J. Moll Stud.

(1998), 64,173-181

The Malacological Society of London 1998

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

ANNETTE GOMOT
Laboratoire de Biologie des Organismes et Ecosystimes, U.F.R. des Sciences et Techniques, place Marichal Leclerc, 25030 Besancon Cedex, France (Received 16 September 1996; accepted 24 May 1997)

to the species, there is no information on their comparative chemical composition. Only the This paper describes the biochemical composition of nutritive value of farmed Helix aspersa of different species (Helix lucorum. Helix pomatia) and relatively low bodyweight has been evaluated sub-species of snails (Helix aspersa aspersa, Helix (Clayes & Demeyer, 1986). aspersa maxima) reared in the same conditions with a Many snail farms are being established at the feed ('Helixal') specially designed for edible snails. present time, in part to compensate for the In addition, the composition of wild H. pomatia and decrease in natural populations in certain H. lucorum is presented to allow comparison countries and in part in order to produce goodbetween snails of different origins. Analyses determined the percentages of proteins, lipids and quality snails for consumption. Thus it seemed minerals. They reveal both similarities and differuseful to make a comparative study of differences in composition according to the species and the ences in biochemical composition between the part analysed (whole body, pedal mass, and visceral most commonly eaten species. With that end in mass). H. pomatia contains the highest percentage of view, analyses of snails of clearly identified orimineral matter and the lowest percentage of lipids. gin were made. In addition, these snails were Surprisingly, protein contents are slightly different subject to genetic studies in our laboratory between artificially reared H. aspersa maxima of 3 months old and wild H. pomatia. The results make (Borgo, Souty-Grosset & Gomot, 1995; Borgo, Souty-Grosset, Bouchon & Gomot, 1996) it possible to evaluate nutritional quality of snails which allowed an up-to-date and accurate with the composition of the body of four edible snail species. identification. Some were collected from wild populations (which are still the main source of supply for human consumption), while others were reared in a controlled environment that INTRODUCTION allowed us to study the effect of different factors (species, age and environmental paramWhile detailed analyses are available for many foods (Alais & Linden, 1991), there are few eters) on the biochemical composition of their data on the composition and nutritional value tissues. of edible snails. The essential amino-acid composition of the molluscan pest Achatina fulica Bowdich was determined by Mead & MATERIALS AND METHODS Kemmerer (1953) to determine the possibility of using snail meat as a source of animal The animals protein in feed for poultry and livestock. The study was conducted on four species or subFor Helix pomatia L. and Helix aspersa species of the genus Helix: Helix aspersa aspersa, Muller, the information available up to 1975 on Helix aspersa maxima Taylor, Helix pomatia and their composition of proteins, lipids, carbo- Helix lucorum L. The majority of the analyses were hydrates and minerals was summarized by of animals reared in off-the-ground cages specially Cadart (1975), who remarked on their richness designed for snail rearing and in optimal environin mineral salts. Since that date some informa- mental conditions for growth (photoperiod 18L : 6D; 20C; relative humidity 90%) (Gomot & tion has appeared on species that are not temperature Deray, 1987). The H. a. aspersa came from a strain always well defined, following various treat- from the Cavaillon region of France. The H. aspersa ments of the flesh (Table 1). Thus, in spite of maxima, originally imported from Algeria, had been the fact that there is a significant international raised in France for 30 generations. The H. pomatia trade in snails and prices vary greatly according parent snails were collected in the Choye forest (70ABSTRACT

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174

A. GOMOT

Table 1. Data available in the literature on the biochemical composition of snails for the same or related species. Species Whole Raw Snail Helix pomatia Raw Snail species not defined Raw Giant African Snail Achatina H. pomatia Raw Foot H. pomatia Edible Part H. pomatia Whole Snail H. aspersa Whole Raw Snail small snails (2.5 g) large snails (3.5 g) H. pomatia H. aspersa Reared Snail Raw Snail Oreohelix- strigosa Whole Snail 79.2 89.2 72.8 to 80.7 79 84.9 16 12.3 1 0.7 2 1 1.9 16.1 9.9 Water (%) gper 100 g fresh matter Proteins Lipids Carbohydrates 0.79 1.4 1.4 2 4.4 1.3 2.1 Authors Ash Oudejans & Van der Host, 1974 Watt & Merril, 1975 Watt & Merril, 1975 Wieser & Schuster, 1975 Souci etal., 1981 Avagnina, 1983 Downloaded from http://mollus.oxfordjournals.org/ by guest on December 4, 2013

