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The TIR1/AFB auxin co-receptors mediate diverse responses to the plant hormone auxin. The Arabidopsis genome encodes six TIR1/AFB proteins representing three of the four clades that were established prior to angiosperm radiation. To determine the role of these proteins in plant development we performed an extensive genetic analysis involving the generation and characterization of all possible multiply-mutant lines. We find that loss of all six TIR1/AFB proteins results in early embryo defects and eventually seed abortion, and yet a single wild-type allele of TIR1 or AFB2 is sufficient to support growth throughout development. Our analysis reveals extensive functional overlap between even the most distantly related TIR1/AFB genes except for AFB1. Surprisingly, AFB1 has a specialized function in rapid auxin-dependent inhibition of root growth and early phase of root gravitropism. This activity may be related to a difference in subcellular localization compared to the other members of the family.

Auxin response factors (ARFs) are a family of transcription factors that are responsible for regulating gene expression in response to changes in auxin level. The analysis of ARF sequence and activity indicates that there are 2 major groups: activators and repressors. One clade of ARFs, clade-D, is sister to clade-A activating ARFs, but are unique in that they lack a DNA-binding domain. Clade-D ARFs are present in lycophytes and bryophytes but absent in other plant lineages. The transcriptional activity of clade-D ARFs, as well as how they regulate gene expression, is not well understood. Here, we report that clade-D ARFs are transcriptional activators in the model bryophyte Physcomitrium patens and have a major role in the development of this species. Δarfd dub protonemata exhibit a delay in filament branching, as well as a delay in the chloronema to caulonema transition. Additionally, leafy gametophore development in Δarfd dub lines lags behind wild type. We present evidence that ARFd1 interacts with activating ARFs via their PB1 domains, but not with repressing ARFs. Based on these results, we propose a model in which clade-D ARFs enhance gene expression by interacting with DNA bound clade-A ARFs. Further, we show that ARFd1 must form oligomers for full activity.

The plant hormone auxin regulates a wide variety of transcriptional responses depending on the cell type, environment and species. How this diversity is achieved may be related to the specific complement of auxin-signalling components in each cell. The levels of activators (class-A AUXIN RESPONSE FACTORS) and repressors (class-B ARFs) are particularly important. Tight regulation of ARF protein levels is probably key in determining this balance. Through comparative analysis of novel, dominant mutants in maize and the moss Physcomitrium patens , we have discovered a ~500-million-year-old mechanism of class-B ARF protein-level regulation mediated by proteasome degradation, important in determining cell fate decisions across land plants. Thus, our results add a key piece to the puzzle of how auxin regulates plant development. Comparative analysis of maize and moss reveals a 500-million-year-old piece of the auxin-signalling puzzle that controls the protein level of ARFs, key transcriptional regulators.

A major challenge in plant biology is to understand how the plant hormone auxin regulates diverse transcriptional responses throughout development, in different environments, and in different species. The answer may lie in the specific complement of auxin signaling components in each cell. The balance between activators (class-A AUXIN RESPONSE FACTORS) and repressors (class-B ARFs) is particularly important. It is unclear how this balance is achieved. Through comparative analysis of novel, dominant mutants in maize and the moss , we have discovered a ∼500-million-year-old mechanism of class-B ARF protein level regulation, important in determining cell fate decisions across land plants. Thus, our results add a key piece to the puzzle of how auxin regulates plant development.

Auxin Response Factors (ARFs) are a family of transcription factors that are responsible for regulating gene expression in response to changes in auxin level. The analysis of ARF sequence and activity indicates that there are two major groups- activators and repressors. One clade of ARFs, clade-D, is sister to clade-A activating ARFs, but are unique in that they lack a DNA binding domain. Clade-D ARFs are present in lycophytes and bryophytes but absent in pteridophytes and spermatophytes. The transcriptional activity of clade-D ARFs, as well as how they regulate gene expression, is not well understood. Here, we report that clade-D ARFs are transcriptional activators in the model bryophyte P. patens and have a major role in the development of this species. Δarfd1,d2 protonemata exhibit a delay in filament branching, as well as a delay in a key cell differentiation event. Additionally, leafy gametophore development in Δarfd1,d2 lines lag behind wild-type. We present evidence that ARFd1 interacts with activating, but not repressing, ARFs. An ARFd1 hypomorph, arfd1T653L, cannot multimerize. Therefore, we propose a model by which clade-D ARFs enhance gene expression by oligomerizing to DNA-bound archetypal ARFs. Competing Interest Statement The authors have declared no competing interest.

The TIR1/AFB auxin co-receptors mediate diverse responses to the plant hormone auxin. The Arabidopsis genome encodes six TIR1/AFB proteins representing three of the four clades that were established prior to angiosperm radiation. To determine the role of these proteins in plant development we performed an extensive genetic analysis involving the generation and characterization of all possible multiply mutant lines. We find that loss of all six TIR1/AFB proteins results in defects in embryogenesis as early as the 8-cell stage, and possibly earlier. Mutant embryos progress but exhibit frequent cell division errors followed by proliferation of the suspensor, and eventually seed abortion. Despite this dramatic phenotype, a single wild-type allele of TIR1 or AFB2 is sufficient to support growth throughout plant development. Further, gametophytic expression of the TIR1/AFB genes is not essential for development of the male or female gametophyte. Our analysis reveals extensive functional overlap between even the most distantly related TIR1/AFB genes except for AFB1. Surprisingly, the AFB1 protein has a specialized function in rapid auxin-dependent inhibition of root growth and early phase of root gravitropism. This activity may be related to a difference in subcellular localization compared to the other members of the family. Footnotes * New data has been added that addresses the role of the TIR1/AFB proteins in rapid auxin-mediated inhibition of root elongation. Surprisingly the AFB1 protein has a particularly important role in this response. In addition, we show that AFB1 contributes during early stages of root gravitropism.