THE UPTAKE OF BIOGENIC MONOAMINES BY THE ISOLATED AURICLE OF THE SNAIL (HELIX POMATIA)

—The uptake of [³H]5HT, [³H]dopamine, [³H]noradrenaline and [³H]octopamine into the auricle of Helix pomatia was studied. When tissues were incubated at 25°C in media containing radioactive amines, tissue:medium ratios of about 49:1, 14:1 and 5:1 for 5-HT, dopamine, noradrenaline, and octopamine respectively were obtained after a 20–30 min incubation time. Tissues incubated at 25°C in media containing radioactive amines for 20–30 mins showed that almost all (96%) the radioactivity was present as unchanged [³H]5-HT, [³H]dopamine, [³H]octopamine or [³H]noradrenaline. The high tissue:medium ratios for 5-HT and dopamine, but not for noradrenaline and octopamine, showed saturation kinetics which were dependent upon temperature and sodium ions. From the Lineweaver–Burk plots, two uptake mechanisms for 5-HT at 25°C were resolved; the high affinity uptake process having a K_m1 value of 6.0 ± 10^?8m and a V_m1 value of 0.115 nmol/g/min while the lower affinity process had a K_m2 value of 1.04 ± 10^?6m and a V_m2 value of 0.66nmol/g/min. At 0°C a single uptake mechanism for 5-HT occurred which gave a K_m value of 5.02 ± 10^?8m and a V_m value of 0.0165 nmol/g/min. In the case of dopamine, the Lineweaver–Burk plot at 25°C showed a single uptake process with values for K_m and V_m of 1.55 ± 10^?7m and 0.086 nmol/g/min respectively. This process did not function at 0°C. The effect of various agents and ions upon the accumulation processes for all amines was also studied, and the data indicate that the same neurons probably accumulate more than one amine type. It is concluded that 5-HT and dopamine uptake in the auricle is a mechanism for inactivating these substances at 25°C and that an uptake mechanism for 5-HT also functions at 0°C. The results are discussed from the point of view of 5-HT's being the cardioexcitatory substance in the snail heart.     

THE LOVE DART OF HELIX ASPERSA MULLER IS NOT A GIFT OF CALCIUM

The phenomenon of dart shooting in several species of land snailshas still not been explained. We were interested in whetherthe dart can function as a nuptial gift of calcium, as previouslyproposed. Donating calcium would increase the fitness of theoffspring and thereby result in a higher reproductive successfor the donor. We confirmed in Helix aspersa that the developingembryo takes up calcium from the egg shell for the formationof its embryonic shell. However, other results from behaviouralobservations and calcium measurements in various reproductivestructures do not support the calcium hypothesis. We found thatthe dart penetrated the skin in 91.7% or the shootings, butit was internalized by the recipient in only 6.3% of the shootings.The amount of calcium in one dart is roughly equal to that ofone egg, and thus it would not contribute significantly to anaverage clutch of 59 eggs. The spermatophore contains virtuallyno calcium, and therefore it is unlikely that the dart signalsa donation of calcium with the sperm. The dart is also unlikelyto influence egg laying since dart shooting does not predicteither the latency or the productivity of egg laying in theshooter or the recipient. We conclude that the love dart ofHelix aspersa is not a gift of calcium. Instead, we suggestthat it is a vehicle to introduce a substance into the partnerto influence the fate of the donated sperm. (Received 8 November 1996; accepted 25 April 1997)     

AN ULTRASTRUCTURAL STUDY OF SPERMATOZOA OF HELIX ASPERSA AND HELIX POMATIA (GASTROPODA, PULMONATA)

