FIELD OBSERVATIONS ON FEEDING OF THE LAND SNAIL HELIX ASPERSA MULLER

Feeding of the land snail Helix aspersa (Müller) was observedat monthly intervals. Three natural populations in Galicia (NW-Spain)were studied. At two sites only a few plants constituted thebulk of the diet and in spring the snails' diet had the highestdiversity (H'). In the third population feeding and distributionof Helix aspersa (Müller) were observed in a small plotwith permanent patches of Urtica dioica. Nearly one half offeeding snails fed upon Urtica dioica. Most of the other observations wereon Mentha suaveolens, Ranunculus repens and Gramineae. The diversity ofthe snails' diet showed seasonal variation with the maximumin the autumn months. Comparison between the availability ofthe different plant species and their contribution to the snails'diet showed that the snails did not eat at random; Urtica dioicawas eaten much more than expected from its occurrence and grasseswere strongly under-represented in the snails' diet. Temporalchanges of availability were significantly correlated with the amountseaten in the case of Urtica, but not for the other food plants.The distribution of the snails in the plot was significantlycorrelated with that of Urtica. Chemical analyses of the foodplants revealed Urtica as the species with the higher protein,ash and calcium contents. The strong preference of Helix aspersafor Urtica dioica could be explained by the value of Urticaas food or by its suitability as habitat for the snails. The largestproportions of green material in the snails' diet occurred inthe spring and juveniles ate more green material than adultsin the three populations. (Received 16 March 1998; accepted 30 November 1998)     

VARIABILITY OF EGG CANNIBALISM IN THE LAND SNAIL HELIX ASPERSA IN RELATION TO THE NUMBER OF EGGS AVAILABLE AND THE PRESENCE OF SOIL

Egg cannibalism by hatchlings has been demonstrated in somepulmonate land snails; this behaviour is promoted by a highhatching asynchrony within the egg-batch. Under laboratory conditions,the percentage of new-born snails Helix aspersa having cannibalisedunhatched eggs was not influenced by the soil factor: about70% of them ingested one egg within their first four days oflife whether soil was present or not. The propensity to eggcannibalism in hatchlings of H. aspersa increased with egg density.However, most of the new-born hatchlings consumed a single eggduring the four days following hatching, and only exceptionallytwo. The consumption of one egg increased the snails wet weightby 38.7% within four days. A weak ingestion of soil componentsalso occurred, but it induced a growth that was three-timesless than that due to the consumption of an egg. In addition,the survival of newly hatched snails maintained under non-dehydratingthermohygrometric conditions was high, even when they were submittedto four days food-deprivation. (Received 22 July 1999; accepted 24 November 1999)     

THE INFLUENCE OF INGESTIVE CONDITIONING ON FOOD CHOICES IN THE LAND SNAIL HELIX ASPERSAMULLER (GASTROPODA: PULMONATA: STYLOMMATOPHORA)

An experiment of food choice in one-day-old naïve landsnail Helix aspersa Müller was carried out, involving threephases. First, a training period on a monophagous diet of leafdiscs of Taraxacum officinale or Urtica dioica was given for1, 5, 10, 15 or 30 days. The area of leaf discs consumed byeach snail during this stage was assessed. In the intermediatephase, the animals were not fed for 24hours. Finally, food choicewas tested over 24 hours, when animals could eat leaf discsof each species of plant usedfor training. A coefficient offood preference for Urtica dioica was calculated (UPC = areaof Vrtica leaf disc consumed/area of Urtica and Taraxacum discsof leaves consumed for 24 hours). The length of the training phase did not reduce the percentageof individuals preferring the plant previously consumed. Onthe contrary, after 15 days of monophagous diet, animals formerlyfed on Urtica showed a coefficient of food preference for Urticasignificantly higher than Taraxacum fed animals, and after 30days of training this phenomenon was more pronounced. Thus,during the choice test, snails ate more of the previously eatenplant leaf. Our experiment highlighted the feeding behaviourplasticity of H. aspersa. The significant correlation betweenthe area of Urtica eaten during the training phase and the coefficientof food preference for Urtica is discussed. (Received 10 April 1994; accepted 10 January 1995)     

