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Porifera, Demospongiae) Zoomorphology (2006) 125: 87-97 DO! 10.1007 js00435-006-00 l 5-z ORIGINAL ARTICLE Elda Gaino . Lidia Scalera Liaci . Margherita Sciscioli Giuseppe Corriero Investigation of the budding processin Tethya citrina and Tethya aurantium (Porifera, Demospongiae) Received: 2 lune 2004 j Accepted: 5 Aprii 2005 j Published online: 24 February 2006 © Springer-Verlag 2006 Abstract The budding process has been studied in two Introduction congeneric Mediterranean species belonging to Tethya from different sampling sites: Marsala and Venice La• Poriferan asexual reproduction by fragmentation, goons (Tethya citrina); Marsala Lagoon and Porto gemmules and buds, together with the constitutively Cesareo Basin (Tethya aurantium). Buds, connected to essenti al elements allowing fragments to reorganize and the adult by a spiculated stalk, differ between the two operate as independent organisms (Wulff 1991), repre• species in morphology and size, since those of T. citrina sents a significant reproductive strategy for some species. 'are small with elongated bodies, showing only a few These processes seem to occur to some extent in all or spicules protruding from the apical region, whereas most sponges (Fell 1993), thereby maximizing the dis• those of T. aurantium are round, larger, and show persal of these animals (Wulff 1995; Tsurumi and Rei• spicules radiating from the peripheral border. In T. ci• swig 1997; Maldonado and Uriz 1999). trina, cells with inclusions, varying in electron density The structure of the gemmules along with the and size, represent the main cell types of the buds. In hatching of a new specimen has been intensively studied T. aurantium, the cell component shows a major diver• (reviews by Simpson 1984 and Weissenfels 1989). Less is sification, resulting from spherulous cells, grey cells, known about the budding process, which typically oc• vacuolar cells and peculiar micro-vesicle cells. Neither curs in marine poriferans (review by Simpson 1984; canals nor choanocyte chambers were observed in the Boury-Esnault 1970; Chen et al. 1997; Corriero et al. buds of the two species. In T. citrina, bud production is 1998), and was first described in the freshwater species similar in both sampling sites. In T. aurantium, budding Radiospongilla cerebellata (Bowerbank 1863), by Saller occurs more rarely in Porto Cesareo Basin, probably in (1990) after experimental induction by injuring the mo• relation with environmental factors, such as the covering ther animaI. of the cortex by sediment and micro-algae. Finally, in Buds consist of cellular masses that sprout out from the buds of both species, the spicule size does not differ the surface of the adults and are able to develop into new from that of the cortex of the adult sponges, further functional individuals. The buds of Tethya were first supporting the main involvement of the cortex in orga• studied by Connes (1967, 1968) in a mixed population of nizing the skeletal architecture of the buds. T. citrina (Sarà and Melone 1965) and T. aurantium (Pallas 1766), where these species were confused in the Keywords Porifera' Tethya . Bud differentiation . entity T. lyncurium (Linné 1767). More recently, marked Spicules . Ultrastructure differences in the number of buds and in the lasting of this asexual process have been described in populations of these two species coexisting in the same environment (Corriero et al. 1996), without any interest in the process leading to bud differentiation. L. Scalera Liaci . M. Sciscioli . G. Corriero (IBI) Dipartimento di Zoologia dell'Università, Via Orabona 4, Tethya citrina and Tethya aurantium conform to the 70125, Bari, Ttaly typical organizational pattern of Tetractinomorpha. E-mail: [email protected] Among them, Tethydae are characterized by Te!.: + 39-80-5443357 Fax: + 39-80-5443357 well-defined cortex and a skeleton consisting of strongyloxeas arranged in bundles, different types of E. Gaino both spherasters and micrasters. According to the Dipartimento di Biolgia Cellulare ed Ambientale, exhaustive revision of Sarà (1987), the two species can Via Elce di Sotto, 06123, Perugia, Italy 88 be distinguished on the basis of colour, surface struc• ulations of T. citrina and T. aurantium, which both ture and spiculation. In particular, T. citrina is light colonize the rhizomes of the phanerogam Posidonia yellow in colour, has a thin cortical layer including oceanica (L.) Delile (Corriero et al. 1989). In both spe• oxyspherasters, and a single category of micrasters cies, the external sponge surface is usually free of sedi• without distinction between cortical and choanosomal ment and epibiontic organisms. The northern area of the elements. T. aurantium is orange in colour, has a strong Venice Lagoon (45°25'N; 12°26'E) is characterized by thick cortical layer including large spherasters, and two the presence of the Dese river which produces a marked categories of micrasters (cortical and choanosomal haline gradient (Sconfietti 1989). At the mouth of the