While components of the pathway that establishes left-right asymmetry have been identified in diverse animals, from vertebrates to flies, it is striking that the genes involved in the first symmetry-breaking step remain wholly unknown in the most obviously chiral animals, the gastropod snails. Previously, research on snails was used to show that left-right signaling of Nodal, downstream of symmetry breaking, may be an ancestral feature of the Bilateria [1, 2]. Here, we report that a disabling mutation in one copy of a tandemly duplicated, diaphanous-related formin is perfectly associated with symmetry breaking in the pond snail. This is supported by the observation that an anti-formin drug treatment converts dextral snail embryos to a sinistral phenocopy, and in frogs, drug inhibition or overexpression by microinjection of formin has a chirality-randomizing effect in early (pre-cilia) embryos. Contrary to expectations based on existing models [3-5], we discovered asymmetric gene expression in 2- and 4-cell snail embryos, preceding morphological asymmetry. As the formin-actin filament has been shown to be part of an asymmetry-breaking switch in vitro [6, 7], together these results are consistent with the view that animals with diverse body plans may derive their asymmetries from the same intracellular chiral elements .
BACKGROUND: Various shapes of gastropod shells have evolved ever since the Cambrian. Although theoretical analyses of morphogenesis exist, the molecular basis of shell development remains unclear. We compared expression patterns of the decapentaplegic (dpp) gene in the shell gland and mantle tissues at various developmental stages between coiled-shell and non-coiled-shell gastropods. RESULTS: We analyzed the expression patterns of dpp for the two limpets Patella vulgata and Nipponacmea fuscoviridis, and for the dextral wild-type and sinistral mutant lineage of the pond snail Lymnaea stagnalis. The limpets had symmetric expression patterns of dpp throughout ontogeny, whereas in the pond snail, the results indicated asymmetric and mirror image patterns between the dextral and sinistral lineages. CONCLUSION: We hypothesize that Dpp induces mantle expansion, and the presence of a left/right asymmetric gradient of the Dpp protein causes the formation of a coiled shell. Our results provide a molecular explanation for shell, coiling including new insights into expression patterns in post-embryonic development, which should aid in understanding how various shell shapes are formed and have evolved in the gastropods.
Creating surfaces capable of resisting liquid-mediated adhesion is extremely difficult due to the strong capillary forces that exist between surfaces. Land snails use this to adhere to and traverse across almost any type of solid surface of any orientation (horizontal, vertical or inverted), texture (smooth, rough or granular) or wetting property (hydrophilic or hydrophobic) via a layer of mucus. However, the wetting properties that enable snails to generate strong temporary attachment and the effectiveness of this adhesive locomotion on modern super-slippy superhydrophobic surfaces are unclear. Here we report that snail adhesion overcomes a wide range of these microscale and nanoscale topographically structured non-stick surfaces. For the one surface which we found to be snail resistant, we show that the effect is correlated with the wetting response of the surface to a weak surfactant. Our results elucidate some critical wetting factors for the design of anti-adhesive and bio-adhesion resistant surfaces.
