First evidence of the presence of chitin in skeletons of marine sponges. Part II. Glass sponges (Hexactinellida: Porifera)

Sponges (Porifera) are presently gaining increased scientific attention because of their secondary metabolites and specific skeleton structures. In contrast to demosponges, whose skeletons are formed from biopolymer spongin, glass sponges (hexactinellids) possess silica-organic composites as the main natural material for their skeletal fibres. Chitin has a crystalline structure and it constitutes a network of organized fibres. This structure confers rigidity and resistance to organisms that contain it, including monocellular (yeast, amoeba, diatoms) and multicellular (higher fungi, arthropods, nematodes, molluscs) organisms. In contrast to different marine invertebrates whose exoskeletons are built of chitin, this polysaccharide has not been found previously as an endogenous biopolymer within glass sponges (Hexactinellida). We hypothesized that glass sponges, which are considered to be the most basal lineage of multicellular animals, must possess chitin. Here, we present a detailed study of the structural and physico-chemical properties of skeletal fragments of the glass sponge Farrea occa. We show that these fibres have a layered design with specific compositional variations in the chitin/silica composite. We applied an effective approach for the demineralization of glass sponge skeletal formations based on an etching procedure using alkali solutions. The results show unambiguously that alpha-chitin is an essential component of the skeletal structures of Hexactinellida. This is the first report of a silica-chitin's composite biomaterial found in nature. From this perspective, the view that silica-chitin scaffolds may be key templates for skeleton formation also in ancestral unicellular organisms, rather than silica-protein composites, emerges as a viable alternative hypothesis.     

First evidence of miniature transposable elements in sponges (Porifera)

Transposable elements play a vital role in genome evolution and may have been important for the formation of the early metazoan genome, but only little is known about transposons at this interface between unicellular opisthokonts and Metazoa. Here, we describe the first miniature transposable elements (MITEs, Queen1 and Queen2) in sponges. Queen1 and Queen2 are probably derived from Tc1/mariner-like MITE families and are represented in more than 3,800 and 1,700 copies, respectively, in the Amphimedon queenslandica genome. Queen elements are located in intergenic regions as well as in introns, providing the potential to induce new splicing sites and termination signals in the genes. Further possible impacts of MITEs on the evolution of the metazoan genome are discussed.     

Screening of the Antarctic marine sponges (Porifera) as a source of bioactive compounds

Sponges (Porifera) currently represent one of the richest sources of natural products and account for almost half of the pharmacologically active compounds of marine origin. However, to date very little is known about the pharmacological potential of the sponges from polar regions. In this work we report on screening of ethanolic extracts from 24 Antarctic marine sponges for different biological activities. The extracts were tested for cytotoxic effects against normal and transformed cell lines, red blood cells, and algae, for modulation of the activities of selected physiologically important enzymes (acetylcholinesterase, butyrylcholinesterase, and α-amylase), and for inhibition of growth of pathogenic and ecologically relevant bacteria and fungi. An extract from Tedania (Tedaniopsis) oxeata was selectively cytotoxic against the cancer cell lines and showed growth inhibition of all of the tested ecologically relevant and potentially pathogenic fungal isolates. The sponge extracts from Isodictya erinacea and Kirkpatrickia variolosa inhibited the activities of the cholinesterase enzymes, while the sponge extracts from Isodictya lankesteri and Inflatella belli reduced the activity of α-amylase. Several sponge extracts inhibited the growth of multiresistant pathogenic bacterial isolates of different origins, including extended-spectrum beta-lactamase and carbapenem-resistant strains, while sponge extracts from K. variolosa and Myxilla (Myxilla) mollis were active against a human methicillin-resistant Staphylococcus aureus strain. We conclude that Antarctic marine sponges represent a valuable source of biologically active compounds with pharmacological potential.     

First evidences of surfactant biodegradation by marine sponges (Porifera): an experimental study with a linear alkylbenzenesulfonate

The main objective of this work was to better appreciate the role of benthic macro-organisms in the biodegradation of detergents in the marine environment. Sponges, which could be highly resistant to pollution and which are highly active filter-feeders, appear as interesting organisms in this topic. An experimental study conducted in aquarium with seawater enriched in a pure linear alkylbenzenesulfonate (LAS), namely 1-(p-Sulfophenyl)nonane, has shown that the primary degradation was 10 times more rapid in the presence of the sponge Spongia officinalis than in the presence of only marine bacteria. The main degradation metabolites, 3-(p-sulfophenyl)propionic acid and p-sulfobenzoic acid were produced in greater amounts in the presence of sponge. The very rapid degradation kinetics observed in this study may be due to the symbiotic microbial community present in S. officinalis. The sponge may also have promoted the activity of marine bacteria by transforming the original molecule in a more available secondary product. This study demonstrates for the first time that benthic macro-organisms can play an important role in the transformation of biodegradable contaminants, such as LAS.     

