Role of chemical signalling in release of motor programs during embryogenesis of freshwater snails Lymnaea stagnalis and Helisoma trivolvis

We have earlier found that freshwater pond snails Helisoma trivolvis and Lymnaea stagnalis, when reared under conditions of starvation, release chemical signals that reversibly suppress larval development of conspecific embryos. Here, we report that (i) these signals are not strictly conspecific and affect also embryos of a closely related species, which occupies a similar environmental niche; (ii) besides the development of embryos, the signals also affect the release of main motor programs, such as locomotion, feeding, and cardiac activity; (iii) action of the signals is bidirectional: they retard the development and release of motor programs at the early larval stages (trochophore to veliger) and accelerate them at later stages (late veliger to hatching). A possible adaptive significance of the described phenomena is discussed.     

Role of chemical signalling in release of motor programs during embryogenesis of freshwater snails <Emphasis Type="Italic">Lymnaea stagnalis</Emphasis> and <Emphasis Type="Italic">Helisoma trivolvis</Emphasis>

We have earlier found that freshwater pond snails Helisoma trivolvis and Lymnaea stagnalis, when reared under conditions of starvation, release chemical signals that reversibly suppress larval development of conspecific embryos. Here, we report that (i) these signals are not strictly conspecific and affect also the embryos of a closely related species, which occupies a similar environmental niche; (ii) besides the development of embryos, the signals also affect the release of main motor programs, such as locomotion, feeding, and cardiac activity; (iii) action of the signals is bidirectional: they retard the development and release of motor programs at the early larval stages (trochophore to veliger) and accelerate them at later stages (late veliger to hatching). A possible adaptive significance of the described phenomena is discussed.     

The suitability of several aquatic snails as intermediate hosts for Angiostrongylus cantonensis.

Sixteen species of aquatic snails of four families were tested by quantitative technique under standardized conditions for their suitability as intermediate hosts for Angiostrongylus cantonensis. These species were the planorbid snails Biomphalaria glabrata, Biomphalaria alexandrina, Planorbis planorbis, Planorbis intermixtus, Bulinus truncatus, Bulinus contortus, Bulinus africanus, Bulinus tropicus and Helisoma sp.; the lymnaeid snails Lymnaea natalensis, Lymnaea tomentosa, Lymnaea stagnalis, and Stagnicola elodes; the physid snail Physa acuta (an Egyptian and a German strain) and the ampullariid snails Marisa cornuarietis and Lanistes carinatus. All these snail species proved to be susceptible to infection with A. cantonensis, and first stage larvae reached the infective third stage in all of them. However, the rate and intensity of infection varied with different species. B. glabrata was the most susceptible snail species with a 100% infection rate and an average percentage recovery of third stage larvae of 26.1. This was followed by S. elodes and B. africanus, with a 100% infection rate and an average percentage recovery of third stage larvae of 15.6 and 14.6 respectively. The rest of snail species proved to be less susceptible. For comparative evaluation of the suitability of the various snail species as intermediate hosts of A. cantonensis a "Capacity Index" was determined. This index should provide a useful method for the evaluation of the suitability of various snails as intermediate hosts of nematode parasites under standardized conditions in the laboratory.     

The presence of predators modifies the larval development of Fasciola hepatica in surviving Lymnaea truncatula

Experimental infections of Lymnaea truncatula with Fasciola hepatica were performed to study the consequences of the presence of predators (sciomyzid larvae or zonitid snails) on the characteristics of larval F. hepatica development in surviving snails. Controls consisted of infected snails that were not subjected to predators. Compared to controls, the survival rate at day 30 post-exposure, the duration of cercarial shedding, and the number of cercariae shed by surviving snails were significantly lower when predators were present in snail breeding boxes, whatever the type of predator used. In contrast, the prevalences of Fasciola infections in snails, and the length of time between exposure and the onset of cercarial shedding showed no significant variation. The progressive development of a stress reaction in surviving snails against predators during the first 30 days of experimental exposure to F. hepatica would influence snail survival during the cercarial shedding period and, consequently, the number of cercariae shed by the snails.     

Yersiniae in pond water and snails

A Yersinia enterocolitica serotype 9 was isolated from pond water; Y. enterocolitica-like bacteria were also isolated from pond water and from three species of snails (Lymnaea palustris elodes, Helisoma sp., Oxyloma retusa) from the Edwin S. George Reserve in southeastern Michigan. There was evidence for biochemical stability among some of the organisms over a period of years. There also was evidence of transmission of these organisms to snails from the water.     