INFLUENCE OF THE QUINOLIZIDINE ALKALOID CONTENT OF LUPINUS ALBUS (FABACEAE) ON THE FEEDING CHOICE OF HELIX ASPERSA (GASTROPODA: PULMONATA)

Adult Helix aspersa snails were maintained individually forone week in plastic cages with 9 living Lupinus albus plantsas their only food. Among these 9 plants, 3 chemotypes bitter,intermediate and sweet which differed in their alkaloid contentwere equally represented. Each day, the leaf surface grazed and the number of leaves attackedby the snails were recorded for each chemotype and each snail.A consumption/ attack (C/A) ratio was calculated by dividing thesurface grazed (C) by the number of attacks (A). The numberof attacks and the grazed area were positively correlated foreach chemotype during the whole experiment, and the snails atesimilar quantities of lupin each day. After 4 and 6 days ofexperiment, we noticed a rejection of the bitter chemotype infavor of the intermediate and sweet ones respectively. Afterthe 6th day, the surface grazed per attack was significantlyhigher on the sweet chemotype than on the bitter plants. Wehypothesize that rejection of the bitter chemotype might berelated to (i) an alkaloids reaction threshold associated withan increase in the amount of alkaloids in the wounded plantsand/or (ii) aversive ingestive conditioning. (Received ; accepted 16 April 1999)     

FORAGING BEHAVIOUR OF TERRESTRIAL GASTROPODS: INTEGRATING FIELD AND LABORATORY STUDIES

The foraging range of adult snails, Helix aspersa, has beenstudied using radio-transmitters. Snails did not return to anexact roost site after foraging, but often remained in the foodpatch, or returned only to the general roost area. Time lapse video films of the slug Deroceras reticulatum madeunder infra-red lighting in arenas were analysed for tracklengthsand degree of turning, in order to simulate slug movements inan unbounded situation. The results suggest that many food itemsare found by random encounter. Slugs usually ate the first food item found, but often ignoredfood items encountered later. If food was scarce, the slugsfed almost every time. Electronic recordings of bites on a wheatflour pellet over 24 hours show that feeding is most intensein the first two and a half hours from starting to feed, andlater meals are both shorter and less regular. Starved slugsdiffered from fed slugs principally by taking a second mealshortly after the first. When given a choice of a more preferredfood (maize pellets) and a less preferred food (pea pellets)in different ratios, the slugs appeared to encounter pelletsat random, but they fed more from the preferred pellets unlessthe ratio was 1 maize: 7 pea. Starved slugs ate twice as muchas fed ones.     

ULTRASTRUCTURAL LOCALIZATION OF COMPLEX CARBOHYDRATES AND PHOSPHATASES IN DIGESTIVE GLAND CELLS OF FEEDING AND HIBERNATING SNAILS, HELIX LUCORUM (GASTROPODA: HELICIDAE)

The digestive gland of normally-fed snails Helix lucorum, aswell as that of snails which had hibernated for 4 months wereexamined by the use of cytochemical techniques for detectionof acid and alkaline phos-phatase, as well as of periodate-reactive(PA-TCH-SP technique), sulfated (HID-TCH-SP technique) and carboxylatedcarbohydrates (LID-TCH-SP technique). The cytochemical resultssupport the hypothesis of intracellular digestion via lysosomalactivity of material taken up by endocytotic processes by thecells of the digestive gland. Four months hibernation did notaffect the intracellular distribution of polysac-charides andphosphatases in the cells of the digestive gland of H. lucorumcompared to that in the control snails. In addition, hibernationaffected the percentage of the calcium cells which significantlyincreased compared to the non-hibernating snails, whiie thepercentages of the digestive and excretory cells remained almoststable. However, the periodate-reactive sulfated and carboxylatedpolysaccharides of the digestive gland cells decreased in thehibernated snails compared to the controls. The results suggestthat the cytochemistry of periodate-reactive, sulfated and carboxylatedpolysaccharides used in the present study could, also, be appliedto the study of lysosomal activities. (Received 1 October 1991; accepted 4 December 1991)     

