Y. Akamatsu, N. Tsurushita, F. Nagawa, M. Matsuoka, K. Okazaki et al., Essential residues in V(D)J recombination signals, J. Immunol. Balt. Md, vol.1950, issue.153, pp.4520-4529, 1994.

J. C. Almagro, I. Hernandez, M. Del-carmen-ramirez, and E. Vargas-madrazo, The differences between the structural repertoires of VH germ-line gene segments of mice and humans: implication for the molecular mechanism of the immune response, Mol. Immunol, vol.34, 1997.

P. M. Alzari, S. Spinelli, R. A. Mariuzza, G. Boulot, R. J. Poljak et al., Three-dimensional structure determination of an anti-2-phenyloxazolone antibody: the role of somatic mutation and heavy/light chain pairing in the maturation of an immune response, EMBO J, vol.9, pp.3807-3814, 1990.

A. G. Amit, R. A. Mariuzza, S. E. Phillips, and R. J. Poljak, Three-dimensional structure of an antigen-antibody complex at 2.8 A resolution, Science, vol.233, pp.747-753, 1986.

S. Barr-e, A. S. Greenberg, M. F. Flajnik, and C. Chothia, Structural conservation of hypervariable regions in immunoglobulins evolution, Nat. Struct. Biol, vol.1, pp.915-920, 1994.

M. Bascove and J. Frippiat, Molecular characterization of Pleurodeles waltl activation-induced cytidine deaminase, Mol. Immunol, vol.47, 2010.

M. Bascove, N. Gu-eguinou, B. Schaerlinger, G. Gauquelin-koch, and J. Frippiat, Decrease in antibody somatic hypermutation frequency under extreme, extended spaceflight conditions, FASEB J. Off. Publ. Fed. Am. Soc. Exp. Biol, vol.25, 2011.

M. Bascove, C. Huin-schohn, N. Gu-eguinou, E. Tschirhart, and J. Frippiat, Spaceflight-associated changes in immunoglobulin VH gene expression in the amphibian Pleurodeles waltl, FASEB J. Off. Publ. Fed. Am. Soc. Exp. Biol, vol.23, 2009.

A. G. Betz, C. Rada, R. Pannell, C. Milstein, and M. S. Neuberger, Passenger transgenes reveal intrinsic specificity of the antibody hypermutation mechanism: clustering, polarity, and specific hot spots, Proc. Natl. Acad. Sci. U. S. A, vol.90, 1993.

M. Bogue, S. Gilfillan, C. Benoist, and D. Mathis, Regulation of N-region diversity in antigen receptors through thymocyte differentiation and thymus ontogeny, Proc. Natl. Acad. Sci. U. S. A, vol.89, 1992.

N. Boudarra, C. Frippiat, C. Dournon, and J. Frippiat, An alternative internal splicing site defines new ikaros isoforms in Pleurodeles waltl, Dev. Comp. Immunol, vol.26, pp.659-673, 2002.

R. Boxio, C. Dournon, and J. Frippiat, Effects of a long-term spaceflight on immunoglobulin heavy chains of the urodele amphibian Pleurodeles waltl, J. Appl. Physiol. Bethesda Md, issue.98, 1985.

P. H. Brodeur and R. Riblet, The immunoglobulin heavy chain variable region (IghV) locus in the mouse. I. One hundred Igh-V genes comprise seven families of homologous genes, Eur. J. Immunol, vol.14, 1984.

R. Castro, L. Jouneau, H. Pham, O. Bouchez, V. Giudicelli et al., Teleost fish mount complex clonal IgM and IgT responses in spleen upon systemic viral infection, PLoS Pathog, vol.9, 2013.
URL : https://hal.archives-ouvertes.fr/hal-00779835

J. Charlemagne, Noninbred axolotls used the same unique heavy chain and a limited number of light chains for their anti-2,4-dinitrophenyl antibody responses, Eur. J. Immunol, vol.17, 1987.

J. Charlemagne, Thymus independent anti-horse erythrocyte antibody response and suppressor T cells in the Mexican axolotl (Amphibia, Urodela, ambystoma mexicanum), Immunology, vol.36, pp.643-648, 1979.

J. Charlemagne and A. Tournefier, Humoral response to Salmonella typhimurium antigens in normal and thymectomized urodele amphibian Pleurodeles waltli Michah, Eur. J. Immunol, vol.7, 1977.

Y. C. Ching and R. J. Wedgwood, Immunologic responses in the axolotl, Siredon mexicanum, J. Immunol. Balt. Md, vol.1950, issue.99, pp.191-200, 1967.

C. Chothia and A. M. Lesk, Canonical structures for the hypervariable regions of immunoglobulins, J. Mol. Biol, vol.196, pp.901-917, 1987.