Persistent larval sensory neurones are required for the normal development of the adult sensory afferent projections in Drosophila

We have tested the hypothesis that larval neurones guide growth of adult sensory axons in Drosophila. We show that ablation of larval sensory neurones causes defects in the central projections of adult sensory neurones. Spiralling axons and ectopic projections indicate failure in axon growth guidance. We show that larval sensory neurones are required for peripheral pathfinding, entry into the CNS and growth guidance within the CNS. Ablation of subsets of neurones shows that larval sensory neurones serve specific guidance roles. Dorsal neurones are required for axon guidance across the midline, whereas lateral neurones are required for posterior growth. We conclude that larval sensory neurones pioneer the assembly of sensory arrays in adults.     

A comparative study of mechanisms underlying digenean-snail specificity: in vitro interactions between hemocytes and digenean larvae

A panel of 4 digenetic trematode species (Echinostoma paraensei, E. trivolvis, Schistosoma mansoni, and Schistosomatium douthitti) and 5 snail species (Biomphalaria glabrata, Helisoma trivolvis, Lymnaea stagnalis, Stagnicola elodes, and Helix aspersa) was examined to determine if known patterns of host specificity could be explained by the tendency of digenean larvae to be bound by snail hemocytes, or by the ability of larvae to influence the spreading behavior of hemocytes. In short-term (1 hr) in vitro adherence assays, there was no overall pattern to suggest that sporocysts were more likely to be bound by hemocytes from incompatible than compatible snails. Compared with the other parasites, sporocysts of E. paraensei were less likely to be bound by hemocytes from any of the snail species tested. All rediae examined, including those of another species Echinoparyphium sp., were also remarkably refractory to binding by hemocytes from any of the snails. Of all the larvae examined, only sporocysts and young daughter rediae of E. paraensei caused hemocytes to round up in their presence. This was true for hemocytes from the compatible species B. glabrata and the incompatible lymnaeid species S. elodes and L. stagnalis. The patterns of host specificity shown by this particular panel of parasites and snails were not predicted by either the extent of hemocyte adherence to digenean larvae or by the ability of larvae to affect hemocyte spreading behavior. The results of this study suggest that a role for hemocytes, although likely, may require different assays, possibly of a more prolonged nature, for its detection. Also, different parasite species (notably E. paraensei) and intramolluscan stages have distinctive interactions with host hemocytes, suggesting that the determinants of specificity vary with the host-parasite combination, and with the parasite life cycle stage.     

Postembryonic development of the auditory system of the cicada Okanagana rimosa (Say) (Homoptera: Auchenorrhyncha: Cicadidae)

Cicadas (Homoptera: Auchenorrhyncha: Cicadidae) use acoustic signalling for mate attraction and perceive auditory signals by a tympanal organ in the second abdominal segment. The main structural features of the ear are the tympanum, the sensory organ consisting of numerous scolopidial cells, and the cuticular link between sensory neurones and tympanum (tympanal ridge and apodeme). Here, a first investigation of the postembryonic development of the auditory system is presented. In insects, sensory neurones usually differentiate during embryogenesis, and sound-perceiving structures form during postembryogenesis. Cicadas have an elongated and subterranian postembryogenesis which can take several years until the final moult. The neuroanatomy and functional morphology of the auditory system of the cicada Okanagana rimosa (Say) are documented for the adult and the three last larval stages. The sensory organ and the projection of sensory afferents to the CNS are present in the earliest stages investigated. The cuticular structures of the tympanum, the tympanal frame holding the tympanum, and the tympanal ridge differentiate in the later stages of postembryogenesis. Thus, despite the different life styles of larvae and adults, the neuronal components of the cicada auditory system develop already during embryogenesis or early postembryogenesis, and sound-perceiving structures like tympana are elaborated later in postembryogenesis. The life cycle allows comparison of cicada development to other hemimetabolous insects with respect to the influence of specially adapted life cycle stages on auditory maturation. The neuronal development of the auditory system conforms to the timing in other hemimetabolous insects.     