Spermatozoa of the pulmonates Helix aspersa Müller andH. pomatia Linnaeus are examined in detail using transmissionelectron microscopy (TEM). Important features such as the acrosome,perinuclear sheath, nucleus and terminal region of the midpieceare described for the first time. Also presented are the firstultrastructural observations on spermatozoa from spermatophoresin any pulmonate gastropod (H. aspersa). No morphological differencescould be found between sperm taken from spermatophores and thosewithin the hermaphrodite duct in H. aspersa. Spermatozoa ofH. aspersa and H. pomatia snow all the characteristics of euthyneuranspermatozoa, namely: a helically-keeled nucleus; distinctivearrangement of acrosomal components (apical vesicle, acrosomalpedestal), and extremely elongate midpiece (axoneme and glycogenhelix enclosed by matrix and paracrystalline layers). The spermnucleus of both species is short, and the midpiece also formsthe terminal portion of the spermatozoon (glycogen piece absent).The extraordinary positioning of the acrosome in H. aspersa—reflectedbackwards from the nuclear apex—is not observed in H.pomatia, though a perinuclear sheath (possibly another acrosomalcomponent) is present in sperm of both species. Helix spermatozoaare compared with other euthyneuran sperm and briefly discussedfrom the systematic viewpoint. Present address: Department of Zoology, St. Lucia, 4067, Brisbane,OLD, Australia (Received 23 May 1988; accepted 17 August 1988)     

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)     

GENETIC STUDIES OF ALLOZYME VARIATION IN LEUCINE AMINOPEPTIDASE IN THE LAND SNAIL HELIX ASPERSA (MULLER)

In the land snail H. aspersathe enzyme LAP has two loci, LAP-1and LAP-2, both of which arc monomeric enzymes under the controlof multiple alleles, the alleles being codominant. None of theobserved ratios in the pheno types in the experimental progenywere significantly different from Mendelian expectation. ^* Present Address: Bournside School, Cheltenham, Glos. (Received 1 September 1981;     

IN VITRO EXPERIMENTS ON THE METABOLISM, ACCUMULATION AND RELEASE OF 5-HT IN THE NERVOUS SYSTEM OF THE SNAIL HELIX POMATIA

Abstract— The accumulation, metabolism and stimulated-induced release of 5-HT in the nervous system of the snail was studied. When nervous tissue was incubated at 24°C in a medium containing [¹⁴C]5-HT or [³H]tryptophan, tissue: medium ratios of about 25:1 and 4:1 respectively were obtained after 45 min incubation. The process responsible for [¹⁴C]5-HT accumulation showed properties of an active transport system: it was temperature sensitive and was greatly inhibited by dinitrophenol and ouabain. Furthermore, the accumulation process was inhibited by imipramine and desipramine. Of a number of analogues of indole, N-acetyl-5-HT and 5-hydroxytryptophan were the most potent in the inhibition of the accumulation of [¹⁴C]5-HT. The presence of a large molar excess of amino acids had little effect. A small amount (less than 14 per cent) of the accumulated [¹⁴C]5-HT was metabolized to form 5-hydroxyindole acetic acid, even after long periods (2 h) of incubation. The accumulated [³H]tryptophan was metabolized to form 5-hydroxytryptophan and 5-HT; the content of formed [³H]5-HT increased with incubation time whilst the [³H]5-hydroxytryptophan remained more or less constant. The presence of p-chlorophenylalanine in the incubation medium did not interfere with the accumulation of [³H]tryptophan, though it inhibited the formation of [³H]5-hydroxytryptophan and to a greater extent [³H]5-HT. A rapid efflux of the accumulated [¹⁴C]5-HT from snail nervous tissue was observed on electrical stimulation. Slower release resulted when the Ca²⁺ ion content of the incubation medium was replaced by Mg²⁺ ions. There is also a slight efflux of radioactive substances following electrical stimulation in tissues previously incubated in [³H]tryptophan. Most of this radioactivity was attributed to the formed [³H]5-HT. The data support the idea that 5-HT is a transmitter-substance in the snail Helix pomatia, and that re-uptake of the substance is a method of inactivating the released amine.     

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)     

THE UPTAKE OF [3H]CHOLINE BY SNAIL (HELIX POMATIA) NERVOUS TISSUE IN VITRO

Abstract— When suboesophageal ganglia of the snail Helix comalia were incubated at 25°C in a medium containing [³H]choline, tissue: medium ratios of about 14:1 were obtained after 20 min incubation, and only 15°, of the accumulated choline was metabolized to form [³H]acetylcholine. The uptake of [³H]choline showed saturation kinetics and was dependent upon temperature and sodium ions. Kinetic analysis suggested the existence of a high affinity uptake process (K_m= 1.7 μM, V_max= 0.21 nmol/g/min) and a low affinity process (K_m= 100 μM, V_max= 1.2 nmol/g/min). The high affinity uptake differed from the low affinity system in that it was sensitive to various metabolic inhibitors and was competitively inhibited by low concentrations of hemicholinium- and acetylcholine. Neither uptake system was greatly influenced by the absence of calcium, potassium or magnesium ions or by the presence of low concentrations of 5-HT, dopamine. tetrabenazine, chlorpromazine, decamethonium, nalaxone or imipramine. The high affinity uptake process may be important in supplying choline for the biosynthesis of acetylcholine in cholinergic neurons.     

