Comparative genomic studies are revealing that, in sharp contrast with the strong stability found in birds and mammals, sex determination mechanisms are surprisingly labile in cold-blooded vertebrates, with frequent transitions between different pairs of sex chromosomes. It was recently suggested that, in context of this high turnover, some chromosome pairs might be more likely than others to be co-opted as sex chromosomes. Empirical support, however, is still very limited. Here we show that sex-linked markers from three highly divergent groups of anurans map to Xenopus tropicalis scaffold 1, a large part of which is homologous to the avian sex chromosome. Accordingly, the bird sex determination gene DMRT1, known to play a key role in sex differentiation across many animal lineages, is sex linked in all three groups. Our data provide strong support for the idea that some chromosome pairs are more likely than others to be co-opted as sex chromosomes because they harbor key genes from the sex determination pathway. K E Y W O R D S :Amphibian, Bufo siculus, convergent evolution, conserved synteny, DMRT1, Hyla arborea, Rana temporaria, sex chromosome turnover.

Contrasting with birds and mammals, most ectothermic vertebrates present homomorphic sex chromosomes, which might be due either to a high turnover rate or to occasional X-Y recombination. We tested these two hypotheses in a group of Palearctic green toads that diverged some 3.3 million years ago. Using sibship analyses of sex-linked markers, we show that all four species investigated share the same pair of sex chromosomes and a pattern of male heterogamety with drastically reduced X-Y recombination in males. Phylogenetic analyses of sex-linked sequences show that X and Y alleles cluster by species, not by gametolog. We conclude that X-Y homomorphy and fine-scale sequence similarity in these species do not stem from recent sex-chromosome turnovers, but from occasional X-Y recombination.

In sharp contrast with birds and mammals, the sex chromosomes of ectothermic vertebrates are often undifferentiated, for reasons that remain debated. A linkage map was recently published for Rana temporaria (Linnaeus, 1758) from Fennoscandia (Eastern European lineage), with a proposed sex-determining role for linkage group 2 (LG2). We analysed linkage patterns in lowland and highland populations from Switzerland (Western European lineage), with special focus on LG2. Sibship analyses showed large differences from the Fennoscandian map in terms of recombination rates and loci order, pointing to large-scale inversions or translocations. All linkage groups displayed extreme heterochiasmy (total map length was 12.2 cM in males, versus 869.8 cM in females). Sex determination was polymorphic within populations: a majority of families (with equal sex ratios) showed a strong correlation between offspring phenotypic sex and LG2 paternal haplotypes, whereas other families (some of which with female-biased sex ratios) did not show any correlation. The factors determining sex in the latter could not be identified. This coexistence of several sex-determination systems should induce frequent recombination of X and Y haplotypes, even in the absence of male recombination. Accordingly, we found no sex differences in allelic frequencies on LG2 markers among wild-caught male and female adults, except in one high-altitude population, where nonrecombinant Y haplotypes suggest sex to be entirely determined by LG2. Multifactorial sex determination certainly contributes to the lack of sex-chromosome differentiation in amphibians.

We investigated the ability of European sea bass (Dicentrarchus labrax) to respond simultaneously to the metabolic demands of specific dynamic action (SDA) and aerobic exercise and how this was influenced by moderate hypoxia (50% air saturation). At 3 h after feeding in normoxia at 20 degrees C, SDA raised the instantaneous oxygen uptake (Mo(2)) of sea bass by 47% +/- 18% (mean +/- SEM, N = 7) above their standard metabolic rate (SMR) when fasted. This metabolic load was sustained throughout an incremental exercise protocol until fatigue, with a 14% +/- 3% increase in their maximum aerobic metabolic rate (MMR) relative to their fasted rate. Their incremental critical swimming speed (U(crit)) did not differ between fasted and fed states. Thus, in normoxia, the bass were able to meet the combined oxygen demands of SDA and aerobic exercise. In hypoxia, the sea bass suffered a significant decline in MMR and U(crit) relative to their normoxic performance. The SDA response was similar to normoxia (84% +/- 24% above fasted SMR at 3 h after feeding), but although this load was sustained at low swimming speeds, it gradually disappeared as swimming speed increased. As a result, the hypoxic sea bass exhibited no difference in their fasted versus fed MMR. Hypoxic U(crit) did not, however, differ between fasted and fed states, indicating that the sea bass deferred their SDA to maintain exercise performance. The results demonstrate that, in normoxia, the sea bass possesses excess cardiorespiratory capacity beyond that required for maximal aerobic exercise. The excess capacity is lost when oxygen availability is limited in hypoxia, and, under these conditions, the sea bass prioritize exercise performance. Thus, environmental conditions (oxygen availability) had a significant effect on patterns of oxygen allocation in sea bass and revealed intrinsic prioritization among conflicting metabolic demands.