Yves, Carton

Directeur de Recherches Emérite CNRS

Membre du CA de la Société Entomologique de France

Diplôme d’Immunologie et Sérologie, Instirut Pasteur, 1965, Paris
Papidou avec micro 5  Thèse d’Etat, 1968, Université de Paris

lab phone : 33 1 69 15 49 67
cell phone : 06 81 65 62 60



The model system consisting of Drosophila and parasitic wasps was chosen because it was well suited for studies on development of resistance in hosts and development of virulence (protection) of parasitoids at the molecular, genetical and populational levels.

Drosophila immune reaction against a parasitoid egg                                                      Drosophila immune reaction against a parasitoid egg

larve avec capsule 4

adulte avec capsule
Drosopila adult with the mark of its immune capacity
larve avec capsule
Drosophila larva showing an immune reaction against parasitoid egg


The genetic mechanism of resistance of D. melanogaster is based on a monogenic dominant system. This resistant character is inherited autosomally, suggesting a single major segregating locus with two alleles (resistance and susceptibility). This system is highly specific and 2 genes of resistance are now localized (Rlb and Rat against L. boulardi and Asobara tabida, respectively) on the right arm of the second chromosome. We also demonstrated with the parasitic wasp Leptopilina boulardi that all the genetic components of virulence variation are comprised in a single segregating factor integrated to the wasp chromosome (as a “gene”). Two genetic systems of virulence have been detected (Ism and ISy), each against a host species, D. melanogaster and D. yakuba respectively. Encapsulation of wasp egg by its host appears to be governed by major genes in the host and in the parasitoid populations. The gene for resistance against L. boulardi in the host is counteracted by a gene for virulence against D. melanogaster immune system in the parasitoid population. Species specificity of these genes have been shown both for the host and the parasitoid, according to a “gene-for-species” model. From an evolutionnary point of view, local adaptations of the immune suppressive genes toward the host species present in the locality were detected.

These investigations of the biochemical and genetic aspects of the Drosophila-parasitoid relationships were successfully developed in part from fruitful cooperations with Professor A. Nappi (Loyola University, Chicago) and Professor Marylene Poirié (Nice University, INRA, Sophia Antipolis).
Since 2008 our collaborative efforts have produced 5 published reviews in various journals concerning co-evolutionary processes, specially “The Advances in Parasitology“.



The chapters concern the immune reaction of Drosophila larvae against

parasitoid species


I have been firstly interested in the historical development of entomology and entomologists, especially of Reaumur (1683-1757) and Buffon (1707-1788).

I also investigated various aspects of the scientific cooperation between French and American entomologists during the period 1830-1940. Perhaps one of the most successful cooperations between these two groups was between Professor Jules Planchon (1823-1888) and Professor Charles Valentine Riley (1843-1895), the famous American entomologist who saved French vineyards from attack by the louse Phylloxera by introducing American resistant Vitis species in France. It has been certainly a model scientific cooperation between the two countries

photo Riley 001
Charles valentine Riley (1843-1845), the most famous american entomologist

                                         1878-09-22 p 305

forme gallicole 2
  AMERICAN VINES :    galls on leaves (gallicole form)
forme radicicole
FRENCH VINES : galls on roots (radicicole form)

 Planchon and Riley            visting  the attacked        vineyards in                     Montpellier



I gave the opening lecture at the  conference “6th International Symposium on Phylloxera“, held at Bordeaux, on august 28-30, 2013, untitled “An exemplary partnership of American entomologist (C. V. Riley) with French entomologists in control of the Grape Phylloxera in France (1868–1895)”.

On year 2006, I discovered in the Riley’s archives (National Agricultural Library, Beltsville, MD, USA) the dedicated photograph of his French colleague friend, H. de Varigny (1855-1934) who had chosen to introduce the Darwinian theory to the French people. I recounted this relationship in my book published in 2009 dealing with the scientific life of H. de Varigny and his efforts to introduce these new evolutionary concepts in France despite the hostility of French biologists to Darwinian principals, an hostility that delayed by 8 years the election of Darwin to the French Academy of Sciences !
Preface written by Claude Combes, Professor emeritus at Perpignan university, member of the French Academy of sciences

                         couverture ouvrage Darwinisme de Y. CARTON                                      photo au café Riche

