) of Monorhaphis chuni. During this study, a major GBS protein component (27 kDa) was isolated and analyzed by MALDI-TOF-MS. The sequences were used to isolate and clone the encoding cDNA via degenerate primer PCR. Bioinformatic analyses revealed a significant sequence homology to silicatein. In addition, the native GBS protein was able to mediate bio-silica synthesis in vitro. We conclude that the syntheses of bio-silica in M. chuni, and the subsequent fusion of nanoparticles to lamellae, and finally to spicules, are enzymatically-driven by a silicatein-like protein. In addition, evidence is now presented that in hexactinellids those fusions involve sintering-like processes.     

Cyathophycus and the origin of demosponges

The relationships between the poriferan classes are currently obscure. Molecular phylogenies appear to be reaching a consensus that the hexactinellids and demosponges are closely related, despite previous attempts to separate the Hexactinellida from other sponges on cytological grounds, but the details of the transition are unknown. Similarities of spicule morphology and structure are used to infer that the transition probably occurred after the onset of silicification, and should therefore be seen in fossils. The similarity between protosponges and early demosponges has been noted previously, based largely on a thin, reticulated wall of simple spicules (monaxons in Leptomitus, stauracts in Protospongia). A close relationship is, however, unlikely, since the protospongiids possessed a precise geometric arrangement of multiple spicule size orders that is lacking in demosponges. A close morphological similarity exists between the skeletons of transitional protosponge-dictyosponge reticulosids, such as Cyathophycus, and the early hazeliid demosponges. The inner spicule layer of primitive dictyosponges consists of a cross-hatched array of fine monaxons, as seen in the wall of some simple hazeliids. Although most described species of hazeliids were morphologically complex, a simple globose species is here recorded from the Caradoc of Wales. The evolutionary link between the classes is suggested to lie within the dictyospongioid and hazeliid lineages. Although the direction of evolution cannot be certainly fixed, it is conceptually much easier to derive the demosponges from the hexactinellids, rather than vice versa.     

The dorid nudibranchs Peltodoris lentiginosa and Archidoris odhneri as predators of glass sponges

The dorid nudibranchs Peltodoris lentiginosa and Archidoris odhneri were found on glass sponges (Porifera, Hexactinellida) during remotely operated vehicle surveys of three reefs in the Strait of Georgia, British Columbia, Canada. Eight nudibranchs were sampled from 2009 to 2011. Identification of sponge spicules found in their gut and fecal contents confirmed the nudibranchs to be predators of the reef‐forming hexactinellids Aphrocallistes vastus and Heterochone calyx, as well as of the demosponge Desmacella austini, which encrusts skeletons of the glass sponges. Four of five nudibranchs dissected for gut content analysis had stomachs containing sponge spicules. Counts from high‐definition video footage taken during systematic surveys done in 2009 showed that nudibranchs were found in only two of the three glass sponge reefs. These data provide the first quantitative evidence of a molluscan predator on glass sponges found outside of Antarctica, and establish the first trophic link between glass sponges and their associated community of animals in a sponge reef ecosystem on the western Canadian continental shelf.     

皖南早寒武世荷塘组海绵骨针化石

本文报道皖南休宁县早寒武世荷塘组黑色页岩中产出的海绵骨针化石组合,这些海绵骨针化石具有较高的丰度和分异度,它们以二轴四射针、T型针、三轴六射针和三轴五射针为主。骨针形态完整,并保存了内部轴丝、轴管以及同心圈层等微细构造。黄铁矿化在化石的保存中起了重要的作用,化石产出的时代可能为梅树村阶至筇竹寺阶（Tommotian-Atdabanian),这个化石组合证实了海绵动物在早寒武世已开始迅速分异。     

Middle and Late Cambrian sponge spicules from Hunan, China

Abundant and well-preserved assemblages of disarticulated sponge spicules occur in Middle and Late Cambrian platform carbonates of western Hunan, China. Assemblages recovered from 11 stratigraphic horizons include calcisponges, demosponges, and hexactinellids. Hexactinellida, in particular, are both abundant and diverse in Upper Cambrian carbonates. Comparison with spicule assemblages from Australia indicates that many of these taxa have long stratigraphic ranges, limiting their use in correlation. The morphological diversity of these spicules exceeds that known for living siliceous sponges, supporting the observation that during the Cambrian radiation, sponges, like other metazoans, evolved a variety of architectural forms not observed in later periods. Like conodonts, individual sponges can produce more than one spicule form; thus, an "apparatus genus" concept based on multiple co-occurring elements may eventually prove useful in the biostratigraphic and paleobiological interpretation of disarticulated sponge spicules. Four distinctive forms are recognized as new taxa: Australispongia sinensis new genus and species, Flosculus gracilis new genus and species, Pinnatispongia bengtsoni new genus and species, and Nabaviella paibiensis new species.     

