Auxin-independent effects of apical dominance induce temporal changes in phytohormones

The inhibition of shoot branching by the growing shoot tip of plants, termed apical dominance, was originally thought to be mediated by auxin. Recently the importance of the shoot tip sink strength during apical dominance has re-emerged with recent studies highlighting roles for sugars in promoting branching. This raises many unanswered questions on the relative roles of auxin and sugars in apical dominance. Here we show that auxin regulation of cytokinins, which promote branching, is significant only after an initial stage of branching we call bud release. During this early bud release stage, rapid cytokinin increases are associated with enhanced sugars. Auxin may also act through strigolactones which have been shown to suppress branching after decapitation, but here we show that strigolactones do not have a significant effect on initial bud outgrowth after decapitation. We report here that when sucrose or cytokinin is abundant, strigolactones are less inhibitory during the bud release stage compared to later stages and that strigolactone treatment rapidly inhibits cytokinin accumulation in pea axillary buds of intact plants. After initial bud release, we find an important role of gibberellin in promoting sustained bud growth downstream of auxin. We are therefore able to suggest a model of apical dominance that integrates auxin, sucrose, strigolactones, cytokinins and gibberellins and describes differences in signalling across stages of bud release to sustained growth.