Effects of diet and larval trematode parasitism on lutein and beta-carotene concentrations in planorbid snails as determined by quantitative high performance reversed phase thin layer chromatography

High performance thin layer chromatography (HPTLC) was used to quantify the concentrations of beta-carotene and lutein in Biomphalaria glabrata and Helisoma trivolvis (Colorado and Pennsylvania strains) snails under various conditions. These conditions were: snails fed a lettuce (L) vs. a yolk (Y) diet; B. glabrata infected with Echinostoma caproni vs. uninfected snails; and H. trivolvis (PA) infected with Echinostoma trivolvis vs. uninfected snails. The pigments were extracted from the snail whole bodies and digestive gland-gonad complexes, separated by reversed phase HPTLC, and quantified by densitometric scanning with standard calibration curves. Snails on the L-diet showed significant increases (Student's t-test, P<0.05) in the concentrations of beta-carotene and lutein compared to snails on the Y-diet. Snails infected with echinostomes showed no significant differences (Student's t-test, P>0.05) in the concentrations of lutein and beta-carotene compared to the uninfected cohorts. Our results were compared with previous studies that analyzed beta-carotene and lutein in snails infected with larval trematodes. Variations in the results of our study compared with others reflect intrinsic differences in the larval trematode-snail systems used.     

Relative vulnerability of six freshwater gastropods to the leech Nephelopsis obscura (Verrill)

SUMMARY. 1. The vulnerability of six species of freshwater snails to the leech Nephelopsis obscura was determined in laboratory predation experiments.
2. Nephelopsis was unable to prey on prosobranch snails with an operculum, but did consume certain pulmonate species though predation rates were low, ranging from one to two snails per leech per night at 20°C. Apparently, Nephelopsis does not actively select prey, but merely consumes those species most easily handled.
3. Among pulmonates, leeches captured species in the following order: Physa gyrina (Say), Helisoma anceps (Menke). Lymnaea emarginata (Say) and Helisoma trivolvis (Say). Susceptibihty of size classes varied among snail species.
4. Neither environmental structure, such as cobble or macrophytes, nor gastropod escape behaviors, such as clamping to substrates or vertical migration away from bottom-feeding leeches, lowered capture rates by Nephelopsis.
5. In 2 years sampling, only one of seven lakes, however, had appreciable abundances of Nephelopsis. It is therefore suggested that the rarity, low feeding rates, and lack of strong gastropod preference limit the effect of Nephelopsis on field gastropod distributions.     

Larval development and metamorphosis in the marine pulmonate Amphibola crenata (Mollusca: Pulmonata)

The development of the free-swimming veliger of Amphibola is followed from hatching to settlement, and the larval structures compared with those of post-metamorphic juveniles and adult snails. Observations of living specimens and light-microscope sections were combined with scanning electron microscopy to build up a composite picture of veliger structure.
Four stages in the development of veligers are recognized, each being characterized by the appearance of organ systems such as the mantle cavity, larval heart, adult heart and kidney, and larval pallial gland. At or after metamorphosis, the larval systems (heart, kidney and pallial gland) disappear, and the developing adult organs move to the positions characteristic of adult snails.
Organogenesis in Amphibola veligers is compared with that of prosobranch and opisthobranch larvae, and with that of pulmonate larvae with direct development. The closest similarity is seen to be with opisthobranch veligers.     

Snail predation by larvae of Sepedon scapularis Adams (Diptera: Sciomyzidae), a potential biocontrol agent of snail intermediate hosts of schistosomiasis in South Africa

Malacophagous larvae of the fly Sepedon scapularis Adams were shown experimentally to be effective predators of three species of aquatic pulmonate snails tested as prey: Bulinus africanus (Krauss) an important intermediate host of Schistosoma haematobium (Bilharz), Bulinus tropicus (Krauss) and the invasive species Physa acuta Draparnaud. Survival of S. scapularis larvae from instar to instar was negatively affected by the size of prey snails, since larvae tended to be asphyxiated by the mucous secretions of the snails, or by the larval hydrofuge hairs becoming entangled in snail faeces. In experiments to test the choice of S.scapularis larvae for different species and sizes of snails, B.africanus was significantly disfavoured compared with the other two snail species. Small snails (< 3 mm) were more frequently killed by all three larval instars of S.scapularis. First instars killed few, if any, large snails (> 7 mm), whereas second and third instars preyed effectively on all sizes of snails. Third instars killed significantly more snails than younger instars. For larvae offered only one species of snail as prey, the mean total number of snails killed per larva during its entire development was 49 B.africanus, 45 B.tropicus or 34 P.acuta. It is concluded that S.scapularis is potentially useful as a biological control agent for use against indigenous Bulinus and exotic Physa snails.     