                                                                                A picture dedicated on a meeting of H. de Varigny with C.V. Riley in Paris (café Riche) on august 24, 1889

BIBLIOTHEQUE NATIONALE DE FRANCE, “Les rendez-vous du samedi”, site François-Mitterrand,  The french national library organize each week (on saturday afternoon, 17h-18h pm) an appointment with some well-known persons such as authors, movie producers, scientists, artists, whom production has retained attention.
Saturday, 23 may 2009, 17h – 18h : Yves Carton “accueil et diffusion du darwinisme en France” Yves Carton is the guest of this new appointment for his book : “ Henry de Varigny, darwinien convaincu, médecin, chercheur et journaliste (1855-1934) ” published by Hermann editor. Hervé Colinmaire, director of the department “Sciences et techniques de la BnF”, with the collaboration of Jérôme Petit, responsible of the science history collections, will animate this meeting. Darwinian theory concerning species evolution has been more or less well understood in France, with a hostile reception by the french biologists. Henry de Varigny, physician and PhD, researcher, will choose scientific journalism to better explain darwinism to the french poeple. This meeting will be follow by a dedication of his book in the library.

More recently, I have investigated a lot of english and french archives to better understand the relationships between Charles Darwin, darwinism and entomology, and specially the French entomologists. It is related in a book published  :

2011 Couverture ouvrage Ent. Darwin et Darwinisme                     Agraceum - Xanthopan

           A Book published  on “ENTOMOLOGY and DARWINISM”

Charles Darwin (1809-1882) is partially unknown for his scientific investigations as an entomologist. Entomology has had a great importance for Darwin fostering the development his theory of Natural Selection. In this book, Yves Carton investigates the influence of entomology on the different aspects of Darwin’s work.
With Darwin’s original observations, notably on bees and butterflies, the first part of the book details Darwin’s attraction for entomology, first during period spent at Cambridge University, and during the voyage of the Beagle around the world, and then in 1842, at Down House.
In the second part of the book, the author examines the importance and role of entomology in four of Darwin’s books. Publishing The Origin of the Species in 1859 induced a complete change in his relationships with English entomologists, his friends. The book was considered by many individuals as highly controversial. However, some friendly entomologists who understood veracity of Darwin’s arguments were very instrumental in fostering the spread of this new theory on species evolution.
In the third and last section of the book, the author analyzes the connections of French entomologists with Darwin and Darwinism. Generally, hostility among entomologists against the new Darwinian concepts was very common, with neo-Lamarckism representing until recently a strong influence in France. However, various institutions, such as the National History Museum, the French Entomological Society, and universities in which entomology plays an important role, all functioned, sometimes in a negative way, in advancing Darwin’s theory.
No book, even in english, has previously examined these different aspects of entomology and Darwinism.
Preface written by Patrick Blandin, Professor emeritus of the national Museum of natural History, senior Director of the “Grande Galerie de l’Evolution”, vice-President of the french entomological Society
A partial translation has been published recently in American Entomologist, 2012, 58, 4, pp. 208-212. click here


A book was  published on 2015, untitled “Histoire de l’Entomologie. Relations entre biologistes français et américains (1830-1940)


This is the story (in french !)of the relationships forged between French and American entomologists on more than a century (1830-1940)which is reported. French biology has benefited from these early contacts and exchanges without any competitive spirit at the time, but instead with pleasure to bring to his colleague assistance and knowledge. We then see build this ideal of internationalization of science.

From 1830 to 1860, the French and American entomologists want to see the other side of the Atlantic. For the former, it is above all the desire to discover a new territory. For the latter, come to France is to discover the French scientific community and somehow be dubbed by it.

From 1860 to 1900, one grasps how the American presence has to fight against this disaster in the making for French viticulture, with the discovery of phylloxera, the aphid sucker roots. Some entomologists, French but also American, were able to have a visionary approach, proposing the graft of the french vines on american vine roots recognized as resistant, and that, paradoxically, had brought evil.

From 1900 to 1920, the popularity of the biological control against crop pests came from another continent. These exchanges between entomologists have allowed them to find the most effective biological agent for acclimatization on the continent where there is the introduced pest.

From 1920 to 1940, period in which some biology visionaries,  perceived the revolution that is taking place in biological sciences in America, especially in genetics, thanks to this exceptional material that is Drosophila. One also observes the fracture that gradually develops between the two communities, that of France still believing in their successes of the 19th century.

