The hormone auxin plays a crucial role in plant morphogenesis. In the shoot apical meristem, the PIN-FORMED1 (PIN1) efflux carrier concentrates auxin into local maxima in the epidermis, which position incipient leaf or floral primordia. From these maxima, PIN1 transports auxin into internal tissues along emergent paths that pattern leaf and stem vasculature. In Arabidopsis thaliana, these functions are attributed to a single PIN1 protein. Using phylogenetic and gene synteny analysis we identified an angiosperm PIN clade sister to PIN1, here termed Sister-of-PIN1 (SoPIN1), which is present in all sampled angiosperms except for Brassicaceae, including Arabidopsis. Additionally, we identified a conserved duplication of PIN1 in the grasses: PIN1a and PIN1b. In Brachypodium distachyon, SoPIN1 is highly expressed in the epidermis and is consistently polarized toward regions of high expression of the DR5 auxin-signaling reporter, which suggests that SoPIN1 functions in the localization of new primordia. In contrast, PIN1a and PIN1b are highly expressed in internal tissues, suggesting a role in vascular patterning. PIN1b is expressed in broad regions spanning the space between new primordia and previously formed vasculature, suggesting a role in connecting new organs to auxin sinks in the older tissues. Within these regions, PIN1a forms narrow canals that likely pattern future veins. Using a computer model, we reproduced the observed spatio-temporal expression and localization patterns of these proteins by assuming that SoPIN1 is polarized up the auxin gradient, and PIN1a and PIN1b are polarized to different degrees with the auxin flux. Our results suggest that examination and modeling of PIN dynamics in plants outside of Brassicaceae will offer insights into auxin-driven patterning obscured by the loss of the SoPIN1 clade in Brassicaceae.

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HighlightSurvival of A. tequilana, under arid conditions depends on a failsafe mechanism for asexual reproduction based on changes in auxin mobility controlled by two functionally distinct PIN1-related proteins.

Auxin plays a vital role in plant organ development, influencing organ initiation and patterning across all axes. The diversity in auxin patterning results from changes in the activities and expression of auxin signaling components, including the AUX/IAA repressors. Higher land plants have multigene AUX/IAA families, which leads to functional redundancy and a lack of phenotype in loss of function mutants. Instead, dominant mutations, which prevent AUX/IAA degradation in response to auxin, have highlighted the importance of these proteins in organ development. Here we report a new dominant AUX/IAA mutant in maize, Hoja loca1 (Oja). Oja has a mutation in the degron motif of ZmIAA28 and affects aerial organ initiation and medio-lateral patterning in the leaf. These phenotypes contrast with other maize AUX/IAA mutants that affect the root or inflorescence only. Oja illustrates the role of auxin signaling in the tight coordination of phytomer unit development and provides evidence of species-specific sub-functionalization of the AUX/IAAs.One Line SummaryThe maize AUX/IAA ZmIAA38 is involved in phytomer coordination, aerial organ initiation and mediolateral patterning, and illustrates species specific sub-functionalization of the AUX/IAAs.