BACKGROUND: The risks of fish-borne zoonotic trematodes (FZT) to human health constitute an important problem in Vietnam. The infection of humans with these trematodes, such as small liver trematodes (Clonorchis sinensis and Opisthorchis viverrini), intestinal trematodes (Heterophyidae) and others is often thought to be linked to fish culture in areas where the habit of eating raw fish is common. Juvenile fish produced in nurseries are often heavily infected with FZT and since fishes are sold to aquaculture facilities for growth, control of FZT in these fishes should be given priority. Controlling the first intermediate host (i.e., freshwater gastropods), would be an attractive approach, if feasible. The black carp, Mylopharyngodon piceus, is a well-known predator of freshwater snails and is already used successfully for biological control of snails in various parts of the world including Vietnam. Here we report the first trials using it for biological control of intermediate host snails in nursery ponds stocked with 1-week old fry (10–12 mm in length) of Indian carp, Labeo rohita. METHODS: Semi-field and field experiments were set up to test the effect of black carp on snail populations. In the semi-field experiment a known quantity of snails was initially introduced into a pond which was subsequently stocked with black carp. In the field trial in nursery ponds, density of snails was estimated prior to a nursing cycle and at the end of the cycle (after 9 weeks). RESULTS: The results showed that black carp affect the density of snail populations in both semi-field and field conditions. The standing crop of snails in nursery ponds, however, was too high for 2 specimens to greatly reduce snail density within the relatively short nursing cycle. CONCLUSIONS: We conclude that the black carp can be used in nursery ponds in Northern Vietnam for snail control. Juvenile black carp weighing 100 - 200g should be used because this size primarily prey on intermediate hosts of FZT and other studies have shown that it does not prey on fish fry of other species. It may be necessary to use a high stocking density of black carp or to reduce snail density in the nursery ponds using other measures (e.g. mud removal) prior to stocking fry in order for the black carp to keep the density of intermediate host snails at a very low level.
Animals have evolved many ways to enhance their own reproductive success. One bizarre sexual ritual is the love dart shooting of helicid snails, which has courted many theories regarding its precise function. Acting as a hypodermic needle, the dart transfers an allohormone that increases paternity success. Its precise physiological mechanism of action within the recipient snail is to close off the entrance to the sperm digestion organ via a contraction of the copulatory canal, thereby delaying the digestion of most donated sperm. In this study, we used the common garden snail Cornu aspersum to identify the allohormone that is responsible for this physiological change in the female system of this simultaneous hermaphrodite. The love dart allohormone (LDA) was isolated from extracts derived from mucous glands that coat the dart before it is stabbed through the partners body wall. We isolated LDA from extracts using bioassay-guided contractility measurement of the copulatory canal. LDA is encoded within a 235 amino acid precursor protein containing multiple cleavage sites that, when cleaved, releases multiple bioactive peptides. Synthetic LDA also stimulated copulatory canal contractility. Combined with our finding that the protein sequence resembles previously described molluscan buccalin precursors, this indicates that LDA is partially conserved in helicid snails and less in other molluscan species. In summary, our study provides the full identification of an allohormone that is hypodermically injected via a love dart. More importantly, our findings have important consequences for understanding reproductive biology and the evolution of alternative reproductive strategies.
Although diet is believed to be a major factor underlying the evolution of venom, few comparative studies examine both venom composition and diet across a radiation of venomous species. Cone snails within the family, Conidae, comprise more than 700 species of carnivorous marine snails that capture their prey by using a cocktail of venomous neurotoxins (conotoxins or conopeptides). Venom composition across species has been previously hypothesized to be shaped by (a) prey taxonomic class (i.e., worms, molluscs, or fish) and (b) dietary breadth. We tested these hypotheses under a comparative phylogenetic framework using ecological data from past studies in conjunction with venom duct transcriptomes sequenced from 12 phylogenetically disparate cone snail species, including 10 vermivores (worm-eating), one molluscivore, and one generalist.
The parasitic nematode Angiostrongylus cantonensis is a major cause of eosinophilic meningitis in humans, and has been documented in other incidental hosts such as birds, horses, dogs and non-human primates. It is endemic in Hawaii, and there have been sporadic reports in the southern continental United States. This parasite uses rats as definitive hosts and snails as intermediate hosts. In this study, we collected potential definitive and intermediate hosts throughout Florida to ascertain the geographic distribution in the state: Rats, environmental rat fecal samples, and snails were collected from 18 counties throughout the state. Classical diagnostics and morphological identification, along with molecular techniques were used to identify nematode species and confirm the presence of A. cantonensis. Of the 171 Rattus rattus collected, 39 (22.8%) were positive for A. cantonensis, and 6 of the 37 (16.2%) environmental rat fecal samples collected in three of the surveyed counties were also positive for this parasite by real time PCR. We examined 1,437 gastropods, which represented 32 species; 27 (1.9%) were positive for A. cantonensis from multiple sites across Florida. Three non-native gastropod species, Bradybaena similaris, Zachrysia provisoria, and Paropeas achatinaceum, and three native gastropod species, Succinea floridana, Ventridens demissus, and Zonitoides arboreus, which are newly recorded intermediate hosts for the parasite, were positive for A. cantonensis. This study indicates that A. cantonensis is established in Florida through the finding of adult and larval stages in definitive and intermediate hosts, respectively, throughout the state. The ability for this historically subtropical nematode to thrive in a more temperate climate is alarming, however as the climate changes and average temperatures rise, gastropod distributions will probably expand, leading to the spread of this parasite in more temperate areas. Through greater awareness of host species and prevalence of A. cantonensis in the United States, potential accidental infections may be avoided.