About the contents, click on : https://cdn.shopify.com/s/files/1/0245/3579/files/203_Histoire_de_l_entomologie_tdm.pdf?6706722594174740758

About the preface, click on : https://cdn.shopify.com/s/files/1/0245/3579/files/203_Histoire_de_l_entomologie_preface.pdf?6706722594174740758


A book concerning a review of historical aspects of innate Immunity from 1865 to 2011 was edited (with an english and french versions)

Innate immunity is a new branch of immunology, confirmed by three Nobel Prize winners in 2011.
It is the first line of defense against pathogens and is in a way the preliminary step of adaptive
immunity which occurs later, and only present in vertebrates.This book examines the way in which innate immunity was discovered in invertebrates. As a starting point, it looks at the work of Louis Pasteur on silkworm disease and the findings of Ilya Metchnikov, discoverer of phagocytosis. It also investigates André Paillot, who in 1920 demonstrated the existence of humoral immunity in insects, unrelated to the type of immunity that was initially thought to be present in all vertebrates.
Finally, Innate Immunity shows how the group directed by Jules Hoffmann found strong similarities
between the innate immunity response of insects and mammals. The discovery of a receptor protein
in Drosophila, which is also found in humans, was what led to Jules Hoffmann being awarded the
Nobel Prize in 2011.

CHAPTER I : LOUIS PASTEUR and the SILKWORM DiSEASE  (1865-1870) : This chapter deals with the work undertaken by L. Pasteur on the pebrine, a parasite of the silkworm. For 5 years (1865-1870), he worked with his team in the south of France. With the publication of 25 articles and a book published, we had the material to understand his conversion from a chemist to a biologist, thus discovering the laws governing pathogenicity and immunity in an insect, to apply them later to mammals.

CHAPTER II : ELIE METCHNIKOFF, NOBEL PRIZE at THE INSTITUT PASTEUR  (1888-1916). L. Pasteur, after having highlighted the anti-bacterial immunity, wishes to understand its mechanism. In 1887, he welcomed to Paris a Russian researcher, E. Metchnikoff, who presented his phagocytic theory of resistance. Pasteur, convinced, recruited him into the new Institute that had just opened. For 30 years, Metchnikoff developed his research on immunity, training a hundred young researchers, which led him to the Nobel Prize in 1908.

ChapTER III : THE POST-METCHNIKOFF  ERA AT THE PASTEUR INSTITUTE (1920-1940) : Mechnikoff’s concepts on immunity were taken up by his students, including the most famous A. Besredka, J. Bordet (future Nobel Prize winner) and J. Cantacuzène. One of them, S. Metalnikov. recruited in 1920 at the Pasteur Institute, will develop an entire school on insect immunity, promoting the cellular aspect of this resistance. They discover the mechanisms of this resistance, the basis of innate immunity

CHAPTER IV : PAILLOT THE DISSIDENT IN THE ASSAULT OF THE PHAGOCYTE DOGMA: HUMORAL IMMUNITY IN INSECTS (1920-1935) : The purely cellular conception of immunity could not be maintained, given what was found with antibodies in mammals. A young researcher from agronomic research, A. Paillot, will be able to demonstrate humoral immunity in insects, but totally different from what we observe with antibodies. It will be the humoral component of innate immunity.

CHAPTER V : The BARREN SPELL FOR INVERTEBRATE IMMUNOLOGY (1960-1990) : It was not until the 1950s that research on the immunity of invertebrates resumed. But it will be for the researchers who addressed these questions, a real barren spell. Vertebrate immunity reigned supreme with the discovery of immunoglobulins, their specificity and immunological memory. It had to be believed and various teams in France led the fight, mainly with the transplant tool. 

CHAPTER VI : TOWARDS THE NOBEL PRIZE (2011) OF J. HOFFMANN IN STRASBOURG :  In 1980, the team of H. Boman (Sweden)  isolated the first antimicrobial peptide, cecropin, from a large caterpillar (the butterfly H. cecropia) by biochemical techniques. Later, molecular genetics made it possible to work on Drosophila, which was undertaken by J. Hoffmann’s team in Strasbourg. Several antibacterial peptides will be isolated in Drosophila. The team will then discover the two signalling pathways for the production of these peptides, the Toll pathway and the IMD pathway, as well as the PRRs (pattern recognition receptors) as recognition factors of the pathogenic bacteria. This permitted the discovery of this Toll receptor in mice and humans (on the macrophage membrane). This shows how this innovative results have made it possible to bridge the gap between innate and adaptive immunity.