83.3 87.6 84.9 81.6 79 78 to 81

12 9.9 12.6 16.3 16 10.4

0.7 0.5 0.5 0.8 1 1.5

0.5 0.4

2.7 1.2 1.8 1.3

Claeys & Demeyer, 1986 Fontanillas, 1989 Bonnet etal., 1990 Feinberg etal., 1991 Rees & Hand, 1993

2 4.6

France) and the H. lucorum came from a population established at Caluire in the Lyon area (69-France). The rearing method was as follows: Following mating of the parents, the eggs laid were collected and the young that hatched were reared in cages above ground on a meal (E3-2) prepared specially for snails (Trademark Helixal: Ets Lupine Alivor - 39 Clairvaux les Lacs - France). The analyses of the snail food was: proteins 13.4%, crude fat 4.3%, cellulose 2.5%, ash 31.4%, calcium 10.8%, vitamins A, D3, E respectively 15 000, 2 000 IU.kg"1 and 20 mg.kg"1. Sample preparation This was performed on a series of artificially reared animals from 3 to 5 months of age and also on adults of the species H. pomatia and H. lucorum collected from wild populations in order to determine whether there are differences in these species that are a function of their origin and age. The snails for analysis were starved for 24 hours, then decapitated and separated from their shell. The whole body or its different parts were put into plastic beakers kept in ice. Analyses were performed both on the whole body and on the two principal parts: the pedal mass, with the head and mantle edge; and the visceral mass, containing the digestive gland, gonad,

albumen gland, genital ducts, kidney and heart. The mucus and hemolymph leaking during separation of the two parts was combined with the foot. The number of animals sampled per lot was chosen to give 130-200 g of organs before dehydration, the quantity necessary to perform the various analyses on the same pool of snails. The soft tissues were rapidly frozen at -70C and stored at that temperature. They were then freezedried to constant weight to determine the moisture content. The freeze-dried tissues were then ground in a knife mill (Janke & Kundel A10) and reduced to powder by passing through a sieve with holes 0.25 mm in diameter. The shells were dried at 60C to constant weight and the percentage shell to total liveweight of the animal was calculated. The crude ash content of the samples was determined by calcination of 2 g of powder in a muffle furnace (Pyrecton KY type KY 2C4 Prolabo) at 550 5C for 6-7 hours. Analysis of the organic matter Protein content was determined by the colorimetric method of Lowry, Rosebrough, Farr & Randall (1951). Total lipids were determined by extraction with a 2:1 V/V chloroform:methanol mixture and

BIOCHEMICAL COMPOSITION OF HELIX titrated according to Folch, Lees & Stanley (1957). Total monosaccharides were determined at Biological Chemistry Laboratory of Lille University (Dr Michalsky and F. Delplace) by gas chromatography after methanolysis and trimethylsilylation following Kemerling, Gerwig, Vligenthart & Clamp (1975).