HETEROGENIC COPULATORY BEHAVIOUR PRODUCES NON-RANDOM MATING IN LABORATORY TRIALS IN THE LAND SNAIL HELIX ASPERSA MULLER

This study reports the results of mate-choice experiments betweenresident and non-resident individuals of the simultaneous hermaphonteland snail Helix aspersa. Snails from different sites differedin their mating proclivity, which resulted in non-random matingin laboratory mate-choice tests. Those snails with the highermating propensity therefore tended to mate with their own typeand tended to be chosen as partner because they were also activelyseeking mates. Keeping snails under crowded conditions did noteffect a lasting influence on mating behaviour in laboratorytests, despite the decline in copulatory behaviour under crowdedfield conditions. (Received 7 November 1994; accepted 31 August 1995)     

EFFECTS OF WOUND HEALING ON ZINC DISTRIBUTION AND ALKALINE PHOSPHATASE ACTIVITY OF HELIX ASPERSA (GASTROPODA: PULMONATA)

Helix aspersa were loaded with ⁶⁵zinc and after 2 and 5 daysstarvation there was no significant reduction in the zinc concentrationof the digestive gland and shell. Wounding of the optic tentacle,foot tissue and partial shell removal of H. aspersa resultedin deposition of ⁶⁵zinc in the wound area. Alkaline phosphataseshowed no increase in concentration at the sites of wounding. (Received 15 October 1985;     

WATER EXCHANGE, TEMPERATURE TOLERANCE, OXYGEN CONSUMPTION AND ACTIVITY OF THE NAMIB DESERT SNAIL, TRIGONEPHRUS SP.

Water exchange, temperature tolerance and oxygen consumptionof the snail, Trigonephrus sp., from the southern Namib desertof Namibia were examined and related to activity. At 25°Cand 15% R.H. mean water loss and food and water uptake were5.95 mg. day^–1 and 630 mg.day^–1, respectively. Bodytemperature tracked sand temperature. Snails tolerated sandtemperatures as high as 45°C. Mean ± S.D. oxygenconsumption rates were 32.0 ± 2.94 µlO₂.g totalbody mass^–1.h^–1 at 15°C, when the snails wereactive, and 11.27 µlO₂.g total body mass^–1.h^–1at 25°C, when the snails were inactive. These values are2-6 times lower than those recorded for the similarly sizedmesic snail, Helix aspersa. Activity experiments indicated thatlow ambient temperatures and high humidities were favoured bythe snails. This, together with the burying behaviour of thesesnails during high temperatures, suggests that they limit stressby restricting activity to physiologically-favourable periods,even though more-extreme conditions may be tolerated. (Received 7 June 1990; accepted 20 November 1990)     

GROWTH, MORTALITY AND FECUNDITY IN SUCCESSIVE GENERATIONS OF HEIX ASERSA MULLER CULTURED INDOORS AND CROWDING EFFECTS ON FAST-, MEDIUM- AND SLOW-GROWING SNAILS OF THE SAME CLUTCH

