Suzanne C. Dufour

Gill Anatomy and the Evolution of Symbiosis in the Bivalve Family Thyasiridae

Among families of bivalves with chemoautotrophic symbionts, the Thyasiridae may vary the most in their anatomical characters and in the extent of their nutritional reliance upon symbionts. Since only a fraction of thyasirid species are symbiotic, and the symbionts are mostly observed to be extracellular, this group may be representative of early stages in the evolution of bacterium- bivalve symbioses. To better understand the distribution of symbiosis among thyasirid genera, and the relationships between gill structure and symbiont occurrence, the gills of 26 thyasirid species were studied by light and electron microscopy. Observations revealed three gill types, which are generally constrained within genera or subgenera. Symbionts were found in two gill types: the most simple, homorhabdic filibranch morphotype, and the most derived and thickened morphotype, which resembles the gill structure of other chemosymbiotic bivalves. In all observable cases, the symbionts were located extracellularly among the microvilli of the bacteriocytes. Among individuals of the species Thyasira (Parathyasira) equalis, the quantity of symbionts varied. The results suggest an evolutionary sequence: a homorhabdic filibranch gill structure with few symbionts among the epithelial cell microvilli eventually thickened abfrontally, and thereby offered a larger surface for colonization by symbionts. Eventually, the symbionts persisted and grew in vacuoles within epithelial cells.

Sulphide mining by the superextensile foot of symbiotic thyasirid bivalves

In a symbiotic association between an invertebrate host and chemoautotrophic bacteria, each partner has different metabolic requirements, and the host typically supplies the bacteria with necessary reduced chemicals (sulphide or methane). Some combination of anatomical, physiological and behavioural adaptations in the host often facilitates uptake and transport of reduced chemicals to the symbionts. We have studied five species of bivalve molluscs of the family Thyasiridae (that is, thyasirids) three of which harbour chemoautotrophic bacteria. Here we show that the symbiotic bivalves extend their feet to form elongated and ramifying burrows in the sediment, most probably to gain access to reduced sulphur. Closely related bivalves (including some thyasirid species) without bacterial symbionts show no comparable foot extension behaviour. The length and number of burrows formed by chemosymbiotic thyasirids are related to the concentration of hydrogen sulphide in the sediment. The burrows are formed by the foot of each bivalve, which can extend up to 30 times the length of the shell, and may be the most extreme case of animal structure elongation documented to date.

Evolutionary trajectories of a redundant feature: lessons from bivalve gill abfrontal cilia and mucocyte distributions

Abstract Recent data on the distributions of cilia and mucocytes on the bivalve gill abfrontal surface are analysed with respect to evolutionary relationships of the principal autobranch gill types. From the primitive function as a mucociliary cleaning surface in the protobranchs, two evolutionary trajectories are evident: (1) progressive reduction of both cilia and mucocytes with resultant loss of surface function, seen in the homorhabdic filibranchs studied; (2) reduction of cilia but retention or increase in acid mucopolysaccharide-secreting (AMPS) mucocyte density in the eulamellibranchs, corresponding to the assumption of a new function, probably in the reduction of frictional resistance to flow in the water canals. Heterorhabdic gill abfrontal surfaces present a mixture of these characteristics: reduction of cilia and mucocytes on the ordinary filaments, and retention of both on the principal filaments. The retention of AMPS mucocytes on the abfrontal surface of the pseudolamellibranchs may be related to the degree of interlamellar fusion, reducing frictional resistance to water flow as in the eulamellibranchs. The gill abfrontal surface thus constitutes an excellent candidate for the study of the different evolutionary options and trajectories of a redundant feature.

Trophic analyses of opportunistic polychaetes (Ophryotrocha cyclops) at salmonid aquaculture sites

A new species of dorvilleid polychaete, Ophryotrocha cyclops , has been observed on the rocky seafloor underneath deep salmonid aquaculture sites on the south coast of Newfoundland, Canada. The distribution of these opportunistic worms is likely related to organic matter accumulation on the seafloor, and this species may have a role in remediation processes. To better understand the functional role of O. cyclops at aquaculture sites, it is important to know what they feed upon. Here, stable isotope analyses (δ 13 C, δ 15 N and δ 34 S) and trace element analyses were performed on dorvilleids and their potential food sources at three aquaculture sites. Stable isotope analyses revealed spatial and temporal variation in the isotopic carbon signature of O. cyclops , highlighting possible differences in the food sources of individual dorvilleids within and between sites. The isotopic composition of dorvilleids was closest to that of fish pellets; the presence of abundant lipid droplets in gut epithelial cells of O. cyclops suggests the assimilation of fish pellet-derived lipids. Trace element analysis indicated that O. cyclops does not concentrate the aquaculture tracers Zn or Cu to a large extent. However, concentrations of sulphur were high in O. cyclops compared with other sources. Taken together, results show that O. cyclops most likely consume both fish pellets and flocculent matter-associated bacteria. As such, they are involved in sulphur cycling and fish pellet degradation at aquaculture sites.

Spermatogenesis in the archaic hydrothermal vent bivalve, Bathypecten vulcani, and comparison of spermatozoon ultrastructure with littoral pectinids

Summary Bathypecten vulcani is considered a relict species from the Paleozoic, based on shell characteristics such as the presence of calcite prisms. To date, it is the only pectinid species reported from hydrothermal ecosystems. Histological and ultrastructural studies show that spermatogenesis is identical to that of littoral pectinids. The spermatozoon has a 2.7 μm long pyriform head and a 40 μm flagellum. The four mitochondria of the mid-piece are about 1.2 μm in diameter. The nucleus contains dense chromatin fibres and possesses a wide, shallow (0.1 μm) anterior fossa and a narrow, deeper (0.2 μm) posterior nuclear fossa. Comparison of the ultrastructural characteristics of the spermatozoon of B. vulcani with those of littoral pectinids shows that they can be used as a diagnostic feature of this species. In particular, its acrosome characters will be a useful complement to the shell characters in the study of the phylogenetic position of this species in relation to other pectinids.

Spatial distribution of Alitta virens burrows in intertidal sediments studied by axial tomodensitometry

Relationships between sediment characteristics and the spatial organization of biogenic structures remain poorly understood, albeit important for characterizing bioturbation impacts and small-scale ecological patterns. Using axial tomodensitometry (CT-scan) and core sectioning, we studied the spatial distribution of Alitta virens burrows in sediment cores from two mud-flats with different degrees of exposure along the St Lawrence Estuary, Canada. A variety of burrow morphologies was observed at both sites, with most being I-shaped. Most values measured (organic matter content, mean tomographic intensity, the number and diameter of burrow shafts and the percentage of space they occupy per transverse section) covaried with depth. The more sheltered site had higher organic matter and mud contents, and lower average tomographic intensity values. The spatial distribution of burrow shafts also differed between sites, with the more sheltered site having a higher number of burrow shafts and percentage of biogenic space in the upper sediment column, as well as a greater volume of biogenic structures that were connected to the surface (although intra-site variability was great). Along the horizontal plane and at various depths within sediments, burrow shafts were more randomly distributed at the exposed site, compared to a more even, maximal spacing at the sheltered site. In addition to finding differences in burrow organization at two spatial scales, we found significant correlations between tomographic intensity and (1) the number of burrow shafts, (2) biogenic space and (3) organic matter content. CT-scan data, including tomographic intensity, are useful for examining and comparing biogenic structures in sediment cores.