DELLA protein function in growth responses to canopy signals

Plants can sense neighbour competitors through light-quality signals and respond with shade-avoidance responses. These include increased shoot elongation, which enhances light capture and thus competitive power. Such plant-plant interactions therefore profoundly affect plant development in crowded populations. Shade-avoidance responses are tightly coordinated by interactions between light signals and hormones, with essential roles for the phytochrome B photoreceptor [sensing the red:far red (R:FR) ratio] and the hormone gibberellin (GA). The family of growth-suppressing DELLA proteins are targets for GA signalling and are proposed to integrate signals from other hormones. However, the importance of these regulators has not been studied in the ecologically relevant, complex realm of plant canopies. Here we show that DELLA abundance is regulated during growth responses to neighbours in dense Arabidopsis stands. This occurs in a R:FR-dependent manner in petioles, depends on GA, and matches the induction kinetics of petiole elongation. Similar interactions were observed in the growth response of seedling hypocotyls and are general for a second canopy signal, reduced blue light. Enhanced DELLA stability in the gai mutant inhibits shade-avoidance responses, indicating that DELLA proteins constrain shade-avoidance. However, using multiple DELLA knockout mutants, we show that the observed DELLA breakdown is not sufficient to induce shade-avoidance in petioles, but plays a more central role in hypocotyls. These data provide novel information on the regulation of shade-avoidance under ecologically important conditions, defining the importance of DELLA proteins and GA and unravelling the existence of GA- and DELLA-independent mechanisms.