Strigolactones (SLs), or their derivatives, were recently demonstrated to act as endogenous shoot branching inhibitors, but their biosynthesis and mechanism of action are poorly understood. Here we show that the branching phenotype of mutants in the Arabidopsis P450 family member, MAX1, can be fully rescued by strigolactone addition, suggesting that MAX1 acts in SL synthesis. We demonstrate that SLs modulate polar auxin transport to control branching and that both the synthetic SL GR24 and endogenous SL synthesis significantly reduce the basipetal transport of a second branch-regulating hormone, auxin. Importantly, GR24 inhibits branching only in the presence of auxin in the main stem, and enhances competition between two branches on a common stem. Together, these results support two current hypotheses: that auxin moving down the main stem inhibits branch activity by preventing the establishment of auxin transport out of axillary branches; and that SLs act by dampening auxin transport, thus enhancing competition between branches.

Plant species have evolved different requirements for environmental/endogenous cues to induce flowering. Originally, these varying requirements were thought to reflect the action of different molecular mechanisms. Thinking changed when genetic and molecular analysis in Arabidopsis thaliana revealed that a network of environmental and endogenous signaling input pathways converge to regulate a common set of “floral pathway integrators.” Variation in the predominance of the different input pathways within a network can generate the diversity of requirements observed in different species. Many genes identified by flowering time mutants were found to encode general developmental and gene regulators, with their targets having a specific flowering function. Studies of natural variation in flowering were more successful at identifying genes acting as nodes in the network central to adaptation and domestication. Attention has now turned to mechanistic dissection of flowering time gene function and how that has changed during adaptation. This will inform breeding strategies for climate-proof crops and help define which genes act as critical flowering nodes in many other species.

Significance: Strigolactones are a new class of plant hormones regulating plant shoot and root architecture in response to the environment. Also present in root exudates, strigolactones stimulate the germination of parasitic plant seeds. This report describes a genomic polymorphism—associated with the Indica / Japonica subspecies divide in rice that has a major impact on the biosynthesis of strigolactones, plant tillering, and germination of the parasitic plant Striga hermonthica —consisting of the deletion of two strigolactone biosynthetic genes orthologous to Arabidopsis MAX1 . Both of these genes rescued the Arabidopsis max1-1 highly branched mutant phenotype and increased the strigolactone level when overexpressed in the Indica rice variety Bala. This finding is of great interest for plant physiologists, plant evolutionary biologists, and breeders.

Strigolactones (SLs) are carotenoid-derived phytohormones with diverse roles. They are secreted from roots as attractants for arbuscular mycorrhizal fungi and have a wide range of endogenous functions, such as regulation of root and shoot system architecture. To date, six genes associated with SL synthesis and signaling have been molecularly identified using the shoot-branching mutants more axillary growth (max) of Arabidopsis (Arabidopsis thaliana) and dwarf (d) of rice (Oryza sativa). Here, we present a phylogenetic analysis of the MAX/D genes to clarify the relationships of each gene with its wider family and to allow the correlation of events in the evolution of the genes with the evolution of SL function. Our analysis suggests that the notion of a distinct SL pathway is inappropriate. Instead, there may be a diversity of SL-like compounds, the response to which requires a D14/D14-like protein. This ancestral system could have been refined toward distinct ligand-specific pathways channeled through MAX2, the most downstream known component of SL signaling. MAX2 is tightly conserved among land plants and is more diverged from its nearest sister clade than any other SL-related gene, suggesting a pivotal role in the evolution of SL signaling. By contrast, the evidence suggests much greater flexibility upstream of MAX2. The MAX1 gene is a particularly strong candidate for contributing to diversification of inputs upstream of MAX2. Our functional analysis of the MAX1 family demonstrates the early origin of its catalytic function and both redundancy and functional diversification associated with its duplication in angiosperm lineages.