Influence of lake trophic status on the occurrence of microcystin-LR in the tissue of pulmonate snails

1. To assess the influence of lake trophic status on the occurrence of microcystin-LR (MCLR) in the tissue of resident pulmonate snails, we sampled seven lakes of varying primary productivity in Alberta, Canada. Parameters associated with productivity were measured every 2 weeks from mid-May through mid-September 1995. Phytoplankton and resident pulmonate snails were collected and analysed for MCLR concentration via high-performance liquid chromatography.
2. For all species of gastropod (i.e. Lymnaea stagnalis , Helisoma trivolvis and Physa gyrina ), the concentration of MCLR in the tissue was correlated ( P  ≤ 0.03) with toxin in the phytoplankton, but not with extracellular aqueous microcystin ( P > 0.28). The concentrations of toxin in the tissues of L. stagnalis and P. gyrina were also correlated with the relative abundance of Microcystis spp. ( P < 0.01).
3. Given that Microcystis spp. abundance and MCLR concentration within phytoplankton are correlated with indicators of productivity, we conclude that trophic status is important in influencing the occurrence and concentration of MCLR in pulmonate snails.     

Why are there few algae on snail shells? The effects of grazing,nutrients and shell chemistry on the algae on shells of Helisoma trivolvis

1. Freshwater snails often lack visible growths of algae on their shells. We tested three possible mechanisms that may account for this (grazing, snail-derived nutrients and chemical defences), using the ramshorn snail Helisoma trivolvis .
2. The experiments were carried out in floating plastic enclosures in a pond and comprised seven treatments. Grazing treatments were: a lone snail (ungrazed, as self-grazing does not occur), Helisoma with conspecifics, Helisoma with the co-occurring pond snail Physa sp., empty shells with Helisoma , and ungrazed empty shells. Nutrient effects were possible in all treatments with occupied shells (lone snail; Helisoma with conspecifics, and with Physa ) versus absent in other treatments. Testing for chemical defences compared algae on fresh empty shells, weathered shells (outer organic periostracum layer absent) and boiled fresh shells (with denaturization of susceptible proteins).
3. Diatoms dominated algal assemblages on snail shells. Although the upright diatom Gomphonema gracile was abundant on all shells, it was dominant on the shells of snails housed with other snail grazers (either Helisoma or Physa ).
4. Only the lone snail (nutrients but no grazing) showed higher algal biomass, so presumably any nutrient effect in the treatments with grazers was masked. Both Helisoma and Physa were observed apparently grazing on Helisoma shells, and consequently algal biomass in multi-snail treatments was similar to that on empty shells. Scanning electron microscopy revealed that algal density was highest near the aperture of live snails, but not empty shells; this is consistent with a nutrient addition effect. There was no evidence of chemical defences against algal growth.
5. In soft-bottomed freshwater habitats with abundant snails, shells of living snails provide nutrient-augmented substrata that may indirectly boost overall snail production.     

Experimental predation studies of malacophagous larvae of Sepedon fuscipennis (Diptera: Sciomyzidae) and aquatic snails

Experimental studies were conducted under both laboratory and field conditions to determine the effects of prey density, three levels of prey aggregation, water depth, and predator density on the number of snails killed per larva of Sepedon fuscipennis. Of these factors, predation rates were most influenced by prey density and water depth. The number of small (2–4.5 mm) Lymnaea palustris killed per larva of S. fuscipennis increased at a decreasing rate as prey density increased under shallow water conditions. Larvae killed a mean of 14 snails at a prey density of 200/m², while an average of 24 snails were killed per larva of S. fuscipennis at a prey density of 4000/m². This functional response to prey density was largely confined to third-instar larvae, and as water depth was increased the response was not apparent.A field study in which larval densities of S. fuscipennis were manipulated showed that the population density of smaller individuals of L. palustris (< 4.5 mm) was reduced when predator density was increased. Populations of Physa integra, Gyraulus parvus, and larger L. palustris were not significantly reduced by the malacophagous larvae at the levels tested.     

The effect of digenea larvae on calcium content in the shells of Lymnaea stagnalis (L.) individuals

The objective of this work was to examine the calcium concentration in the shells of naturally infected individuals of Lymnaea stagnalis. Calcium concentration in the water of lakes and in snail shells was examined by the EDTA method. Calcium concentration in the shells of infected snails was found to be higher than that in the shells of uninfected ones only in the lake with the lowest calcium concentration in water. However, this difference is not evidence of hypercalcification of snail shells caused by trematode larvae but suggests that the limited calcium concentration in the environment can be a very good factor for studying snail shell calcification under natural conditions. Moreover, small adult snails from 2 lakes had more calcium in their shells than did large ones. Different reproductive rates of different-sized snails could be the cause of this phenomenon.     