In vertebrates, the left-and-right pairs of homologous organs are generally present in equal numbers. A remarkable exception is snail-eating snakes in the family Pareidae: almost all the pareid snakes have much more teeth on the right mandible than on the left for functional specialization in feeding on the dextral majority of land snails. Because the only exceptional species with symmetric dentition has been regarded as a slug-eater, the extent of dietary specialization on slugs could shape the degree of the lateral asymmetry of mandibular dentition (dentition asymmetry) even among snail eaters.
Predator-prey interactions are major processes promoting phenotypic evolution. However, it remains unclear how predation causes morphological and behavioural diversity in prey species and how it might lead to speciation. Here, we show that substantial divergence in the phenotypic traits of prey species has occurred among closely related land snails as a result of adaptation to predator attacks. This caused the divergence of defensive strategies into two alternatives: passive defence and active defence. Phenotypic traits of the subarctic Karaftohelix land snail have undergone radiation in northeast Asia, and distinctive morphotypes generally coexist in the same regions. In these land snails, we documented two alternative defence behaviours against predation by malacophagous beetles. Furthermore, the behaviours are potentially associated with differences in shell morphology. In addition, molecular phylogenetic analyses indicated that these alternative strategies against predation arose independently on the islands and on the continent suggesting that anti-predator adaptation is a major cause of phenotypic diversity in these snails. Finally, we suggest the potential speciation of Karaftohelix snails as a result of the divergence of defensive strategies into passive and active behaviours and the possibility of species radiation due to anti-predatory adaptations.
Molluscs (snails, octopuses, clams and their relatives) have a great disparity of body plans and, among the animals, only arthropods surpass them in species number. This diversity has made Mollusca one of the best-studied groups of animals, yet their evolutionary relationships remain poorly resolved. Open questions have important implications for the origin of Mollusca and for morphological evolution within the group. These questions include whether the shell-less, vermiform aplacophoran molluscs diverged before the origin of the shelled molluscs (Conchifera) or lost their shells secondarily. Monoplacophorans were not included in molecular studies until recently, when it was proposed that they constitute a clade named Serialia together with Polyplacophora (chitons), reflecting the serial repetition of body organs in both groups. Attempts to understand the early evolution of molluscs become even more complex when considering the large diversity of Cambrian fossils. These can have multiple dorsal shell plates and sclerites or can be shell-less but with a typical molluscan radula and serially repeated gills. To better resolve the relationships among molluscs, we generated transcriptome data for 15 species that, in combination with existing data, represent for the first time all major molluscan groups. We analysed multiple data sets containing up to 216,402 sites and 1,185 gene regions using multiple models and methods. Our results support the clade Aculifera, containing the three molluscan groups with spicules but without true shells, and they support the monophyly of Conchifera. Monoplacophora is not the sister group to other Conchifera but to Cephalopoda. Strong support is found for a clade that comprises Scaphopoda (tusk shells), Gastropoda and Bivalvia, with most analyses placing Scaphopoda and Gastropoda as sister groups. This well-resolved tree will constitute a framework for further studies of mollusc evolution, development and anatomy.