Controlled auxin biosynthesis and transport are important for developmental decisions in the early diverging land plant Physcomitrella patens

Sammanfattning: The aim of this thesis was to increase our understanding of the importance of the plant hormone auxin in land plant evolution. The role of a number of auxin regulatory network components in developmental decisions during the haploid phase of the life cycle of the model moss Physcomitrella patens, which belongs to the early diverging bryophyte group of land plants, was investigated. We show that the role of SHI/STY transcriptional activators in the regulation of auxin biosynthesis rates in seed plants is conserved in moss. The amount and rates of auxin synthesis are reduced in PpSHI knockout mutants, defects in these mutants can be mimicked by reducing active auxin in the PpSHI expression domain, and several moss homologues of YUC and TAA1/TAR genes, which encode the main auxin biosynthesis enzymes in seed plants, were activated in moss when PpSHI2 was transiently expressed. Our data reveal that these moss YUC and TAA1/TAR genes can induce auxin biosynthesis, and that their expression domains largely overlap with that of the two PpSHIs in certain stages of the moss life cycle. PpSHI2 also directly or indirectly activates homologues of genes involved in polar auxin transport (PAT) in seed plants, PpPINA, PpPINB and PpLAXB. We show that the auxin efflux function of PIN proteins have been conserved during the evolution of land plants, and that they, together with the PpSHI auxin biosynthesis regulators, are important for determining developmentally regulated decisions, such as the switch between two cell types of the filamentous stage in moss. In addition, PIN-mediated distribution of auxin synthesised in the leaves plays an important role in determining a developmentally regulated wave of cell expansion. PpSHI regulated local auxin biosynthesis is crucial for several steps in reproductive organ development, such as egg cell maturation, clearance of a canal to the egg cavity, as well as apical opening of both male and female reproductive organs allowing sperm release and entrance into the egg cavity. If polar auxin transport plays a role during these stages remains to be studied. This demonstrates that regulated distribution of auxin was operational in cell-fate decisions already in the early land plants, and that auxin has played a key role in the evolution of complex land plant structures.

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