Secondary Active Transporters Decrease Cytokinin Flow into Barley Shoots and Inhibit their Growth under Phosphate Deficit

Abstract

Secondary active transport is one of the most important mechanisms controlling cytokinin distribution between shoots and roots which enables the adaptive growth reaction of barley. Concentrations (ng/g of fresh mass) and contents (ng/organ) of different forms of cytokinins were determined in shoots and roots of barley plants by immunoassay. It was shown that under phosphate deficit the maintenance of root growth, inhibition of root branching and decline in shoot mass of barley plants (Hordeum vulgare 'Prairie') was due to decline in shoot cytokinins and their accumulation in the roots. Protonophore carbonyl cyanide-m-chlorophenylhydrazone as an inhibitor of secondary active transport leveled off hormonal reaction to phosphate deficit: it prevented accumulation of cytokinins in roots, increased their content in shoots and changed the percentage of zeatin and its derivatives in total cytokinin content in roots and shoots. A data analysis showed that in roots, protonophore did not change significantly the content of ribosides and nucleotides of zeatin, while uptake of free zeatin by cells was significantly affected by the protonophore and its content decreased dramatically. The role of secondary active trans-membrane transfer is discussed in the context of cytokinins transport from roots to shoots under phosphate deficit