C. Chothia, A. M. Lesk, A. Tramontano, M. Levitt, S. J. Smith-gill et al., Conformations of immunoglobulin hypervariable regions, Nature, vol.342, 1989.

G. P. Cook and I. M. Tomlinson, The human immunoglobulin VH repertoire, Immunol. Today, vol.16, pp.80166-80175, 1995.

P. Dard, S. Huck, J. P. Frippiat, G. Lefranc, A. Langaney et al., The IGHG3 gene shows a structural polymorphism characterized by different hinge lengths: sequence of a new 2-exon hinge gene, Hum. Genet, vol.99, pp.138-141, 1997.

A. Desmyter, T. R. Transue, M. A. Ghahroudi, M. H. Thi, F. Poortmans et al., Analysis of the frequency and pattern of somatic mutations within nonproductively rearranged human variable heavy chain genes, J. Immunol. Balt. Md, vol.3, issue.158, pp.2779-2789, 1996.

J. S. Fellah and J. Charlemagne, Characterization of an IgY-like low molecular weight immunoglobulin class in the Mexican axolotl, Mol. Immunol, vol.25, 1988.

J. S. Fellah, S. Iscaki, J. P. Vaerman, J. Charlemagne, J. S. Fellah et al., Transient developmental expression of IgY and secretory component like protein in the gut of the axolotl (Ambystoma mexicanum), Immunogenetics, vol.39, 1992.

J. S. Fellah, F. Kerfourn, and J. Charlemagne, Evolution of T cell receptor genes. Extensive diversity of V beta families in the Mexican axolotl, J. Immunol. Balt. Md, vol.1950, issue.153, pp.4539-4545, 1994.

C. Frippiat, P. Kremarik, A. Ropars, C. Dournon, and J. P. Frippiat, The recombination-activating gene 1 of Pleurodeles waltl (urodele amphibian) is transcribed in lymphoid tissues and in the central nervous system, Immunogenetics, vol.52, pp.264-275, 2001.

M. Gellert, V(D)J recombination: RAG proteins, repair factors, and regulation, Annu. Rev. Biochem, vol.71, 2002.

R. Golub, S. Andr-e, A. Hassanin, P. Affaticati, M. Larijani et al., Early expression of two TdT isoforms in the hematopoietic system of the Mexican axolotl. Implications for the evolutionary origin of the N-nucleotide addition, Immunogenetics, vol.56, 2004.

R. Golub, J. Charlemagne, R. Golub, J. S. Fellah, J. ;. Charlemagne et al., Somatic mutation of immunoglobulin lambda chains: a segment of the major intron hypermutates as much as the complementarity-determining regions, 402e409. Gonz alez-Fern andez, vol.1950, 1994.

N. Gu-eguinou, C. Huin-schohn, N. Ouzren-zarhloul, S. Ghislin, and J. Frippiat, Molecular cloning and expression analysis of Pleurodeles waltl complement component C3 under normal physiological conditions and environmental stresses, Dev. Comp. Immunol, vol.46, 2014.

R. N. Haire, C. T. Amemiya, D. Suzuki, and G. W. Litman, Eleven distinct VH gene families and additional patterns of sequence variation suggest a high degree of immunoglobulin gene complexity in a lower vertebrate, Xenopus laevis, J. Exp. Med, vol.171, pp.1721-1737, 1990.

K. R. Hinds-frey, H. Nishikata, R. T. Litman, and G. W. Litman, Somatic variation precedes extensive diversification of germline sequences and combinatorial joining in the evolution of immunoglobulin heavy chain diversity, J. Exp. Med, vol.178, pp.815-824, 1993.

S. Hoegg, M. Vences, H. Brinkmann, and A. Meyer, Phylogeny and comparative substitution rates of frogs inferred from sequences of three nuclear genes, Mol. Biol. Evol, vol.21, 2004.

M. Houdayer and M. Fougereau, Phylogeny of immunoglobulins: immune reaction of the axolotl, Ambystoma mexicanum. Kinetics of the immune response and characterization of antibodies, Ann. Inst. Pasteur, vol.123, pp.3-28, 1972.

E. Hsu, J. Schwager, F. W. Alt, O. Ignatovich, I. M. Tomlinson et al., The creation of diversity in the human immunoglobulin V(lambda) repertoire, Proc. Natl. Acad. Sci. U. S. A, vol.86, 1989.

I. I. Ivanov, R. L. Schelonka, Y. Zhuang, G. L. Gartland, M. Zemlin et al., Development of the expressed Ig CDR-H3 repertoire is marked by focusing of constraints in length, amino acid use, and charge that are first established in early B cell progenitors, J. Immunol. Balt. Md, vol.1950, issue.174, pp.7773-7780, 2005.

