Shovel roots: a unique stress-avoiding developmental strategy of the legume plant Hedysarum coronarium L.

Hedysarum coronarium (sulla) is a legume native to the Mediterranean basin, known for its broad tolerance to various environmental stresses, and its ability to thrive without signs of chlorosis when growing in arid and alkaline soils up to pH 9.6. A unique but poorly known morphological feature of its root system is the production of "shovels", modified lateral roots that acquire a curved and flattened shape. A combined structural and functional analysis was undertaken to define the nature and role of the shovel roots using various microscopy techniques, histochemical stains, STEM - energy dispersive X-ray microanalysis, infrared spectroscopy, and plant cultivation in different conditions. We found that sulla displays remarkable unique rhizosphere-buffering properties at both ends of the pH scale, and that shovels act as efficient calcium-absorbing organs that accumulate this cation intracellularly as insoluble crystalline salts. Such bioaccumulation results in a localized depletion of CaCO3 from the soil. As a consequence of this removal of the pivotal carbonate buffering system, the iron-solubilizing acidification activities of the roots can become effective. Further tests revealed that the factor triggering shovel development is exposure of roots to iron oxide. This signal, reporting at once both iron presence and alkalinity, assures the availability of iron nutrient reserves upon acidification of the local microenvironment surrounding the roots. These findings, besides casting light on a novel and unique botanical phenomenon, offer the potential to exploit sulla's model and genes for the improvement of other crops to sustain productivity in a scenario of climate warming and increasing desertification.