To download the papers pdf, click on :


  • LUE, C.-H., BUFFINGTON M. L., SCHEFFER S., LEWIS M., ELLIOT T. A., LINDSEY A. R. I., DRISKELL A., JANDOVA A., KIMURA M. T., CARTON Y., KULA R. R., Schlenke T. A., Mateos M., Govind S., Varaldi J., Guerrieri E., Giorgini M., Wang X., Hoelmer K., … Hrcek J. (2021).  Molecular voucher database for identification of Drosophila parasitoids. Molecular Ecology Resources, pp. 1–18. https://doi.org/10.1111/1755-0998.13435

  • CARTON Y., 2019. Innate immunity : from Louis Pasteur to Jules Hoffmann (1865 – 2011), Ed. Iste editions Ltd, London, 322 pages
  • CARTON Y., 2019. Immunité Innée : de Louis Pasteur à Jules Hoffmann, prix Nobel (1865 – 2011), Iste éditions Ltd, London, 317 pages. (à paraitre)
  • CARTON Y., 2016. Histoire de l’Entomologie. Histoire de l’Entomologie. Relations entre biologistes français et américains (1830-1940), Iste editions, London, 208 pages.
  • CARTON Y., 2014. L’oeuvre de Charles Bocquet (1918-1977), Directeur du LGEB du CNRS de 1965 à 1977, dans “Le laboratoire CNRS de génétique Evolutive de Gif : de part et d’autre de l’oeuvre de Georges Teissier“, sous la direction de laurent Loison, ed. Hermann Paris, 198 pages.
  •  DAVID J. et Y. CARTON, 2014. Génétique des populations et allométrie dans l’œuvre scientifique de Georges Teissier, dans “Le laboratoire CNRS de génétique Evolutive de Gif : de part et d’autre de l’oeuvre de Georges Teissier“, sous la direction de laurent Loison, ed. Hermann Paris, 198 pages.
  • CARTON Y., 2013. L’entomologie française et le Muséum face au transformisme (1832 – 1900), Bull. Amis du Muséum natinal Histoire naturelle, n° 253, pp. 33-36
  • CARTON, Y. 2013. Le Phylloxera de la vigne, dans Des Insectes et des Plantes, ed. N. Sauvion, P.A. Calatayud, Thiery D., Marion-Poll F., Editions Publibook et IRD, pp. 698-704
  • DUPAS  S.,  POIRIE M., FREY F. and Y. CARTON, 2013. Is parasitoid virulence against multiple hosts adaptive or constrained by phylogeny ? The study of Leptopilina spp. (Hymenoptera: Figitidae) / Drosophila (Diptera: Drosophilidae) interactions.,  Annls Soc. Entomol. Fr, vol. 49, (2), pp. 222-231.
  • CARTON Y., 2013. Une histoire de la lutte biologique contre les ravageurs, Biofutur, 343, pp. 27-32
  • CARTON Y. 2012 Tne Entomological Society of France with regard to darwinism, after 1859, American  Entomology,  vol .  4,    pp. 208-212
  • CARTON, Y. 2011 Entomologie, Darwin et Darwinisme, editions Hermann Paris, coll. Histoire des Sciences, 247 pages
  • A. DUBUFFET, D. COLINET, C. ANSELME, S. DUPAS,Y. CARTON AND M. POIRIE 2009 Variation of Leptopilina boulardi Success in Drosophila Hosts: What is Inside the Black Box? Advances in Parasitology 70: 148
  • A. NAPPI, M. POIRIE and Y. CARTON 2009 The Role of Melanization and Cytotoxic Byproducts in the Cellular Immune Responses of Drosophila Against Parasitic Wasps Advances in Parasitology 70: 100
  • CARTON Y. 2009 Accueil et diffusion du darwinisme en France. Henry de Varigny(1855-1934), médecin, Chercheur et journaliste, un darwinien Convaincu Médecine et Sciences, 25(4): 413
  • CARTON Y. 2009 Henry de Varigny : un journaliste contre l’antidarwinisme. Pour la Science 379(4): 80
  • CARTON Y., FREY F. AND A. NAPPI 2009 Parasite-induced changes in nitric oxide levels in Drosophila paramelanica Journal of Parasitology 95(5): 1134
