Save your TIRs - more to auxin than meets the eye.

Auxin has long been known as an important regulator of plant growth and development. Classical studies in auxin biology have uncovered a 'canonical' transcriptional auxin-signalling pathway involving the TRANSPORT INHIBITOR RESPONSE1/AUXIN SIGNALING F-BOX (TIR1/AFB) receptors. TIR1/AFB perception of auxin triggers the degradation of repressors and the derepression of auxin-responsive genes. Nevertheless, the canonical pathway cannot account for all aspects of auxin biology, such as physiological responses that are too rapid for transcriptional regulation. This Tansley insight will explore several 'non-canonical' pathways that have been described in recent years mediating fast auxin responses. We focus on the interplay between a nontranscriptional branch of TIR1/AFB signalling and a TRANSMEMBRANE KINASE1 (TMK1)-mediated pathway in root acid growth. Other developmental aspects involving the TMKs and their association with the controversial AUXIN-BINDING PROTEIN 1 (ABP1) will be discussed. Finally, we provide an updated overview of the ETTIN (ETT)-mediated pathway in contexts outside of gynoecium development.

Noncanonical Auxin Signaling.

Auxin influences all aspects of plant growth and development and exerts its function at scales ranging from the subcellular to the whole-organism level. A canonical mechanism for auxin signaling has been elucidated, which is based on derepression of downstream genes via ubiquitin-mediated degradation of transcriptional repressors. While the combinatorial nature of this canonical pathway provides great potential for specificity in the auxin response, alternative noncanonical signaling pathways required to mediate certain processes have been identified. One such pathway affects gene regulation in a manner that is reminiscent of mechanisms employed in animal hormone signaling, while another triggers transcriptional changes through auxin perception at the plasma membrane and the stabilization of transcriptional repressors. In some cases, the exact perception mechanisms and the nature of the receptors involved are yet to be revealed. In this review, we describe and discuss current knowledge on noncanonical auxin signaling and highlight unresolved questions surrounding auxin biology.