title: Gamete types, sex determination and stable equilibria of all-hybrid populations of diploid and triploid edible frogs (Pelophylax esculentus).
abstract: BACKGROUND
  Triploid individuals often play a key role in speciation by hybridization. An understanding of the gamete types (ploidy and genomic content) and stability of hybrid populations with triploid individuals is therefore of importance for exploring the role of hybridization in evolution. The all-hybrid populations of the edible frog, Pelophylax esculentus, are unique in their composition and genetic dynamics: Diploid (genotype LR) and triploid (LLR and LRR) hybrids depend on each other's different gamete contributions for successful reproduction and maintenance of the populations, as the parental genotypes P. lessonae (LL) and P. ridibundus (RR) are absent among adults. This study provides data and interpretations on gamete types and sex determination that are essential for understanding the function, evolutionary potential and threats of this intriguing system.
  
  
  RESULTS
  Dissection of metamorphs from a crossing experiment confirmed that sex determination is an XX-XY system with the Y confined to the L genome. From microsatellite analysis of parents and offspring from the crossings, gamete frequencies could be deduced: Triploids of both sexes mostly made haploid gametes with the genome they had in double dose, however LLR females also made approximately 10% LL gametes by automixis. LR frogs showed much variation in their gamete production. In LRR-rich populations, their LR sperm production was sufficiently high (22%) to explain the observed proportion of LRR males, the formation of which has not previously been understood. A model was constructed to calculate equilibrium genotype proportions for different population types on the basis of the gamete proportions found. These equilibria agreed well with empirical literature data.
  
  
  CONCLUSIONS
  If population differentiation with respect to genotype proportions is really driven by gamete patterns, as strongly suggested by the present study, all-hybrid populations constitute not one, but several intrinsically different breeding systems. Tetraploidization could occur if the survival or fertility of both males and females increased. Whether introduction of hybrid or parental species individuals would threaten the all-hybrid populations cannot be predicted without further knowledge on the mechanisms behind non-hybrid inviability, but at least R genomes with Y factor are predicted to be invasive, if introduced, and could bring the populations to collapse.
authors.affiliation: Ecology, Zoological Institute, University of Zurich, Winterthurerstrasse 190, CH-8057 Zurich, Switzerland. ditte.christiansen@zool.uzh.ch
authors.forename: Ditte G
authors.initials: DG
authors.lastname: Christiansen
mesh_terms: D000818:Animals; D003433:Crosses, Genetic; D004171:Diploidy; D005260:Female; D005718:Gametogenesis; D005838:Genotype; D005854:Germ Cells / Q000166:cytology*; D006824:Hybridization, Genetic; D008297:Male; D018895:Microsatellite Repeats; D008957:Models, Genetic; D011123:Polyploidy; D011898:Ranidae / Q000235:genetics*; D012098:Reproduction / Q000235:genetics; D019849:Sex Determination Processes