VARIABILITY OF THE GENITAL SYSTEM OF HELIX POMATIA L., 1758 AND H. LUCORUM L., 1758 (GASTROPODA: STYLOMMATOPHORA)

This paper examines the variability of the genital characters ofHelix pomatia and H. lucorum. Some of these characters are commonlyused to distinguish the two closely related species. Variationswithin and between populations are also investigated. The highvariability of these characters can frequently lead to identificationerrors, especially in eastern Europe. Correct identificationof these edible snails is also of commercial importance, differentspecies having a different price. The reciprocal nature of themale parts of the genitalia and the bursa stalk and diverticulumcomplex is pointed out. A possible function of the diverticulumis suggested on the basis of size relations between genitalsystem features. (Received 12 August 1999; accepted 11 April 2000)     

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)     

ANNUAL ACTIVITY CYCLES OF THE LAND SNAIL HELIX ASPERSA MULLER IN NATURAL POPULATIONS IN NORTH-WESTERN SPAIN

Activity and inactivity (dormancy) periods of the land snailHelix aspena MÜller were monitored at monthly intervalsduring two years in two natural populations in Galicia (NW-Spain),one m an area of Mediterranean climatic influence and anotherin an Atlantic -climate area The annual activity cycles of Haspena in Galicia are consistent with latitudinal adjustmentsof the life cycle of this species which shows a very wide distributionrange, but also showed some special features related to theparticular microclimate of each locality (Received 4 December 1995; accepted 6 May 1996)     

A POPULATION STUDY OF AN EQUATORIAL LAND SNAIL, LIMICOLARIA MARTENSIANA (ACHATINIDAE)

Near the equator in Uganda, the herbivorous land snail, Limicolaria martensiana , occurs in well-defined populations. These populations may be very large and the snails may in places occur at exceedingly high densities. The snails breed all the year round, but with two peaks that correspond with the two annual dry seasons. They enter dispause whenever the weather is very dry. The size of adult snails seems environmentally determined and it varies in different populations : the largest snails occur on soil rich in calcium. The shells of the snails are highly variable and polymorphic in colour and pattern. In particular, there is a streaked form that occurs in all populations, and four distinct pallid dilute forms. The relative frequency of the forms varies markedly in different populations. Polymorphism is greatest at high population densities and the snails are all streaked where they are rare. Polymorphism can be distinguished in Pleistocene fossil populations.     

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)     

GROWTH AND REPRODUCTION OF THE MINUTE LAND SNAIL PUNCTUM PYGMAEUM (DRAPARNAUD)

Reproductive characters and growth of Punctum pygmaeum wereinvestigated under laboratory conditions. Eggs were laid singlyat intervals of 2–8 days. Egg size averaged 0.41 by 0.50mm. Due to a variable period of egg retention, newly depositedeggs contained differently developed embryos resulting in anincubation period varying from 1 to 34 days. Each snail on averagelaid 6.0 eggs (range 1–16 eggs) during its life. Egg productionrate and hatching success in individuals kept in isolation werethe same as in individuals kept in pairs. Hatchlings measured0.49 mm in shell breadth. Snails grew throughout their life,but the growth rate declined significantly at a shell breadthof 1.3–1.4 mm, the point at which the onset of sexualmaturity occurred. Comparison of reproductive characters ofsnails living in leaf litter revealed that very small-sizedspecies produce few, but in relation to their shell size largeeggs, which they oviposit singly. ^*Present address: Zoological Institute, University of Basel,Rhein-sprung 9. CH-4051 Basel, Switzertand (Received 23 May 1988; accepted 11 July 1988)     

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     

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)