Annie Mercier

Settlement preferences and early migration of the tropical sea cucumber Holothuria scabra.

Settlement and post-settlement processes of the sea cucumber Holothuria scabra Jaeger were studied in the laboratory. Independent and paired choice experiments revealed that several substrates could induce metamorphosis into pentactulae, but that specific substrates favoured settlement. Leaves of seagrass Thalassia hemprichii, with or without their natural bio-film, yielded the highest settlement rates (4.8-10.5%). T. hemprichii was preferred as a settlement substrate over sand, crushed coral, several other plant species and artificial seagrass leaves with or without a bio-film. Only settlement on the seagrass, Enhalus acoroides, was similar to that recorded for T. hemprichii. In the absence of a substrate, the larvae delayed settlement for nearly 96 h and survival was less than 0.5%. Sand and crushed coral, either alone or together, induced settlement from <1.5% of the available larvae. The pentactulae found on sand, coral and in bare containers were 10-35% smaller than those on T. hemprichii leaves. Soluble extracts from T. hemprichii and E. acoroides successfully induced metamorphosis and settlement on clean plastic surfaces. Newly settled juveniles remained on the seagrass leaves for 4-5 weeks before migrating to sand at around 6 mm in length. Prior to this, the juveniles spent 4-5 days moving on and off the leaves. Once on the sand, the juveniles became deposit-feeders, but did not show the typical burrowing behaviour of older specimens until they reached around 11 mm in length. The larvae of H. scabra appear to actively select seagrass leaves, possibly through chemical detection. We hypothesise that larvae settling on seagrass have an increased chance of growth and survival because they are provided with a suitable substrate on which to grow, and a bridge to sand substrates as they become deposit-feeders.

Periodic movement, recruitment and size-related distribution of the sea cucumber Holothuria scabra in Solomon Islands

Field studies of the sea cucumber Holothuria scabra conducted in Kogu Veke, Solomon Islands, showed monthly recruitment of newly-settled juveniles on seagrass and indicated that size distribution was a function of substratum type and depth. Adults >250 mm body length were found mainly on sand, with <5% organic matter (OM), at depths of >1–3 m. Individuals >10–250 mm were found mostly in 30–120 cm of water, on mud and muddy sand with OM content between 5 and 10%. Specimens >40–150 mm were also found in the intertidal zone, sometimes burrowed on exposed sandflats at low tide. Holothuria scabra avoided substrata of fine silt or shell and coral pebbles, and sediment with an organic content ≥30%. Juveniles ≤100 mm burrowed at sunrise and surfaced at sunset, whereas individuals >100 mm burrowed and surfaced a few hours earlier. Holothuria scabra tended to burrow when salinity decreased, whereas increased water temperatures reduced normal burrowing behaviour. Spatial distribution, observed during tank experiments, suggested that adult H. scabra aggregated prior to spawning and in response to the lunar cycle. The formation of pairs, trios or larger groups increased during the new moon and was most common just before the full moon. Newly-settled juveniles up to ca. 9 mm were found on seagrass leaves. Typically, maximum densities and smallest recruits were observed a couple of weeks after the full moon, lower densities and slightly larger recruits were found a few days later. Juveniles with a mean length around 65 mm released on sand moved less and grew faster than juveniles released in seagrass beds or on substrata of shells and crushed coral.

Internal brooding favours pre-metamorphic chimerism in a non-colonial cnidarian, the sea anemone Urticina felina

The concept of intraorganismal genetic heterogeneity resulting from allogeneic fusion (i.e. chimerism) has almost exclusively been explored in modular organisms that have the capacity to reproduce asexually, such as colonial ascidians and corals. Apart from medical conditions in mammals, the natural development of chimeras across ontogenetic stages has not been investigated in any unitary organism incapable of asexual propagation. Furthermore, chimerism was mainly studied among gregarious settlers to show that clustering of genetically similar individuals upon settlement promotes the occurrence of multi-chimeras exhibiting greater fitness. The possible occurrence of chimeric embryos and larvae prior to settlement has not received any attention. Here we document for the first time the presence of natural chimeras in brooded embryos and larvae of a unitary cnidarian, the sea anemone Urticina felina. Rates of visible bi- and multi-chimerism of up to 3.13 per cent were measured in the broods of 16 females. Apart from these sectorial chimeras, monitored fusion events also yielded homogeneous chimeric entities (mega-larvae) suggesting that the actual rates of natural chimerism in U. felina are greater than predicted by visual assessment. In support of this assumption, the broods of certain individuals comprised a dominant proportion (to 90%) of inexplicably large embryos and larvae (relative to oocyte size). Findings of fusion and chimerism in a unitary organism add a novel dimension to the framework within which the mechanisms and evolutionary significance of genetic heterogeneity in animal taxa can be explored.

Supplementary material from "Full allogeneic fusion of embryos in a holothuroid echinoderm"

Whole-body chimaeras (organisms composed of genetically distinct cells) have been directly observed in modular/colonial organisms (e.g. corals, sponges, ascidians); whereas in unitary deuterostosmes (including mammals) they have only been detected indirectly through molecular analysis. Here, we documented for the first time the step-by-step development of whole-body chimaeras in the holothuroid Cucumaria frondosa , a unitary deuterostome belonging to the phylum Echinodermata. To the best of our knowledge, this is the most derived unitary metazoan in which direct investigation of zygote fusibility has been undertaken. Fusion occurred among hatched blastulae, never during earlier (unhatched) or later (larval) stages. The fully fused chimaeric propagules were two to five times larger than non-chimaeric embryos. Fusion was positively correlated with propagule density and facilitated by the natural tendency of early embryos to agglomerate. The discovery of natural chimaerism in a unitary deuterostome that possesses large externally fertilized eggs provides a framework to explore key aspects of evolutionary biology, histocompatibility and cell transplantation in biomedical research.