Axial growth of hexactinellid spicules: formation of cone-like structural units in the giant basal spicules of the hexactinellid Monorhaphis

The glass sponge Monorhaphis chuni (Porifera: Hexactinellida) forms the largest bio-silica structures on Earth; their giant basal spicules reach sizes of up to 3 m and diameters of 8.5 mm. Previously, it had been shown that the thickness growth proceeds by appositional layering of individual lamellae; however, the mechanism for the longitudinal growth remained unstudied. Now we show, that the surface of the spicules have towards the tip serrated relief structures that are consistent in size and form with the protrusions on the surface of the spicules. These protrusions fit into the collagen net that surrounds the spicules. The widths of the individual lamellae do not show a pronounced size tendency. The apical elongation of the spicule proceeds by piling up cone-like structural units formed from silica. As a support of the assumption that in the extracellular space silicatein(-like) molecules exist that associate with the external surface of the respective spicule immunogold electron microscopic analyses were performed. With the primmorph system from Suberites domuncula we show that silicatein(-like) molecules assemble as string- and net-like arrangements around the spicules. At their tips the silicatein(-like) molecules are initially stacked and at a later stay also organized into net-like structures. Silicatein(-like) molecules have been extracted from the giant basal spicule of Monorhaphis. Applying the SDS–PAGE technique it could be shown that silicatein molecules associate to dimers and trimers. Higher complexes (filaments) are formed from silicatein(-like) molecules, as can be visualized by electron microscopy (SEM). In the presence of ortho-silicate these filaments become covered with 30–60 nm long small rod-like/cuboid particles of silica. From these data we conclude that the apical elongation of the spicules of Monorhaphis proceeds by piling up cone-like silica structural units, whose synthesis is mediated by silicatein(-like) molecules.     

深海六放海绵大骨针的结构与特性

在海绵动物(多孔动物)中,六放海绵和寻常海绵为硅质骨骼.生活在深海(1 000 m)中的六放海绵是最古老的海绵动物,其中间单根海绵和春氏单根海绵有长达3 m的骨针,是地球上最长的生物硅结构.利用电子显微技术观测, 这些直径达8 mm的巨大根须骨针具有同心层状结构,其横截面显示明显的构造分界:中间为含有轴丝的轴管,外围是一50-150 μm厚的轴筒,最外面为区状区(300-500层,每层厚度3-5 μm).生物化学研究显示其主要的蛋白质为35 kD大分子,另外,还检测到23-24 kD 多肽,可能是硅蛋白相关蛋白.依据现有的红血球凝聚活性,从骨针提取物中也检测到了凝集素.由电子探针获得其化学成分主要为Si,K和Na.此外,骨针的光传输实验表明,该巨大根须骨针用作光纤可传输600 nm至1 400 nm范围的光,而滤掉小于600 nm的光(类似高通滤波器)和大于1 400 nm 的红外光(类似低通滤波器).另外,从六放海绵的空囊泡沫海绵中分离出一个基因并确证了其推导的编码蛋白序列,该蛋白编码一个光裂合酶相关蛋白,蛋白相似性比较结果显示属于光裂合酶相关蛋白中多细胞动物隐色素一类.基于以上数据给出了六放海绵硅质骨针形成的示意图.另外,由单根海绵骨针可作为波导传输光/电和/或化学信号,推断在海绵动物中有类似神经系统的网络系统[动物学报 53(3):557-569,2007].     

Growth patterns of Lower Palaeozoic sponges

Detailed studies of the growth patterns of modern siliceous sponges are restricted to demosponges and theoretical models. It is generally assumed that sponge growth is essentially incremental, with completion of one arbitrary unit being followed by external addition. All recent species are thick-walled, but Lower Palaeozoic sponges are dominated by thin-walled hexactinellids, with most Cambrian taxa consisting of a single spicule layer. Large populations of a primitive dictyospongiid have allowed the reconstruction of the growth patterns of their spicules and body morphology. The results indicate that growth occurred through continuous expansion of the globose body, accompanied by continuous enlargement of existing spicules, with a spicule size limit being reached only during the lifetime of a few individuals. It is noted that this skeletal growth pattern is otherwise restricted to deuterostomes. Consecutive appearance of successive spicule size orders appears to have maintained a maximum inhalant pore area. Comparisons with more limited data from two acanthose hexactinellids and a hazeliid demosponge indicate that an identical growth pattern operated in these species. The subsequent evolution of growth patterns is discussed, with various mechanisms producing the later thick-walled morphologies of hexactinellids and demosponges. The implications of these observations are discussed with reference to identification and systematics, since spicule size and arrangement are shown to vary during growth.     

