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Phylogenetic Analysis and Identification of Pseudogenes Reveal a Progressive Loss of Zona Pellucida Genes During Evolution of Vertebrates¹

Physiologie de la Reproduction et des Comportements,³ INRA-CNRS-Université de Tours-Haras Nationaux, IFR 135, 37380 Nouzilly, France Département de Biologie Structurale,⁴ IMPMC UMR 7590, Universités Pierre et Marie Curie-Paris 6 et Denis Diderot-Paris 7, CNRS, Campus Boucicaut, 75015 Paris, France

ABSTRACT

Vertebrate eggs are surrounded by an extracellular matrix with similar functions and conserved individual components: the zona pellucida (ZP) glycoproteins. In mammals, chickens, frogs, and some fish species, we established an updated list of the ZP genes, studied the relationships within the ZP gene family using phylogenetic analysis, and identified ZP pseudogenes. Our study confirmed the classification of ZP genes in six subfamilies: ZPA/ZP2, ZPB/ZP4, ZPC/ZP3, ZP1, ZPAX, and ZPD. The identification of a Zpb pseudogene in the mouse genome, Zp1 pseudogenes in the dog and bovine genomes, and Zpax pseudogenes in the human, chimpanzee, macaque, and bovine genomes showed that the evolution of ZP genes mainly occurs by death of genes. Our study revealed that the extracellular matrix surrounding vertebrate eggs contains three to at least six ZP glycoproteins. Mammals can be classified in three categories. In the mouse, the ZP is composed of three ZP proteins (ZPA/ZP2, ZPC/ZP3, and ZP1). In dog, cattle and, putatively, pig, cat, and rabbit, the zona is composed of three ZP proteins (ZPA/ZP2, ZPB/ZP4, and ZPC/ZP3). In human, chimpanzee, macaque, and rat, the ZP is composed of four ZP proteins (ZPA/ZP2, ZPB/ZP4, ZPC/ZP3, and ZP1). Our review provides new directions to investigate the molecular basis of sperm-egg recognition, a mechanism which is not yet elucidated.

evolution, fertilization, gamete biology, oocyte development, ovum, pseudogene, zona pellucida

¹Supported by grants from Les Haras Nationaux in France and La Région Centre in France to S.M.

Correspondence: ²FAX: 33 2 47 42 77 43; e-mail: goudet{at}tours.inra.fr

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