175

stituents of the tissue, the latter are expressed as percentage dry matter. They are also given as a ratio to fresh weight to provide a comparison with the flesh of other animals. The ash content was highest in the wild H. pomatia and H. lucorum. Among the artificially reared snails, H. pomatia had the highest ash Determination of minerals (Ca, Mg, P) and trace content (11.7% at 3 months and 12.6% at 5 elements (Cu, Fe, Zn) months). The other three species are similar in their ash content (8.7 to 9.4%). With H. pomaFollowing ashing of the samples, Ca, Mg, Fe, Cu and Zn were determined by atomic absorption tia the influence of age can be seen on the ash spectrometer (Perkin-Elmer 1100). Phosphorus was content, dry matter and proportion of the shell determined using a Technicon autoanalyser with to liveweight as the animal goes from 3 to 5 nitro-vanado-molybdic reagent. months of age with the same feed (Table 2). Initially, determinations were made on 5 snails The percentage of protein, like that of moisanalysed individually to calculate the standard deviature, differs little between artificially reared H. tion and on pools of from 5 to 10 snails. The results aspersa maxima and wild H. pomatia; these are being very close, the figures in the tables are the the lowest percentages (55 to 59.3%). Artifivalues obtained from analyses of pools of animals, which are mean values. cially reared H. pomatia has a higher percentage protein, close to that of artificially reared H. a. aspersa and H. lucorum (65.2 to 68.2% at 3 months and 72.5% for H. pomatia at 5 RESULTS months). For H. lucorum one notes a difference between the two lots collected from wild Composition of the body offour species of populations, the cause of which might be a Helix snails difference in the development of the reproducThe composition of the whole body was tive apparatuslarger or smaller albumen studied on artificially reared snails of the same gland according to whether the snails were age (3 months) of each of the 4 species (Table collected before or after egg-laying. 2). Determinations were also made on H. The percentage of lipids is lowest in wild H. pomatia at 4 months and 5 months, since this lucorum and H. pomatia, with differences of species, like H. lucorum, grows more slowly 1.9% and 3.1% between the two lots analysed than H. a. aspersa or H. aspersa maxima. This for these species. The percentage lipids in the permitted identification of changes in composi- artificially reared snails of all the species is tion with age. higher than in the wild ones, in spite of the fact The weight of the shell as a percentage of the that the latter are older. Of the 4 species, H. total liveweight was between 10.3% and 11.5% pomatia contains the least lipids and one also notes a decrease between the ages of 3 and 5 for snails of the same age of the 4 species fed on the same feed E3-2 (Table 2). The shell as a months. percentage of liveweight is much higher in the The percentage of total sugars is commonly snails (H. pomatia and H. lucorum) collected obtained as a difference between 100% and from wild populations (17.5% and 22%), the sum of protein + lipids + ash. With this largely due to the animals being older (over 2 calculation one obtains 23.7% of total sugars years of age) but also to other factors like the for H. aspersa maxima. In contrast, an accurate nature of the soil and vegetation, as well as the determination of sugars by gas chromatoginfluence of the seasons (interrupted growth, raphy gives 13.8% for this species (Table 2), hibernation, etc.). which shows that the calculation by difference is unsatisfactory. The moisture content of snails varies with the temperature and humidity of their environment. However, in the identical conditions in which sampling was performed, artificially Comparison of the biochemical composition of reared H. aspersa maxima has a moisture the foot and viscera in the four species content very close to that of wild H. pomatia, Snails are prepared for human consumption in whereas H. a. aspersa has a slightly lower different ways, depending on the species and moisture content, close to that of H. lucorum. the region. Petits Gris (//. a. aspersa) are To prevent differences in moisture content generally eaten whole. With the larger species affecting the percentages of the different con- (H. pomatia, H. lucorum and H. aspersa

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Table 2. Analyses of the whole body of four snail species. Results : g per 100 g Lipids /FM 65.2 58.5 57.6 9.1 7.2 8.1 10.8 11.9 10.7 11.7 12.6 12.6 15.8 17.7 14.6 13.1 1.8 2.2 1.8 10.7 12.8 8.6 7.3 58 67.8 68.2 60 68.1 72.5 59.3 55 1.6 1.8 1.2 1.5 1.5 1.1 0.6 1.1 0.9 0.7 0.4 Proteins (Lowry) /FM /DM 10.5

Species Shell 11.2 1.5 1.3 1.1 1.2 1.4 2.3 8.8 8.7 9 11.5 10.9 11.2 10.8 17.5 19 10.3 12.7 22.2 18.1 84.2 82.3 85.3 86.8 84 82.4 82.1 16 17.5 17.8 85.8 14.1 84.3 87.4 15.6 12.5 83.7 16.2 9.4

Age (months) 9.1 16.3 15.7 16 4.8 15.9 19.6 5.6 8.9 24.2 26.9

Snail feeds

Number of animals

# ** Water
# Ash Dry matter /FM /DM

Mean weight <g>

/DM 10 12 9.7 10.8 9.7 6.5 3.4

Carbohydrates /FM /DM

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H.aspersa maxima

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8 10

>

1.7

13.8

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3 ?