Effects of diet and<Emphasis Type="Italic"> Echinostoma revolutum</Emphasis> infection on energy allocation patterns in juvenile<Emphasis Type="Italic"> Lymnaea elodes</Emphasis> snails

Resource allocation strategies may be influenced by both biotic and abiotic factors. The purpose of this study was to investigate the effects of both parasitism and diet quality on the growth, reproduction, and survival of the pond snail, Lymnaea elodes. In addition, we assessed parasite growth and reproduction. High-protein (high diet) or low-protein diets (low diet) were fed to juvenile L. elodes snails that were either exposed or sham-exposed to the castrating trematode, Echinostoma revolutum. Host growth was assessed weekly; reproduction and survival were recorded every 2-3 days. We estimated parasite development as the time to parasite release from the host (patency), and parasite reproduction as the number of larvae shed from infected snails at two time points. Diet and infection status had significant effects on snail growth. Infected snails produced few eggs and tended to grow to larger sizes than uninfected snails regardless of diet. In contrast, exposed-uninfected individuals displayed diet-dependent patterns of growth and reproduction. On the high-protein diet, uninfected and exposed-uninfected snails exhibited similar patterns of growth and reproduction, whereas in the low-diet treatment, exposed-uninfected snails exhibited reduced growth and delayed reproduction relative to uninfected individuals. Survival differed among treatments in the latter stages of the study with infected snails exhibiting reduced survival relative to snails from other treatments. Moreover, infected low-diet snails exhibited lower survival than infected high-diet snails. Parasite development and reproduction did not appear to be directly influenced by the quality of host diet. Results from this study suggest that energy allocation patterns are context-dependent in juvenile snails, influenced by parasite exposure and diet quality. Furthermore, parasite reproduction appears to depend more on host size than on the quality of host diet.     

Infectivity of Echinostoma friedi miracidia to different snail species under experimental conditions

The infectivity of Echinostoma friedi (Trematoda: Echinostomatidae) miracidia was studied experimentally in a range of laboratory-reared snails that coexist in the same natural locality, namely Radix peregra, Lymnaea fuscus, L. truncatula (Lymnaeidae), Gyraulus chinensis, Helisoma duryi (Planorbidae) and Physella acuta (Physidae), and snails from different geographical origins acting naturally or experimentally as intermediate hosts of Schistosoma spp., namely Planorbarius metidjensis (from Málaga, Spain), Biomphalaria glabrata (Guadeloupe), B. alexandrina (Egypt) (Planorbidae), Bulinus cernicus (Mauritius), B. globosus (Zambia), B. natalensis (South Africa) and B. truncatus (Niger) (Bulinidae). Six species of snails were found to be susceptible, with the rate of infection ranging from 0 to 36.7%. The highest infection was detected in R. peregra. The low host specificity of E. friedi might have an epidemiological significance as a requisite for a recent establishment in a new geographical area.     

Incompatibility between miracidia of Schistosoma mansoni and Helisoma duryi occurs at two stages: penetration and intramolluscan establishment

Helisoma spp. snails are not susceptible to infection with miracidia of Schistosoma mansoni because the miracidia do not penetrate them. However, in view of the phylogenetic proximity and histocompatibility between Helisoma spp. and the normal intermediate host, Biomphalaria glabrata , schistosome miracidia conceivably could survive if experimentally introduced into the hemocoel of Helisoma spp. To test this hypothesis, schistosome-susceptible NIH albino B. glabrata, schistosome-resistant Salvador B. glabrata, and Helisoma duryi were injected with miracidia of S. mansoni, and the outcome was followed both by monitoring snails for infection for several weeks and by histological examination at 24 and 48 hr post-injection (PI). Patent infections developed in most NIH albino snails but in none of the Salvador B. glabrata or H. duryi individuals. Histological analysis showed a higher proportion of normal sporocysts in various tissues of NIH albino snails at both time periods relative to Salvador snails, which contained mostly sporocysts undergoing hemocytic encapsulation. In H. duryi , nearly all sporocysts were dead by 48 hr PI.