This paper examines the optimum conditions for edible snailsHelix aspersa to be cultured indoors successfully in successivegenerations (originating from the crossing of snails comingfrom different clutches of a previous generation), and the effectof crowding on growth and reproduction in fast-, medium-, andslow-growing snails coming from the same clutches. The timeneeded for the snails to reach marketable size (25–32mm)varied from 2.5 to 5 months up to the 7th generation. The timeneeded for the snails to mature and reproduce from 4 to 7 monthsuntil the fifth generation. After the F5 x F5 generation, thefinal size of the snails decreased. The number of eggs did notdiffer statistically among the different generations but thereproductive success (how many snails reproduced/cage) increasedfrom Fl = Fl generation onwards to F5 x F5. In F6 x F6 onlythree (out of 26) snails reproduced and in F7 x F7 none, althoughthe snails remained under controlled conditions for 15 moremonths. Mortality in the different generations varied from 0–10%up to F5 x F5 but from F6 x F6 onwards increased and reached25%. Concerning the origin of snails, it was found that largersnails (originating from Kyparissia, Peloponnesos) lay statisticallymore eggs (138.40 ± 29.60, N =5) than smaller ones (77.38± 40.42, N=4) (originating from Hania, island of Crete).Hatching success was greater, too. (Received 10 September 1996; accepted 24 March 1997)     

VARIATION OF WATER CONTENT IN HELIX ASPERSA MULLER IN A NATURAL ENVIRONMENT

1. Daily and seasonal variations occur in the water content ofHelix aspersa Müller.
2. The water content is not constantthroughout 24 hours but fluctuatesby 2 to 10%. There is noapparent daily rhythm.
3. Humidity has an effect on the watercontent. Falls in relativehumidity to 60% or less, triggera decrease of the snails' hydration,over a range of temperatures,as well as a brief pause in activity.
4. Only external climaticconditions produce noticeable fluctuationsof water content,and they more particularly affect juvenileswhich undergo greaterevaporo-transpiration.
5. A seasonal rhythm in water contentexists in Helix aspersa,with maxima during spring and autumnand minima in June andJanuary.
6. The water content of juvenilesis always superior to that ofthe adults.
7. The water contentaffects the duration of the snail's activephase.

(Received 22 September 1988; accepted 5 February 1989)     

BIOCHEMICAL COMPOSITION OF HELIX SNAILS: INFLUENCE OF GENETIC AND PHYSIOLOGICAL FACTORS

This paper describes the biochemical composition of differentspecies (Helix lucorum, Helix pomatia) and sub-species of snails(Helix aspersa aspersa, Helix aspersa maxima) reared in thesame conditions with a feed (‘Helixal’) speciallydesigned for edible snails. In addition, the composition ofwild H. pomatia and H. lucorum is presented to allow comparisonbetween snails of different origins. Analyses determined thepercentages of proteins, lipids and minerals. They reveal bothsimilarities and differences in composition according to thespecies and the part analysed (whole body, pedal mass, and visceralmass). H. pomatia contains the highest percentage of mineralmatter and the lowest percentage of lipids. Surprisingly, proteincontents are slightly different between artificially rearedH. aspersa maxima of 3 months old and wild H. pomatia. The resultsmake it possible to evaluate nutritional quality of snails withthe composition of the body of four edible snail species. (Received 16 September 1996; accepted 24 May 1997)     

Tales of two snails: sexual selection and sexual conflict in Lymnaea stagnalis and Helix aspersa

Sexual selection and sexual conflict have been shown to playkey roles in the evolution of species with separate sexes. Experimentalevidence is accumulating that this is also true for simultaneoushermaphrodites. For example, many species of land snails forcefullystab their mating partners with love darts. In the brown gardensnail (Helix aspersa, now called Cantareus asperses), this dartincreases sperm storage and paternity, probably via the transferof an allohormone that inhibits sperm digestion. A recent interspeciescomparison of dart-possessing land snails revealed coevolutionbetween darts and spermatophore-receiving organs that is consistentwith counteradaptation against an allohormonal manipulation.The great pond snail (Lymnaea stagnalis) seems to use a seminalproduct to manipulate its partner and mates in the male rolewhen enough seminal fluid is available in the prostate gland.Receipt of semen not only initiates egg laying in virgin animals,but also feminizes the mating partner later in life. These increasesin the female function have been shown to go at the expenseof growth and seminal fluid production of the sperm recipient.Although in Helix, and probably also Lymnaea, the sperm donorbenefits from the induced changes through increased fertilizationsuccess, the sperm recipient may experience injury, imposedreallocation of resources, and altered sperm storage. Thesefindings support the existence of sexual conflict in simultaneouslyhermaphroditic snails, and its importance for the evolutionof mating behaviors and reproductive morphologies is discussed.     