  • POIRIE M., CARTON Y. AND A. DUBUFFET 2009 Virulence strategies in parasitoid Hymenoptera as an example of adaptative diversity. Comptes Rendus Biologies. Spécial issue : Darwin’s theory revisited by today’s biology 332(4): 311
  • S. DUPAS, A. DUBUFFET, Y. CARTON, AND M. POIRIE 2009 Local, Geographic and Phylogenetic Scales of Coevolution in Drosophila–Parasitoid Interactions Advances in Parasitology 70: 282
  • CARTON Y. 2008 Henry de Varigny, Darwinien convaincu, médecin, chercheur et journaliste (1855-1934). Editions Hermann, Paris, 126 pages 0: 0
  • CARTON Y., POIRIE M. AND NAPPI A.J. 2008 Insect immune resistance to parasitoids. Insect Science , 15: 68
  • DUBUFFET A., G. DOURY, LABROUSSE C., J.M. DREZEN, Y. CARTON AND M. POIRIE 2008 Variation of success of Leptopilina boulardi in Drosophila yakuba: the mechanisms explored. Developmental and Comparative Immunology, 32: 598
  • SORENSEN C., SMITH E., SMITH J. AND CARTON Y. 2008 Charles V. Riley, France and Phylloxera. American Entomologist 3: 134
  • CARTON Y., SORENSEN C., SMITH J. SMITH E. 2007 Une coopération exemplaire entre entomologistes français et américains pendant la crise du Phylloxera en France (1868-1895) Ann. Soc. Entomol. Fr., 43(1): 103
  • CARTON Y. 2007 Le Phylloxera de la vigne. Annales Muséum Sciences naturelles de Perpignan 15: 47
  • DAVID J. CARTON Y. 2007 Georges-Louis Leclerc, comte de Buffon (1707-1788) un homme d’influence au siècle des Lumières. Medecine Sciences, 23(11): 1057
  • KOHLER L.J., CARTON Y., MASTORE M. and NAPPI A.J. 2007 Parasite suppression of the oxidations of eumelanin precursors in Drosophila melanogaster. Arch. Insect Biochem. Physiol., 66: 64
  • CARTON Y. 2006 La découverte du Phylloxera en France : Un sujet de polémique : les archives parlent (Hemiptera, Chermesidae).111, (3), 305-316 Bull. Soc. Entomol. Fr., 111(3): 305
  • DUBUFFET A., DUPAS, S.,, FREY, F., and DREZEN, J.-M., POIRIE, M. AND CARTON, Y. 2006 Genetic interactions between a parasitoid wasp and its Drosophila hosts. Heredity, 97(4): 1
  • HITA M.,, ESPAGNE E.,, LEMEUNIER F., and L. PASCUAL, Y. CARTON, G. PERIQUET AND M. POIRIE. 2006 Mapping candidate genes for Drosophila melanogaster resistance to the parasitoid Leptopilina boulardi. Genetical Research, Camb.,, 88: 81
  • CARTON Y. 2005 . Reaumur (1683-1757) :le véritable fondateur de l’Entomologie en France. Bull. Soc. Entomol. Fr., 109: 445
  • CARTON Y.,, NAPPI A., and POIRIE M. 2005 Genetics of anti-parasite resistance in Invertebrates. Developmental and Comparative Immunology, 29: 9
  • CARTON, Y. 2005 Reaumur (1683-1757) : The discoverer of parasitoids in France Biological Control 32: 40
  • NAPPI A.J., FREY F. and CARTON Y. 2005 Drosophila serpin 27A is a likely target for immune suppression of the blood cell-mediated melanotic encapsulation response. J. Insect Physiol., 51, 197-205 0: 0
  • FLEURY, F., RIS, N., FOUILLET, P., CARTON, Y. and M. BOULETREAU 2004 Genetic and ecological interactionsin Drosophila -Parasitoids communities : a case study with Drosophila melanogaster, D. simulans and their Leptopilina species. Genetica (Special issue on Genetics of Speciation). 120(1): 181
  • NAPPI A.J., VASS E., MALIGOLI D. and CARTON Y. 2004 The effects of parasite-derived immune suppressive factors on the cellular innate immune and autoimmune responses of Drososphila melanogaster Jounal of Parasitology 90(2): 1138
  • DUPAS, S., CARTON, Y. and POIRIE, M 2003 The genetic dimension of the coevolution of virulence resistance in Drosophila – parasitoid wasp relationships. Heredity 90(1): 84