Molecular systematics of sponges (Porifera)

The first application of molecular systematics to sponges was in the 1980s, using allozyme divergence to dis-criminate between conspecific and congeneric sponge populations. Since this time, a fairly large database has been accumulated and, although the first findings seemed to indicate that sponge species were genetically more divergent than those of other marine invertebrates, a recent review of the available dataset indicates that levels of interspecific gene identities in most sponges fall within the normal range found between species of other invertebrates. Nevertheless, some sponge genera have species that are extremely divergent from each other, suggesting a possible polyphyly of these genera. In the 1990s, molecular studies comparing sequences of ribosomal RNA have been used to reappraise the phylogenetic relationships among sponge genera, families, orders and classes. Both the 18S small subunit and the 28S large subunit rRNA genes have been sequenced (41 complete or partial and 75 partial sequences, respectively). Sequences of 18S rRNA show good support for Porifera being true Metazoa, but they are not informative for resolving relationships among genera, families or orders. 28S rRNA domains D1 and D2 appear to be more informative for the terminal nodes and provide resolution for internal topologies in sufficiently closely related species, but the deep nodes between orders or classes cannot be resolved using this molecule. Recently, a more conserved gene, Hsp70, has been used to try to resolve the relationships in the deep nodes. Metazoan monophyly is very well supported. Nevertheless, the divergence between the three classes of Porifera, as well as the divergence between Porifera, Cnidaria and Ctenophora, is not resolved. Research is in progress using other genes such as those of the homeodomain, the tyrosine kinase domain, and those coding for the aggregation factor. For the moment the dataset for these genes is too restricted to resolve the phylogenetic relationships of these phyla. However, whichever the genes, the phylogenies obtained suggest that Porifera could be paraphyletic and that the phylogenetic relationships of most of the families and orders of the Demospongiae have to be reassessed. The Calcarea and Hexactinellida are still to be studied at the molecular level.     

Spicules of hexactinellid sponges (Hexactinellida: Porifera) as natural composite materials

This paper reviews studies on the hexactinellid glass sponges (Hexactinellida: Porifera) that have organic silica spicules. According to its physical properties (microdensity, Young’s modulus, and light transmission), the material of the spicules is similar to amorphous silica; however, sponge spicules are birefringent, which suggests that they have a highly ordered crystal-like nature. Mineralized remnants of siliceous spicules composed of chemically inert materials are preserved in sedimentary rocks and provide evidence of the ecological state of the ancient biosphere. Sponges occur in waters with low temperatures; therefore, they grow very slowly and live for hundreds of years. The organic silica spicules exhibit the capacity for triboluminescence. The generated light emission may be used by symbiotic bacteria on the spicule surface.     

Space as a limiting resource in freshwater systems: competition between zebra mussels (Dreissena polymorpha) and freshwater sponges (Porifera)

Species interactions between two types of sessile benthic invertebrates, the zebra mussel (Dreissena polymorpha) and freshwater sponges (Porifera), were evaluated in Michigan City IN Harbor in southern Lake Michigan during 1996. The study objective was to define whether competition plays a role in structuring benthic communities using experimental techniques commonly employed in marine systems. Sponges were uninhibited by zebra mussel presence and overgrew zebra mussel shells on hard vertical substrata. In contrast, zebra mussels did not overgrow sponge colonies, but did show an ability to re-capture hard substrata if relinquished by the sponge. The negative affect of sponges on zebra mussels through overgrowth and recruitment suggests interactions that could eventually displace zebra mussels from these benthic communities. However, seasonal reduction of sponge biomass from autumn through winter appears to allow the zebra mussel a periodic respite from overgrowth, preventing exclusion of zebra mussels from the community and allowing these two taxa to co-exist.     

On functional morphology of the skeleton in lychnisc sponges (Porifera,Hexactinellida)

By comparison with architectural models it can be shown that both the “euretoid” arrangement of fused spicules and lychnisc node itself should enhance the strength of the skeleton of the Lychniscosa. Contrary to expectation however, the Lychniscosa do not inhabit more turbulent waters than the Hexactinosa with a simpler skeletal structure. Possible reasons for this are discussed.     