E3-2 [Natural mid July]

36 8 6

9.2 13.4

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Mean

E3-2

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3 5 ?

[Natural end June]

24 17 6 6

7 5.5 5 3.1

*: % in weight of healthy snails; **: % in weight of deshelled snails.; FM: Fresh matter.; DM: Dry matter.

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BIOCHEMICAL COMPOSITION OF HELIX maxima) the visceral hump is removed, since it has a bitter taste, and only the fleshy part 6 .o foot, head and mantle edgeis eaten. To take CD Q account of these differences in culinary u (A <D I t Z practice, the foot and viscera of snails fed with (0 the same food were always analysed separately. ^^ m in CO CM Table 3 reveals common features but also some Q CO specific differences. XI The common features include: a relatively 00 CO 'a LJ o co constant foot/viscera fresh-weight ratio of 72.8 1.2% for the foot, 27.1 1.2% for the co r^ viscera; the dry matter of the viscera exceeds in co CO CO that of the foot, with different ratios according 1 to the species (x 1.9 on average): 1.64:1 (H. CO - CO lucorum); 1.82:1 (//. pomatia); 1.96:1 (H. aspersa 0 0 OO o o | maxima); 2.36:1 (H. a. aspersa); the lipids are CL o> always at a higher concentration in the viscera than in the foot (x 1.8 on average), but as for oo CM d en the dry matter the ratio varies with the species: CO "~ LU 1.2:1 (H. pomatia); 1.8-1.9:1 (H. lucorum, H. a. aspersa); 2.5:1 (H. aspersa maxima). CO CO t - CM tt The differences include: the protein propor tion, which varies with the species and part of CD the body. The percentage of proteins is similar CO CD e CD . C CM en in the foot and viscera of H. a. aspersa and H. Q - CM lucorum, the latter being richer in proteins than c H. a. aspersa in both parts. In H. pomatia the a > in co visceral mass contains a higher protein level 03 * that the foot, while for H. aspersa maxima the * reverse is true. In addition, the percentage ash is similar in the viscera and foot of H. aspersa maxima, while in the other 3 species, mineral r- CM matter is more abundant in the foot than in the co ~T viscera. o x> Overall, for both parts of the body, one notes O -O that the dry matter of artificially reared H. *pomatia and H. lucorum is richer in proteins o cq than that of H. a. aspersa and H. aspersa maxima, while the opposite is true for lipids. * V)
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For P, the differences are less marked than for Ca, since the concentration in H. pomatia is respectively 1.6, 1.7 and 2.3 times that in H.

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The comparative analysis of the biochemical composition of raw snails presented here is the first complete study on these animals. It reveals

BIOCHEMICAL COMPOSITION OF HELIX

179

both similarities and differences between species. Until now only fragmentary data have been available, sometimes without clear identification of the species (Table 1). In addition, heterogeneous sampling methods and analysis of animals of differing origins have led to major differences in reported protein content for the same species (from 9.9% to 16.3% for H. aspersa). All this provides justification for the present analysis of different species reared in the same conditions. The results are much more homogeneous for each species (Tables 2, 3, 4) and clearly show the role of genetic and physiological factors on the composition of the snail. Account must be taken of these factors in interpretation of biochemical analyses and in the development of a snail food that promotes the nutritional qualities of their meat. The principal genetic and physiological characteristics of the four species of snail studied are summarized and discussed below. The genetic factor that should be most emphasised is growth rate. This allows us to distinguish two groups of species: H. a. aspersa and H. aspersa maxima with a rapid growth rate and adult weight attained at the age of 3 months in favourable environmental conditions; and H. pomatia and H. lucorum with a slower growth rate and more than 5 months required to attain adult weight. Among the most significant differences, we can underline that when fed on the same feed (E3-2) H. pomatia and H. lucorum were the richest in protein ( 68% of dry matter) followed by H. a. aspersa (65%) and H. aspersa maxima (58%) (Table 2) whereas for lipids, H. pomatia contains the least. Lastly, H. pomatia has the highest content of Ca, P, Cu and Fe (Table 4). In fact, this species often presents the clearest differences. Among the physiological factors, age and environmental factors are those that appear most clearly. This can be seen in the relative weight of the shell in H. pomatia, which rises from 10.3% of liveweight at 3 months to 12.7% at 5 months and 24.2% in the adults collected from wild populations at an age of from 3 to 5 or 6 years. On the other hand, the lipid content of H. pomatia decreases with age and the same phenomenon has been observed with H. a. aspersa and H. aspersa maxima. This finding is new and specific to snails, since in other farmed animals ageing is accompanied by an increase in percentage of lipids (Beitz, 1985).