  • LABROSSE, C., CARTON, Y., DREZEN, J.M. and POIRIE, M. 2003 Active suppression of D. melanogaster immune response by long gland products of the parasitic wasp Leptopilina boulardi J. Insect Pysiology 49: 513
  • Y. CARTON, F. FREY, D.W. STANLEY, E. VASS and A.J. NAPPI 2002 Dexamethasone inhibition of the cellular immune response of Drosophila melanogaster against a parasitoid Journal of Parasitology 88: 405
  • ALLEMAND, R, LEMAITRE, C., FREY, F. and BOULETREAU, M. \VAVRE, F.\NORDLANDER, G.\VAN ALPHEN, J. AND Y. CARTON 2002 Phylogeny of six african Leptopilina species (Hymenoptera, Cynipoidea, Figitidae), parasitoids of Drosophila, with the description of three new species Ann. Soc. Entomol., Fr, 38(4): 319
  • CAMPAN, E, COUTY, A, CARTON, Y and PHAM-DELEGURE, M AND L. KAISER 2002 Variability and genetic components of innate fruit odor recognition in a parasitoid of Drosophila Physiological Entomology 27: 243
  • SORRENTINO, R.P., CARTON, Y. and GOVIND, S. 2002 Cellular immune response to parasite infection in the Drosophila lymph gland is developmentally regulated Developmental Biology 243: 65
  • RUSSO J. /BREHELIN M., CARTON Y. 2001 Haemocyte change in resitant and susceptible strainns of D. melangaster caused by virulent and avirulent strains of the parasitic wasp Leptopilina boulardi. J. Insect physiol. 47: 167
  • Y. CARTON AND A. NAPPI 2001 Immunogeneitic aspects of the cellular immune response of Drosophila against parasitoids Immunogenetics 52: 157
  • BENASSI V., COUSTAU C. and CARTON Y. 2000 Insect immunity: A genetic factor (hrtp) is essential for antibacterial peptide expression in Drosophila after infection by parasitoid wasps. Arch. Insect Biochem. Physiol. 43: 64
  • NAPPI A.J., VASS E., FREY F. and CARTON Y. 2000 Nitric oxide involvement in Drosophila immunity. Nitric Oxide: Biology and Chemistry 4: 423
  • POIRIE M., FREY F., HITA M., HUGUET E., LEMEUNIER F., PERIQUET G. and CARTON Y. 2000 Drosophila resistance genes to parasitoids: chromosomal location and linkage analysis. Proc. Roy. Soc. London 267: 1417
  • DUPAS S. and CARTON Y. 1999 Two non-linked genes for specific virulence of Leptopilina boulardi against Drosophila melanogaster and D. yakuba. Evol. Ecol., 13: 211
  • HITA M., POIRIE M., LEBLANC N., LEMEUNIER F., LUTCHER F., FREY F., PERIQUET G. and CARTON Y. 1999 Genetic localization of a Drosophila melanogaster resistance gene to a parasitoid wasp and physical mapping of the region. Genome Research 9: 471
  • DUPAS, S., FREY, F. and CARTON, Y. 1998 A single parasitoid segregating factor governs the suppression of Drosophila immune reaction, Journal of heredity,  89(4): 306
  • CARTON Y., DUPAS S. 1998 Genetic of resistance and virulence in host parasioid system, inLaboratory manual on techniques in Insect Immunology” A. Wiesner, AG. Dunphy, V.J. marmaras, I. Morishima, M. Sugumaran and M. Yamakawa ed, SOS publications, Fair Haven, NJ “Laboratory manual on techniques in Insect Immunology” 0: 279
  • CARTON Y., A.NAPPI 1997.  Drosophila cellular immunity against parasitoids, Trends in Parasitology, 13, pp. 218-226.
  • CARTON, Y., 1997. Le Phylloxera de la vigne, dans “Des Insectes et des Plantes“, ed. N. Sauvion, P.A. Calatayud, Thiery D., Marion-Poll F., éditions Publibook et IRD, pp. 698-704.

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