INTERACTIONS BETWEEN SOIL BACTERIA AND THE MOLLUSCAN ALIMENTARY TRACT

The alimentary tract is a major site of interaction betweenan animal's ecosystem and its physiology. It may be intermittentlyor permanently populated by microorganisms from its environmentthat could be beneficial or pathogenic in their influence. Thebacterial populations of the snail (H. aspersa) and the woodlouse(O. asellus) have therefore been estimated in terms of colonyforming units. gm body weight^–1. The bacteria in the snailappear to be similar to those found in soil and show wide resistanceto antibiotics. The main types are gram negative rods tentativelyidentified as belonging to Pseudomonas, Xanthomonas, Acinobacter,Vibrio and Enterobacteriaceae. Some gram positive and endosporeforming bacteria of Bacillus, Staphylococcus and Micrococcusspp were also present. The numbers of bacteria increase towardsthe posterior of the alimentary tract although the numbers inthe digestive gland are more constant. An artificial diet is described that can be used to feed snailsand study the effects of known amounts of additives. When starvedor kept under clean conditions the bacterial population of thesnail gut falls and the thickness of the alimentary tract decreases.All the indications are that the bacteria in the alimentarytract of the snail are indiscriminately accumulated from thenatural environment and are transient populations. (Received 7 June 1989; accepted 24 July 1989)     

GROWTH, MORTALITY, AND FEEDING RATES OF THE SNAIL HELIX ASPERSA AT DIFFERENT POPULATION DENSITIES IN THE LABORATORY, AND THE DEPRESSION OF ACTIVITY OF HELICID SNAILS BY OTHER INDIVIDUALS, OR THEIR MUCUS

When reared at high densities, young Helix aspersa show lessshell growth, even if waste products are removed. They alsofeed less, and show increased mortality. It is suggested thatthese effects are linked to reduced activity. Juveniles showreduced activity in the presence of adults or their mucus. Mucusof adult Cepaca nemoralis also depresses the activity of bothadult and young Helix and Cepaea. ^* Present Address: Unit Zoologi, Universiti Kebangsaan Malaysia,Jalan Pantai Baru, Kuala Lumpur, West Malaysia. (Received 12 May 1981;     

Rhabditis maupasi: occurrence in food snails and cultivation

Rhabditis maupasi, a nematode, inhabits the mantle cavity of terrestrial snails, Helix aspersa, that are imported from Morocco and sold in the food markets of New York City. Most of the nematodes found in living snails are stage-3 larvae which mature and reproduce after the snail has died and decayed. The nematodes were isolated from snails and cultivated with a microflora and then in species isolation, i.e., axenically. Sterile slices of rabbit kidney supported by agar slants served as the initial culture system. Subsequently, a liquid medium was used that contained raw rabbit liver extract. When, instead, raw snail extract was substituted in the liquid medium, nematode larvae did not mature unless the snail extract was digested secondarily or inactivated by heating. Nematodes grow abundantly in dead snails and in treated snail extract but not in living snails or in raw snail extract. The reason is probably not the nematodes' lack of suitable digestive enzymes for snail tissue nor the absence of adequate nutrients in living, microbially poor snails, but rather the presence of an inhibitor in snails.     