elements). river, marine conditions prevail and the hard substrates In the present paper, we exami ne the various steps in are mainly colonized by different species of the Bivalvia, the process of bud formation in T. citrina and T. aur• Bryozoa and Porifera. Among 20 poriferan species antium, at the structural and ultrastructurallevels, with (Corriero et al. 2003), T. citrina is present with high the aim of verifying whether the budding process reflects density values, mainly colonizing vertical walls and a species-specific behaviour, and to what extent different small crevices between 3 and 5 m of depth (Corriero environmental conditions can affect this asexual repro• unpublished data). The external surface of the sponge duction in congeneric species. appears free of sediment, or, at most, partially covered From the bulk of data, it has emerged that buds by fine sediment which fills the depression among the significantly differ in their cytological and skeletal cortical tubercles. Epibiontic microalgae are lacking. organization, thereby proving that their differentiation is a species-specific processo Moreover, environmental conditions may locally enhance or depress this mecha• Methods nism of asexual reproduction. In arder to investigate the skeletal elements, spicule preparations were obtained from fragments of animals Material and methods as well as from whole buds dissected from adults. The material was treated by soaking it overnight in chloride Animals and sampling sites bleach (sodium hypochlorite) and then it was dissolved in 60% nitric acido After repeated washing in distilled The study was carri ed out at sites where rich budding water and in a graded series of ethylic alcohol, the iso• populations belonging to T. citrina (Sarà and Melone lated spicules were examined using an Olympus BH-2 1965) and T. aurantium (Pallas 1766) (Tetractinomor• light microscope. For each sampling site, buds were pha, Demospongia) have been previously observed counted and separated into two size classes between O (Corriero 1990; Corriero et al. 1996; Gaino et al. 2002). and 1.50 mm in diameter for T. citrina, and into five size Budding specimens of T. citrina and T. aurantium were classes between O and 3.75 mm in diameter for T. aur• collected in October and November 2001. T. citrina was antium. Spherasters and oxyspherasters of five buds sampled in Venice (Adriatic Sea) and Marsala Lagoons from each sampling site and from each bud size class (Tyrrhenian Sea, Narthern Sicily). T. aurantium was were counted and measured. For comparative analysis, sampled in the lagoons of Porto Cesareo Basin (Apulia, 50 spherasters and 50 oxyspherasters of both the cho• Ionian Sea) and Marsala. For each sampling site, five anosomal and corticallayers of each budding specimen budding specimens were randomly selected and sponges were measured. were removed from the substrate within 2 weeks of the For histological observations, the budding region of onset of the processo the mother sponge and the outermost part of its cho• Porto Cesareo Basin is a small, sheltered basin on the anosome were fixed in a solution of 10% formalin in south-western coast of Apulia (40° l 5'N; l7°54'E) with a filtered sea water, buffered with 0.1 N NaOH to a final maximum depth of 2.5 m. A wide channel system allows pH of7.6-7.8. Desilification ofthe samples was obtained considerable sea water inflow into the basin (Passeri by immersion in 5% hydrofluoric acid (H F) in filtered 1974). Among 42 poriferan species present, a rich pop• sea water for one and a half hours. Samples were then ulation of T. aurantium has been reported (Corriero dehydrated in an ethanol series and embedded in par• 1990; Mercurio 2000). The species colonizes hard hori• affino The 5-7 flm thick histological sections were zontally oriented substrates exposed to direct sunlight stained with haematoxylin-eosin or toluidine, and ob• and it is covered by a continuous layer of sediment served under a Leica microscope. mixed with a dense population of filamentous microal• For ultrastructural investigations (TEM, SEM), fix• gae. This layer, up to 5 mm thick, completely covers the ation was carried out for 2 h in 2.5% glutaraldehyde in a sponge's surface hiding its typical orange colour and the buffer of cacodylate (0.7 M) and filtered seawater to a cortical characters such as tubercles and lacunae. M ar• final pH of 7.4. For transmission electron microscopy sala is a large lagoon along the north-western coast of (TEM), selected material was post-fixed in 1% osmium Sicily (37°l4'N; l2°40'E), widely communicating with tetroxide (1 h) in the same buffer and desilificated the sea. The average depth is about l m. Forty-five (30 min) in 5% HP. Afterwards, samples were dehy• species of Porifera have been recognized with rich pop- drated in a graded ethanol series and embedded in an 89 epon-araldite mixture. Ultra-thin sections obtained with an LKB ultratome were contrasted with uranyl acetate and lead citrate, and examined with a Philips EM 208.
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