Formation of giant spicules in the deep-sea hexactinellid <Emphasis Type="Italic">Monorhaphis chuni</Emphasis> (Schulze 1904): electron-microscopic and biochemical studies

The siliceous sponge Monorhaphis chuni (Hexactinellida) synthesizes the largest biosilica structures on earth (3 m). Scanning electron microscopy has shown that these spicules are regularly composed of concentrically arranged lamellae (width: 3–10 μm). Between 400 and 600 lamellae have been counted in one giant basal spicule. An axial canal (diameter: ~2 μm) is located in the center of the spicules; it harbors the axial filament and is surrounded by an axial cylinder (100–150 μm) of electron-dense homogeneous silica. During dissolution of the spicules with hydrofluoric acid, the axial filament is first released followed by the release of a proteinaceous tubule. Two major proteins (150 kDa and 35 kDa) have been visualized, together with a 24-kDa protein that cross-reacts with antibodies against silicatein. The spicules are surrounded by a collagen net, and the existence of a hexactinellidan collagen gene has been demonstrated by cloning it from Aphrocallistes vastus. During the axial growth of the spicules, silicatein or the silicatein-related protein is proposed to become associated with the surface of the spicules and to be finally internalized through the apical opening to associate with the axial filament. Based on the data gathered here, we suggest that, in the Hexactinellida, the growth of the spicules is mediated by silicatein or by a silicatein-related protein, with the orientation of biosilica deposition being controlled by lectin and collagen. Carsten Eckert was previously with the Museum für Naturkunde, Invalidenstrasse 43, 10115 Berlin, Germany. The collagen sequence from Aphrocallistes vastus reported here, viz., [COL_APHRO] APHVACOL (accession number AM411124), has been deposited in the EMBL/GenBank data base. This work was supported by grants from the European Commission, the Deutsche Forschungsgemeinschaft, the Bundesministerium für Bildung und Forschung Germany (project: Center of Excellence BIOTECmarin), the National Natural Science Foundation of China (grant no. 50402023), and the International Human Frontier Science Program.     

Formation of siliceous spicules in the marine demosponge <Emphasis Type="Italic">Suberites domuncula</Emphasis>

The siliceous skeleton of demosponges is constructed of spicules. We have studied the formation of spicules in primmorphs from Suberites domuncula. Scanning electron microscopy and transmission electron-microscopical (TEM) analyses have revealed, in the center of the spicules, an axial canal that is 0.3–1.6 m wide and filled with an axial filament. This filament is composed of the enzyme silicatein, which synthesizes the spicules. TEM analysis has shown that spicule formation starts intracellularly and ends extracellularly in the mesohyl. At the initial stage, the axial canal is composed only of silicatein, whereas membranous structures and fibrils (10–15 nm in width) can later also be identified, suggesting that intracellular components protrude into the axial canal. Antibodies against silicatein have been applied for Western blotting; intracellularly, silicatein is processed to the mature form (24 kDa), whereas the pro-enzyme with the propeptide (33 kDa) is detected extracellularly. Silicatein undergoes phosphorylation at five sites. Immunohistological analysis has shown that silicatein exists in the axial canal (axial filament) and on the surface of the spicules, suggesting that they grow by apposition. Finally, we have demonstrated that the enzymic reaction of silicatein is inhibited by anti-silicatein antibodies. These data provide, for the first time, a comprehensive outline of spicule formation.This work was supported by grants from the European Commission (SILIBIOTEC), the Deutsche Forschungsgemeinschaft, the Bundesministerium für Bildung und Forschung Germany (project: Center of Excellence BIOTECmarin) and the International Human Frontier Science Program.     