S. L. Kaattari, H. L. Zhang, I. W. Khor, I. M. Kaattari, and D. A. Shapiro, Affinity maturation in trout: clonal dominance of high affinity antibodies late in the immune response, Dev. Comp. Immunol, vol.26, 2002.

J. Kaufman, H. V?-olk, and H. J. Wallny, A "minimal essential Mhc" and an "unrecognized Mhc": two extremes in selection for polymorphism, Immunol. Rev, vol.143, pp.63-88, 1995.

A. Lee and E. Hsu, Isolation and characterization of the Xenopus terminal deoxynucleotidyl transferase, J. Immunol. Balt. Md, vol.1950, issue.152, pp.4500-4507, 1994.

T. Logtenberg, Properties of polyreactive natural antibodies to self and foreign antigens, J. Clin. Immunol, vol.10, pp.137-140, 1990.

S. Marr, H. Morales, A. Bottaro, M. Cooper, M. Flajnik et al., Localization and differential expression of activation-induced cytidine deaminase in the amphibian Xenopus upon antigen stimulation and during early development, J. Immunol. Balt. Md, vol.1950, issue.179, pp.6783-6789, 2007.

M. Matsumoto, S. F. Lo, C. J. Carruthers, J. Min, S. Mariathasan et al., Affinity maturation without germinal centres in lymphotoxin-alpha-deficient mice, Nature, vol.382, pp.462-466, 1996.

R. Mussmann, L. Du-pasquier, and E. Hsu, Is Xenopus IgX an analog of IgA?, Eur. J. Immunol, vol.26, 1996.

W. Ndifon, H. Gal, E. Shifrut, R. Aharoni, N. Yissachar et al., Chromatin conformation governs T-cell receptor Jb gene segment usage, Proc. Natl. Acad. Sci. U. S. A, vol.109, 2012.

V. K. Nguyen, R. Hamers, L. Wyns, and S. Muyldermans, Camel heavy-chain antibodies: diverse germline V(H)H and specific mechanisms enlarge the antigen-binding repertoire, EMBO J, vol.19, 2000.

M. Nielsen, C. Lundegaard, O. Lund, and T. N. Petersen, CPHmodels-3.0eremote homology modeling using structure-guided sequence profiles, Nucleic Acids Res, vol.38, 2010.

Y. Ohta and M. Flajnik, IgD, like IgM, is a primordial immunoglobulin class perpetuated in most jawed vertebrates, Proc. Natl. Acad. Sci. U. S. A, vol.103, 2006.

E. A. Padlan, H. M. Patel, and E. Hsu, Abbreviated junctional sequences impoverish antibody diversity in urodele amphibians, J. Immunol. Balt. Md, vol.31, issue.159, pp.3391-3399, 1994.

J. Pei, B. Kim, and N. V. Grishin, PROMALS3D: a tool for multiple protein sequence and structure alignments, Nucleic Acids Res, vol.36, 2008.

C. Prado, M. Rodríguez, I. Cortegano, C. Ruiz, M. Alía et al., Postnatal and adult immunoglobulin repertoires of innate-like CD19(þ) CD45R(lo) B Cells, J Innate Immun, vol.6, 2014.

A. Ropars, A. Bautz, and C. Dournon, Sequencing and expression of the CD3 gamma/delta mRNA in Pleurodeles waltl (urodele amphibian), Immunogenetics, vol.54, 2002.

K. H. Roux, A. S. Greenberg, L. Greene, L. Strelets, D. Avila et al., Structural analysis of the nurse shark (new) antigen receptor (NAR): molecular convergence of NAR and unusual mammalian immunoglobulins, Proc. Natl. Acad. Sci. U. S. A, vol.95, 1998.

J. F. Ruiz, D. Lucas, E. García-palomero, A. I. Saez, M. A. Gonz-alez et al., Overexpression of human DNA polymerase mu (Pol mu) in a Burkitt's lymphoma cell line affects the somatic hypermutation rate, Nucleic Acids Res, vol.32, 2004.

F. Salvadori and A. Tournefier, Activation by mitogens and superantigens of axolotl lymphocytes: functional characterization and ontogenic study, Immunology, vol.88, pp.586-592, 1996.

B. Sammut, V. Laurens, and A. Tournefier, Isolation of Mhc class I cDNAs from the axolotl Ambystoma mexicanum, Immunogenetics, vol.45, pp.285-294, 1997.

B. Schaerlinger, M. Bascove, and J. Frippiat, A new isotype of immunoglobulin heavy chain in the urodele amphibian Pleurodeles waltl predominantly expressed in larvae, Mol. Immunol, vol.45, 2008.

