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Biomedical subjects

M Schilthuizen

Publications and source records attributed to M Schilthuizen.

9 recordsLinked to original sources

The convoluted evolution of snail chirality.

The direction that a snail (Mollusca: Gastropoda) coils, whether dextral (right-handed) or sinistral (left-handed), originates in early development but is most easily observed in the shell form of the adult. Here, we review recent progress in understanding snail chirality from genetic, developmental and ecological perspectives. In the few species that have been characterized, chirality is determined by a single genetic locus with delayed inheritance, which means that the genotype is expressed in the mother's offspring. Although research lags behind the studies of asymmetry in the mouse and nematode, attempts to isolate the loci involved in snail chirality have begun, with the final aim of understanding how the axis of left-right asymmetry is established. In nature, most snail taxa (>90%) are dextral, but sinistrality is known from mutant individuals, populations within dextral species, entirely sinistral species, genera and even families. Ordinarily, it is expected that strong frequency-dependent selection should act against the establishment of new chiral types because the chiral minority have difficulty finding a suitable mating partner (their genitalia are on the 'wrong' side). Mixed populations should therefore not persist. Intriguingly, however, a very few land snail species, notably the subgenus Amphidromus sensu stricto, not only appear to mate randomly between different chiral types, but also have a stable, within-population chiral dimorphism, which suggests the involvement of a balancing factor. At the other end of the spectrum, in many species, different chiral types are unable to mate and so could be reproductively isolated from one another. However, while empirical data, models and simulations have indicated that chiral reversal must sometimes occur, it is rarely likely to lead to so-called 'single-gene' speciation. Nevertheless, chiral reversal could still be a contributing factor to speciation (or to divergence after speciation) when reproductive character displacement is involved. Understanding the establishment of chirality, the preponderance of dextral species and the rare instances of stable dimorphism is an important target for future research. Since the genetics of chirality have been studied in only a few pulmonate species, we also urge that more taxa, especially those from the sea, should be investigated.

Animal Structures↗

Population structure and coil dimorphism in a tropical land snail.

Tree snails of the subgenus Amphidromus s. str. are unusual because of the chiral dimorphism that exists in many species, with clockwise (dextrally) and counter-clockwise (sinistrally) coiled individuals co-occurring in the same population. Given that mating in snails is normally impeded when the two partners have opposite coil, positive frequency-dependent selection should prevent such dimorphism from persisting. We test the hypothesis that a strong population structure with little movement between tree-based demes may result in the fixation of coiling morphs at a very small spatial scale, but apparent dimorphism at all larger scales. To do so, we describe the spatial structure in a Malaysian population of A. inversus (Müller, 1774) with 36% dextrals. We marked almost 700 juvenile and adult snails in a piece of forest consisting of 92 separate trees, and recorded dispersal and the proportions of dextrals and sinistrals in all trees over a 7-day period. We observed frequent movement between trees (155 events), and found that no trees had snail populations with proportions of dextrals and sinistrals that were significantly different from random. Upon recapture 1 year later, almost two-thirds of the snails had moved away from their original tree. We conclude that population structure alone cannot stabilise the coil dimorphism in Amphidromus.

Animal Structures↗

The 'rare allele phenomenon' in a ribosomal spacer.

We describe the increased frequency of a particular length variant of the internal transcribed spacer 1 (ITS-1) of the ribosomal DNA in a hybrid zone of the land snail Albinaria hippolyti. The phenomenon that normally rare alleles or other markers can increase in frequency in the centre of hybrid zones is not new. Under the term 'hybrizyme' or 'rare allele' phenomenon it has been recorded in many organisms and different genetic markers. However, this is the first time that it has been found in a multicopy locus. On the one hand, the pattern fits well with the view that purifying selection in hybrid populations works on many loci across the genome and should thus have its effect on many independent molecular markers. On the other hand, the results are puzzling, given that the multiple copies of rDNA are not expected to respond in unison. We suggest two possible explanations for these conflicting observations.

Alleles↗

Dualism and conflicts in understanding speciation.

Speciation is a central but elusive issue in evolutionary biology. Over the past sixty years, the subject has been studied within a framework conceived by Ernst Mayr and Theodosius Dobzhansky and subsequently developed further by numerous other workers. In this "isolation" theory, the evolution of reproductive isolation is a key element of speciation; natural selection is given only secondary importance while gene flow is considered prohibitive to the process. In this paper, I argue that certain elements in this approach have produced confusion and irreconcilability among students of speciation. The more prominent debates in speciation (i.e., the species definition, sympatry/allopatry, and the role of reinforcement) all derive from an inherent conflict between the "isolation" theory and Darwin's "selection" view on species and speciation (in which disruptive selection is crucial). New data, mainly from field ecology, molecular population genetics, laboratory studies with Drosophila and computer analysis, all suggest that the isolation theory may no longer be the most desirable vantage point from which to explore speciation. Instead, environmental selection in large populations, often unimpeded by ongoing gene flow, appears to be the decisive element. The traditional preoccupation with reproductive isolation has created gaps in our knowledge of several crucial issues, mainly regarding the role of environmental selection and its connection with mate selection.

Animals↗

Screening mollusks for Wolbachia infection.

We screened 38 species of mollusks for infection by Wolbachia, a bacterium that is a common endosymbiont in arthropods, where it induces alterations in reproduction. Using a PCR assay, we could not detect the symbiont in any of the samples, indicating that, in mollusks, it might be absent.

Animals↗

Horizontal transmission of parthenogenesis-inducing microbes in Trichogramma wasps.

Complete parthenogenesis (thelytoky) in species of the parasitic wasp Trichogramma is usually caused by the cytoplasmically inherited bacterium Wolbachia. This symbiont induces gamete duplication, which, in these haplodiploid organisms, results in all-female broods. Antibiotic treatment "cures' this condition, restoring normal sexual reproduction. Phylogenetic analysis of Wolbachia has shown that, in contrast with the strains in other host organisms (where the symbiont also induces different reproductive alterations), those in Trichogramma form a monophyletic group. This might be an indication of symbiont-host cocladogenesis. To test this, we performed comparative molecular phylogenetics on 20 parthenogenetic Trichogramma cultures and their Wolbachiae. We conclude that there is, in fact, little evidence for cocladogenesis. Instead, the phylogenetic distribution of the symbionts appears to result from occasional horizontal transmission, which probably takes place inside the hosts of Trichogramma parasitoids (usually lepidopteran eggs). This study therefore suggests that parthenogenesis is not only curable, it can sometimes be contagious also.

Alphaproteobacteria↗

Phylogenetic relationships inferred from the sequence and secondary structure of ITS1 rRNA in Albinaria and putative Isabellaria species (Gastropoda, Pulmonata, Clausiliidae).

An analysis of the ITS1 sequence variation among five species of terrestrial pulmonate snails was performed to decide between two conflicting hypotheses concerning the phylogeny of these anatomically similar gastropods. It turned out that the so-called genus Isabellaria is a polyphyletic entity; the diagnostic, apomorphic structure of its clausilial apparatus, enabling a nearly complete obstruction of the shell aperture with the animal at rest, apparently evolved more than once from ancestors currently classified with the speciose genus Albinaria. The classification based on general shell shape and sculpture, and distributional patterns, turns out to be the natural one. This study also provides the first data on ITS1 sequences in gastropods. The recently published ITS1 sequence of another molluscan species, a bivalve, is quite different but similar in length to that of the snails.

Animals↗