Root Engineering in Barley: Increasing Cytokinin Degradation Produces a Larger Root System, Mineral Enrichment in the Shoot and Improved Drought Tolerance

Root size and architecture are important crop plant traits, as they determine access to water and soil nutrients. The plant hormone cytokinin is a negative regulator of root growth and branching. Here, we generated transgenic barley ( Hordeum vulgare ) plants with an enlarged root system by enhancing cytokinin degradation in roots to explore the potential of cytokinin modulations in improving root functions. This was achieved through root-specific expression of a CYTOKININ OXIDASE/DEHYDROGENASE gene. Enhanced biomass allocation to roots did not penalize shoot growth or seed yield, indicating that these plants were not source limited. In leaves of transgenic lines, the concentrations of several macroelements and microelements were increased, particularly those with low soil mobility (phosphorus, manganese, and zinc). Importantly, seeds contained up to 44% more zinc, which is beneficial for human nutrition. Transgenic lines also demonstrated dampened stress responses to long-term drought conditions, indicating lower drought sensitivity. Taken together, this work demonstrates that root engineering of cereals is a promising strategy to improve nutrient efficiency, biofortification, and drought tolerance.