The influence of environmental factors requires further study, especially of snails collected from wild populations in different biotopes, to analyse the influence of the vegetation consumed and the nature of the soil, since we have previously shown that this last plays a very important role on the growth of H. a. aspersa (Gomot, Gomot, Boukraa & Bruckert, 1989). In previous studies we have also determined the optimum conditions of photoperiod, temperature and stocking which induce the endocrine factors necessary for fast and balanced development of edible snails (Gomot & Deray, 1987; Gomot & Gomot, 1995). The results presented in this paper illustrate the way in which genetic differencesrevealed by electrophoresis of the muscle proteins of the foot, to distinguish between the meat of H. pomatia and Achatina fulica (Bracchi, 1988) or by study of mitochondrial DNA (Borgo et al., 1995)translate into differences in the proportions of the essential biochemical constituents of the tissues of these species. In order to position the snail relative to other foodstuffs, we report (Table 5) the average composition of various, types of meat and fish (from Kayser, 1963) compared with our analyses of the flesh of H. aspersa maxima ready for consumption (the majority of the digestive gland being removed and the remaining flesh boiled in a court-bouillon). The snail is an advantageous foodstuff from a dietary point of view in that it is a source of proteins while remaining low in calories, its energy value (calculated from the conversion coefficients proposed by Dupin, Cuq, Malewiak, LeynaudRouaud & Berthier, 1992) being lower than that of the leanest meat or fish. In conclusion, the value of the complete snail rearing system developed in our laboratory for rational study of the influence of factors affecting development and biochemical composition must be emphasised. Control of the various parameters makes it possible on the one hand to influence the quality of the animals reared for human consumption without the possible effects of pollutants which may occur in the field such as bioaccumulation of heavy metals (Martin & Coughtrey, 1982; Dallinger & Wieser, 1984; Campani, Bracchi, Guizzardi, Madarena & Del Bono, 1992).

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ACKNOWLEDGEMENTS This work has been performed in the framework of a CIFRE convention (n 607-90) of the Association

180

A. GOMOT Nationale de la Recherche Technique between the Zoology-Embryology Laboratory of the University of Franche-Comte' and the Institut de Recherches et d'Innovation Scientifiques of Paris. We have benefited from the collaboration of specialists, whom we warmly thank, for analysis of sugars (Dr J.C. Michalsky and F. Delplace: Biological Chemistry Laboratory, Lille University), for the analysis of proteins and minerals (M. Regnier, UCAAB, Chateau-Thierry) and metals (Prof. J.C. Pihan, Ecotoxicology Laboratory of the University of Metz). We thank Pr L. Gomot and Pr C.R. Marchand for many fruitful discussions during the course of these studies. We are also very grateful to Brigitte Jolibois for the preparation of the manuscript and very much indebted to Dr LJ. Elmslie for his help with its translation. REFERENCES
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22
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ALAIS, C. & LINDEN, G. 1991. Biochimie alimentaire.

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Masson, Paris. AVAGNINA, G. 1983. Composizione chimica e valore alimentare. In: Principi di Elicicoltura, 101-103. Edagricole, Bologna. BEITZ, D.C. 1985. Physiological and metabolic systems important to animal growth: an overview. Journal of Animal Science, 6L 1-20.
BONNET, J.C, AUPINEL, P. & VRILLON, J.L. 1990.

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L'escargot Helix aspersa: biologic, ilevage. INRA, Paris.

BORGO, R., SOUTY-GROSSET, C. & GOMOT, L. 1995.

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