SNAIL FARMING: A SURVEY OF BREEDING MANAGEMENT, HYGIENE AND PARASITISM OF THE GARDEN SNAIL, HELIX ASPERSA MULLER

Fifteen farms in 1984 and twenty in 1985 were investigated forbreeding management, hygiene and parasitism. Farm hygiene didnot seem to play an important role on the breeding performancesin our samples. Nematodes were the most frequent parasites:Alloionema appendiculatum were equally prevalent among juvenileand adult snails whereas Angiostoma aspersa and Nemhelix bakeriwere found mostly in reproductive adults. The presence of nematodes,assessed by coproculture, was negatively related to breedingperformances     

CYCLIC ACTIVITY AND EPITHELIAL RENEWAL IN THE DIGESTIVE GLAND TUBULES OF THE MARINE PROSOBRANCH MAORICRYPTA MONOXYLA (LESSON)

Three cell types are present in tubules of the digestive glandof the marine prosobranch Maoricrypta monoxyla (Lesson). Histochemistry,and feeding and starvation experiments established that themain type, the digestive cell, is involved in endocytotic uptakeof food material from the lumen. Digestion of this materialoccurs within vacuoles, and indigestible material (indicatedby the dye trypan blue) accumulates in basal residual bodiescontaining lipofuscin pigment. Another cell type, the cryptcell, appears to secrete a glycoprotein, probably enzymaticin function. The third cell type contains large vacuoles butit has not been Established whether the contents are secretoryor excretory. The tubules undergo a cycle of digestive activity not relatedto the tidal cycle as in some marine molluscs, but probablyan indirect result of the feeding regime. The cycle begins withimmature tubules in which some endocytosis occurs. These developinto absorbing tubules involved mainly in food uptake. In maturetubules intracellular digestion occurs. At a later stage thetubules fragment to produce spherules which may conserve usefulmaterial, and finally they, disintegrate completely. Eighty per cent of the dividing cells in the epithelium occurin crypts, which are therefore considered to be the main sitesof epithelial renewal. The processes by which tubules may bereformed after breakdown are discussed (Received 28 September 1978;     

NATURAL HISTORY AND POPULATION DYNAMICS OF THE LAND SNAIL HELIX TEXTA IN ISRAEL (PULMONATA: HELICIDAE)

Helix texta is endemic to the Mediterranean regions of Israel.It has a seasonal activity pattern which starts in the autumn,with the first rains, and dwindles towards the spring, whenthe snails dig into the ground for a six months long aestivation.A cold spell of 0°C will, however, terminate the activeseason of the adult snail, even in the middle of the rainy season.Survival of the young is very low and most of them (90%) donot survive their first year, because of the winter cold andthe summer drought. Massive predation of adult snails by wild boar was observedin December 1986: within a few days, about 50% of the adultsin the study in the area were eaten. A very rapid growth ofyoung and subadults was observed immediately after this predation.These observations suggest that the extent of recruitment ofnew adults to the population may be partly controlled by existingadults, through a growth-inhibiting pheromone in the mucus.After predation this inhibiting factor disappears, enablinga subsequent rapid growth of the young. In this manner, massive,irregular predation of the adults by a large predator, and changesin juvenile survival, result in sharp fluctuations in the agestructure of the population. The resulting pattern of unstablepopulation dynamics is different from that described for theEuropean species of Helix. (Received 16 January 1989; accepted 17 April 1989)     

Dart shooting influences paternal reproductive success in the snail Helix aspersa (Pulmonata, Stylommatophora)

Although animal courtship behaviors are generally understoodwithin the context of sexual selection, the relevance of manysexual behaviors to sexual selection, and vice versa, remainsunexplained. For example, the adaptive function of the "lovedart" used in the precopulatory behavior of hermaphroditicland snails is only now becoming apparent. Contrary to previous assumptions, dart shooting is unlikely to function as a stimulusfor copulation. In searching for a more ultimate explanationof the dart's function, we tested whether variation in dartshooting influences reproductive fitness in Helix aspersa.Individual mother snails were mated sequentially to two potentialfathers. Dart shooting was closely observed and quantifiedfor all pairings, and percentages of offspring sired by each potential father were determined using allozymes. The resultsindicate that snails that shoot darts effectively have significantlygreater paternal reproductive success than snails that shootpoorly. In contrast, there was no significant effect of matingorder on either dart shooting or paternal reproductive success.