Page 212 - The Toxicology of Fishes
P. 212

192                                                        The Toxicology of Fishes


                                                                0.1
                                                         ZebrafishGSTA
                                                          FHMinnow
                                                          PufferGSTA
                                                           Medaka
                                                                                     Rho class
                                                         plaiceGSTA1
                                                         SeaBream
                                                         LMBass
                                                         PlaiceGSTA
                                                           ZebrafishOmega
                                                                                     Omega class
                                                           PufferOmega
                                                           ZebrafishAlpha            Alpha class
                                                           CatfishPi
                                                         SocksalmonPi
                                                                                     Pi class
                                                          TroutPi
                                                         ZebrafishPi
                                                           ZebrafishMU2
                                                          ZebrafishMU1               Mu class
                                                           MedakaMU
                                                           ZebrafishTheta
                                                                                     Theta class
                                                           PufferTheta
                                                           WFlounderAJ605273
                                                            PufferCA844840
                                                                                     Microsomal
                                                           MedakaAV669487
                                                           Plaicetrypsinogen

                       FIGURE 4.12 Phylogenetic relationships of cloned fish GSTs; derived from deduced amino acid sequences using Clustal
                       W (zeta and mitochondrial kappa isoforms omitted).

                       major isoform in flatfish, mullet, and bass is now designated as Rho-class (Konishi et al., 2005). From
                       nucleotide and protein sequence data, genes for all nine families have been identified in fish to date. A
                       phylogenetic analysis of the sequenced fish genes is shown in Figure 4.12. Table 4.12 shows for each
                       species the number of GST isoforms assigned to each family.
                        From purification studies, pi-class GSTs have been identified as the major hepatic and intestinal GSTs
                       in the livers of salmonids, brown bullhead, channel catfish, and lamprey. These piscine pi-class enzymes
                       display high activities with BaP epoxides and diol epoxides. The most highly characterized piscine GSTs
                       are from the plaice, a pleuronectid flatfish, where purification and cloning studies identified a multigene
                       cluster of enzymes originally designated as GSTA. These are most closely related to the mammalian,
                       insect, and plant theta-class enzymes and have now been designated as a separate class, Rho. They
                       appear to be unique to fish. Their genetic organization appears to be the same in plaice, flounder, sea
                       bream, and largemouth bass. The primary role of these theta-like enzymes is in oxidative defense. GST-A
                       is most active toward hydroxynonenals, and GSTA1 (or GSTR2) exhibits a high peroxidase activity with
                       organic peroxides. They are the major isoforms in most tissues of flatfish. Homologs have been identified
                       by cDNA sequencing, purification, or immunochemical cross-reactivity in a number of other fish species,
   207   208   209   210   211   212   213   214   215   216   217