No abstract

Fusion trials between metamorphs of the aplousobranch compound ascidian Diplosoma listerianum indicated that chimera formation was not dependent on relatedness. Similar, high rates of union were observed between full siblings, half siblings, unrelated individuals from the same population, and individuals from two geographically distant localities. This is in contrast to the well-studied ascidian genus Botryllus, in which a highly polymorphic allorecognition system governing the fusion^non-fusion reaction (colony speci¢city) largely limits fusion to close relatives. Fusion in Botryllus establishes a vascular chimera throughout which stem cells may circulate, promoting cell lineage competition between the fusion partners. The restriction of fusion to close kin in Botryllus is thought to reduce the inclusive ¢tness costs of competitive interactions between cell lineages within the chimera. In contrast to Botryllus, modules (zooids) of a D. listerianum colony are not interlinked by blood vessels, seemingly precluding the exchange of stem cells. The apparent absence of strict colony speci¢city in D. listerianum is thus in keeping with the predictions of the Botryllus model for the maintenance of allorecognition polymorphism.However, colony speci¢city has been reported in other species of aplousobranch ascidian that also lack a common vascular system. In these, the threat of migrating blood-borne stem cells cannot be responsible for the presence of colony speci¢city. One possibility, requiring experimental investigation, is that stem cells could perhaps migrate between zooids by another route, such as through the matrix of the colonial tunic. Even in the absence of stem cell exchange, cheating on the costs of colony maintenance and defence could also produce selective forces favouring colony speci¢city. In compound ascidians, this could involve unequal contribution to extrazooidal structures, principally the tunic and related tissues. This consideration seems potentially relevant to the lack of discrimination during fusion in D. listerianum, since extrazooidal somatic investment in this species appears minimal, severely limiting the scope for this other form of cheating.The various possible modes of exploitative interaction between fused colonies are not mutually exclusive, and o¡er fundamentally similar explanations for colony speci¢city. If none of them can be shown to occur in non-botryllid species possessing colony speci¢city, the generality of the Botryllus model may require re-evaluation.

Many marine invertebrates bud vegetatively to produce a modular colony of individuals derived from a single zygote. Fusion of different colonies to produce a genetically composite entity (a chimera) is known from experiments on sponges, hydroids, corals, bryozoans and ascidians – groups which together dominate sessile faunas on marine hard substrates. Random amplified polymorphic DNA–polymerase chain reaction (RAPD–PCR) analysis was applied to individual modules (zooids) dissected from colonies of a colonial ascidian, Diplosoma listerianum (Milne Edwards), to investigate the presence and extent of chimerism. The technique revealed chimerism in wild material. In total, 288 colonies from eight different natural populations were analysed. Chimeric colonies were present in all populations, at frequencies of up to 61%, with up to six different genotypes present in some colonies. Zooids of different genetic origin often intermingled within a chimeric colony to produce a zooidal mosaic. Although fusion of colonies has been observed directly in the laboratory, an unknown proportion of the chimerism detected in wild populations might have arisen through somatic mutation. To assess this possibility, tissue of 12 clones in culture was sampled repeatedly over a period of 3 years and subjected to RAPD–PCR analysis. RAPD banding patterns were generally very stable; the changes noted mostly involved minor bands that would not, on their own, have been taken as evidence for chimerism under the conservative criteria adopted for the study of wild populations. It was concluded that a large proportion of natural chimerism is attributable to colony fusion. © 2003 The Linnean Society of London. Biological Journal of the Linnean Society, 2003, 79, 183–192.

The importance of sexual compatibility between mates has only recently been realized in zoological research into sexual selection, yet its study has been central to botanical research for many decades. The reproductive characteristics of remote mating, an absence of precopulatory mate screening, internal fertilization and embryonic brooding are shared between passively pollinated plants and a phylogenetically diverse group of sessile aquatic invertebrates. Here, we further characterize the sexual compatibility system of one such invertebrate, the colonial ascidian Diplosoma listerianum. All 66 reciprocal pairings of 12 genetic individuals were carried out. Fecundities of crosses varied widely and suggested a continuous scale of sexual compatibility. Of the 11 animals from the same population c. 40% of crosses were completely incompatible with a further c. 20% having obvious partial compatibility (reduced fecundity). We are unaware of other studies documenting such high levels of sexual incompatibility in unrelated individuals. RAPD fingerprinting was used to estimate relatedness among the 12 individuals after a known pedigree was successfully reconstructed to validate the technique. In contrast to previous results, no correlation between genetic similarity and sexual compatibility was detected. The blocking of many genotypes of sperm is expected to severely modify realized paternity away from ‘fair raffle’ expectations and probably reduce levels of intra‐brood genetic diversity in this obligatorily promiscuous mating system. One adaptive benefit may be to reduce the bombardment of the female reproductive system by outcrossed sperm with conflicting evolutionary interests, so as to maintain female control of somatic : gametic investment.