Analysis of the axial filament in spicules of the demosponge Geodia cydonium: different silicatein composition in microscleres (asters) and megascleres (oxeas and triaenes)

The skeleton of the siliceous sponges (Porifera: Hexactinellida and Demospongiae) is supported by spicules composed of bio-silica. In the axial canals of megascleres, harboring the axial filaments, three isoforms of the enzyme silicatein (-alpha, -beta and -gamma) have been identified until now, using the demosponges Tethya aurantium and Suberites domuncula. Here we describe the composition of the proteinaceous components of the axial filament from small spicules, the microscleres, in the demosponge Geodia cydonium that possesses megascleres and microscleres. The morphology of the different spicule types is described. Also in G. cydonium the synthesis of the spicules starts intracellularly and they are subsequently extruded to the extracellular space. In contrast to the composition of the silicateins in the megascleres (isoforms: -alpha, -beta and -gamma), the axial filaments of the microscleres contain only one form of silicatein, termed silicatein-alpha/beta, with a size of 25kDa. Silicatein-alpha/beta undergoes three phosphorylation steps. The gene encoding silicatein-alpha/beta was identified and found to comprise the same characteristic sites, described previously for silicateins-alpha or -beta. It is hypothesized, that the different composition of the axial filaments, with respect to silicateins, contributes to the morphology of the different types of spicules.     

Brief communication: Investigation of the semicircular canal variation in the Krapina Neandertals

Previous studies comparing bony labyrinth morphology in geographically‐dispersed samples of Neandertals and modern Homo sapiens (H. sapiens) showed that Neandertals generally have smaller semicircular canals than modern H. sapiens (Hublin et al., 1996 ; Spoor et al., 2003 ; Glantz et al., 2008 ). Here we analyze the morphology of a single group of Neandertal specimens from one locale, the Krapina site, to determine the intraspecific variation in Neandertal semicircular canal sizes. Dimensions of the semicircular canals were collected from computed tomography scans of nine temporal bones. With the rare exception, the dimensions of the semicircular canals in the Krapina sample are similar to those previously reported across a geographically‐dispersed sample of Neandertals, further supporting previous studies that suggest low levels of variation in the semicircular canals for Neandertals. Am J Phys Anthropol 154:302–306, 2014. © 2014 Wiley Periodicals, 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 and ontogeny of multiple lateral‐line canals in the rock prickleback,Xiphister mucosus (Cottiformes: Zoarcoidei: Stichaeidae)

The structure and ontogeny of lateral‐line canals in the Rock Prickleback, Xiphister mucosus, were studied using cleared‐and‐stained specimens, and the distribution and morphology of neuromasts within lateral‐line canals were examined using histology. X. mucosus has seven cephalic canals in a pattern that, aside from four branches of the infraorbital canals, is similar to that of most teleostean fishes. Unlike most other teleosts, however, X. mucosus features multiple trunk lateral‐line canals. These include a short median posterior extension of the supratemporal canal and three paired, branching canals located on the dorsolateral, mediolateral, and ventrolateral surfaces. The ventrolateral canal (VLC) includes a loop across the ventral surface of the abdomen. All trunk canals, as well as the branches of the infraorbitals, are supported by small, dermal, ring‐like ossifications that develop independently from scales. Trunk canals develop asynchronously with the mediodorsal and dorsolateral canals (DLC) developing earliest, followed by the VLC, and, finally, by the mediolateral canal (MLC). Only the mediodorsal and DLC connect to the cephalic sensory canals. Fractal analysis shows that the complexity of the trunk lateral‐line canals stabilizes when all trunk canals develop and begin to branch. Histological sections show that neuromasts are present in all cephalic canals and in the DLC and MLC of the trunk. However, no neuromasts were identified in the VLC or its abdominal loop. The VLC cannot, therefore, directly function as a part of the mechanosensory system in X. mucosus. The evolution and functional role of multiple lateral‐line canals are discussed. J. Morphol. 276:1218–1229, 2015. © 2015 Wiley Periodicals, Inc.     

Ultrastructure of siliceous spicules and microsclerocytes in the marine sponge Neofibularia irata N. SP.

The marine sponge Neofibularia irata contains four different categories of siliceous spicules. These spicules are evident in the tissues as distinct bundles that act to increase the structural rigidity of the sponge. All spicules have a normal structural morphology with silica deposition around a hexagonal axial canal containing a crystalline axial filament. The megasclere strongyles are secreted in typical megasclerocytes. The sigma and raphid microscleres are secreted in individual microsclerocytes that are grouped together in parallel to form loose bundles. However, the microxea microscleres are apparently secreted in distinct tight bundles (trichodragmas) within a single cell. These cells, containing between 13 and 39 spicules, are grouped to form large packets of bundles of spicules.