B. Schaerlinger and J. Frippiat, IgX antibodies in the urodele amphibian Ambystoma mexicanum, Dev. Comp. Immunol, vol.32, 2008.

R. L. Schelonka, E. Szymanska, A. M. Vale, Y. Zhuang, G. L. Gartland et al., DH and JH usage in murine fetal liver mirrors that of human fetal liver, Immunogenetics, vol.62, 2010.

V. Schenten, N. Gu-eguinou, S. Baatout, and J. Frippiat, Modulation of Pleurodeles waltl DNA polymerase mu expression by extreme conditions encountered during spaceflight, PloS One, vol.8, 2013.
URL : https://hal.archives-ouvertes.fr/hal-01480625

J. Schwager, N. Bürckert, M. Courtet, and L. Du-pasquier, The ontogeny of diversification at the immunoglobulin heavy chain locus in Xenopus, EMBO J, vol.10, pp.2461-2470, 1991.

J. Schwager, N. Bürckert, M. Courtet, and L. Du-pasquier, Genetic basis of the antibody repertoire in Xenopus: analysis of the Vh diversity, EMBO J, vol.8, 1989.

R. Subrahmanyam, H. Du, I. Ivanova, T. Chakraborty, Y. Ji et al., Localized epigenetic changes induced by DH recombination restricts recombinase to DJH junctions, Nat. Immunol, vol.13, 2012.

A. Tournefier, Incomplete antibodies and immunoglobulin characterization in adult urodeles, Pleurodeles waltl Michah. and Triturus alpestris Laur. Immunology, vol.29, pp.209-217, 1975.

A. Tournefier and J. Charlemagne, Antibodies against Salmonella and SRBC in urodele amphibians: synthesis and characterization, Adv. Exp. Med. Biol, vol.64, pp.161-171, 1975.

A. Tournefier, V. Laurens, C. Chapusot, P. Ducoroy, M. R. Padros et al., Structure of MHC class I and class II cDNAs and possible immunodeficiency linked to class II expression in the Mexican axolotl, Immunol. Rev, vol.166, pp.259-277, 1998.

A. Tramontano, C. Chothia, and A. M. Lesk, Framework residue 71 is a major determinant of the position and conformation of the second hypervariable region in the VH domains of immunoglobulins, J. Mol. Biol, vol.215, 1990.

H. V?-olk, J. Charlemagne, A. Tournefier, S. Ferrone, R. Jost et al., Wide tissue distribution of axolotl class II molecules occurs independently of thyroxin, Immunogenetics, vol.47, pp.339-349, 1998.

G. W. Warr, K. E. Magor, and D. A. Higgins, IgY: clues to the origins of modern antibodies, Immunol. Today, vol.16, issue.95, pp.80008-80013, 1995.

J. A. Weinstein, N. Jiang, R. A. White, D. S. Fisher, and S. R. Quake, Highthroughput sequencing of the zebrafish antibody repertoire, Science, vol.324, 2009.

M. Wilson, E. Hsu, A. Marcuz, M. Courtet, L. Du-pasquier et al., What limits affinity maturation of antibodies in Xenopusethe rate of somatic mutation or the ability to select mutants?, EMBO J, vol.11, pp.4337-4347, 1992.

F. Yang, G. C. Waldbieser, and C. J. Lobb, The nucleotide targets of somatic mutation and the role of selection in immunoglobulin heavy chains of a teleost fish, J. Immunol. Balt. Md, vol.1950, issue.176, pp.1655-1667, 2006.

M. Yasuike, J. De-boer, K. R. Von-schalburg, G. A. Cooper, L. Mckinnel et al., Evolution of duplicated IgH loci in Atlantic salmon, Salmo salar, BMC Genomics, vol.11, 2010.

J. Y-elamos, N. Klix, B. Goyenechea, F. Lozano, Y. L. Chui et al., Targeting of non-Ig sequences in place of the V segment by somatic hypermutation, Nature, vol.376, 1995.

Y. Zhang, I-TASSER server for protein 3D structure prediction, BMC Bioinforma, vol.9, 2008.

Y. Zhang, I. Salinas, J. Li, D. Parra, S. Bjork et al., IgT, a primitive immunoglobulin class specialized in mucosal immunity, Nat. Immunol, vol.11, 2010.

Y. Zhao, Q. Pan-hammarstr?-om, S. Yu, N. Wertz, X. Zhang et al., Identification of IgF, a hinge-region-containing Ig class, and IgD in Xenopus tropicalis, Proc. Natl. Acad. Sci. U. S. A, vol.103, 2006.