Bidirectional hybridization between Ulva prolifera and U. linza (Ulvophyceae, Chlorophyta): Evidence for compatibility and paternal chloroplast inheritance

Ulva prolifera and U. linza are closely related species, with abundant adult thalli and reproductive cells co-occurring extensively in time and space during the Yellow Sea green tides. Elucidating their hybridization compatibility is crucial for species delimitation, assessing interspecific gene flow, and evaluating the ecological impacts of green tides. Previous studies suggested asymmetric gamete compatibility (only U. prolifera mt+ x U. linza mt-), but lacked sex-linked markers to reliably identify hybrid diploids and their reproductive modes, and did not examine chloroplast inheritance. Here, we performed bidirectional crosses using sexual strains of different geographic origins from both parents, with sex-linked markers to quantify progeny genotypes, examine fertility and reproduction pathway of F1 hybrids, and trace chloroplast inheritance using the species-specific petB marker. Our results showed that: (1) F1 hybrids were obtained in both cross directions, with significantly higher frequency in the direct cross (U. linza mt+ x U. prolifera mt-), indicating no complete reproductive isolation in either direction; the biased compatibility likely reflected genetic background differences among the limited strain combinations in a single study. (2) A considerable number of germinated progeny arose from parthenogenesis of parental gametes. (3) F1 hybrids from both crosses could undergo meiosis to form gametes and develop into gametophytes; additionally, F1 from the reciprocal cross produced diploid spores for asexual reproduction, suggesting meiotic disturbance. (4) Chloroplasts were maternally inherited in selfing of U. prolifera parent, but in all F1 hybrids they were paternally inherited, indicating a potential reversal of the inheritance pattern due to interspecific hybridization. These findings provided new insights into the potential for genetic exchange between U. prolifera and U. linza.