Implications from a 28S rRNA gene fragment for the phylogenetic relationships of halichondrid sponges (Porifera: Demospongiae)

Halichondrida is a pivotal demosponge order of which the classification underwent major changes in the recent history. The monophyly of this order and its intra-ordinal phylogeny cannot be reliably determined on the basis of morphology. Here we present a 28Sr RNA gene tree of selected halichondrids, which supports the hypothesis of halichondrid non-monophyly and elucidates further inter-ordinal relationships. We enlarged the analysis by previously published sequences, discuss how previous analyses suffer from taxon bias and analyse the resulting phylogenetic implications. Most halichondrid families (in particular Axinellidae und Dictyonellidae) cluster polyphyletic and the molecular classification of several genera does not agree with the current (morphological) system.     

The sterol composition of the marine sponge Aplysina (= Verongia) archeri: a comparative study of the Verongidae

1. The sterol composition of the marine sponge Aplysina archeri was studied revealing the presence, besides other common sterols, of considerable amounts (75%) of aplysterol (1), verongulasterol (3), and 25-dehydroaplysterol (4). 2. 24,28-Didehydroaplysterol (2) was not present in the sterol mixture as it was previously reported for the Verongidae. 3. A comparison between the characteristic sterols of the Verongidae was carried out.     

Intragenomic variation of the rDNA internal transcribed spacers in sponges (Phylum Porifera): implications for phylogenetic studies

The internal transcribed spacer regions (ITS1 and ITS2) of the tandemly repeated nuclear ribosomal DNA clusters are frequently used as markers for fine scale analyses in diverse animals. In certain taxa, ITS is nearly exclusively used for population level or inter-specific studies, despite the frequent presence of divergent paralogs within individual genomes that can be phylogenetically misleading. For the first time we survey diverse marine sponges to determine the extent and phylogenetic implications of intragenomic polymorphisms (IGPs) exhibited at their ITS loci. We discover that the extent of IGP varies greatly between taxa (with most taxa exhibiting very few) and cannot be predicted by taxonomy. Furthermore, we demonstrate that ITS can be phylogenetically informative between species when moderate levels of IGPs are detected, but that ITS paralogy can interfere with population level studies. We caution against the routine use of ITS in phylogenetic studies of sponges without (1) screening for IGPs in specimens from every population sampled; (2) including all divergent paralogs in phylogenetic analyses; (3) testing ITS data using other single-copy, unlinked loci (such as nuclear introns).     

Spicule form in calcareous sponges (Porifera: Calcarea). The principle of uniplanar curvature

A study of the shapes of secondary sagittal tri- and quadri-radiate spicules of the calcareous sponge Leuconia fistulosa has revealed that the paired rays appear kinked, with straight sections in between the kinks, when the spicules are viewed along the optic axis of the constituent calcite. The secondary sagittal spicules thus conform with monaxons and with per-regular and primary sagittal tri- and quadri-radiates, in general, in that whenever a ray is curved, the curvature is always in a single plane that includes the optic axis. The kinks are not necessarily distributed in symmetrical fashion along the paired rays. A mechanism is proposed whereby uniplanar curvature could be achieved.     

First evidence of polychaete intermediate hosts for Neospirorchis spp. marine turtle blood flukes (Trematoda: Spirorchiidae)

Life cycles of spirorchiids that infect the vascular system of turtles are poorly understood. Few life cycles of these blood flukes have been elucidated and all intermediate hosts reported are gastropods (Mollusca), regardless of whether the definitive host is a freshwater or a marine turtle. During a recent survey of blood fluke larvae in polychaetes on the coast of South Carolina, USA, spirorchiid-like cercariae were found to infect the polychaetes Amphitrite ornata (Terebellidae) and Enoplobranchus sanguineus (Polycirridae). Cercariae were large, furcate, with a ventral acetabulum, but no eyespots were observed. Partial sequences of D1–D2 domains of the large ribosomal subunit, the internal transcribed spacer 2, and the mitochondrial cytochrome oxidase 1 genes allowed the identification of sporocysts and cercariae as belonging to two unidentified Neospirorchis species reported from the green turtle, Chelonia mydas, in Florida: Neospirorchis sp. (Neogen 13) in A. ornata and Neospirorchis sp. (Neogen 14) in E. sanguineus. Phylogenetic analysis suggests that infection of annelids by blood flukes evolved separately in aporocotylids and spirorchiids. Our results support the contention that the Spirorchiidae is not a valid family and suggest that Neospirorchis is a monophyletic clade within the paraphyletic Spirorchiidae. Since specificity of spirorchiids for their intermediate hosts is broader than it was thus far assumed, surveys of annelids in turtle habitats are necessary to further our understanding of the life history of these pathogenic parasites.