Structural changes induced by NaCl in companion and transfer cells of Medicago sativa blades

Medicago sativa var. Gabes is a perennial glycophyte that develops new shoots even in high salinity (150 mM NaCl). In the upper exporting leaves, K(+) is high and Na(+) is low by comparison with the lower leaves, where Na(+) accumulation induces chlorosis after 4 weeks of NaCl treatment. By secondary ion mass spectroscopy, a low Na(+)/K(+) ratio was detected in the phloem complex of blade veins in these lower leaves. By transmission electron microscopy, the ultrastructural features were observed in the phloem complex. In the upper leaves of both control and NaCl-treated plants, companion cells in minor veins were found to be transfer cells. These cells may well be involved in the intravenous recycling of ions and in Na(+) flowing out of exporting leaves. Under the effect of NaCl, companion cells in the main veins develop transfer cell features, which may favor the rate of assimilate transport from exporting leaves toward meristems, allowing the positive balance necessary for the survival in salt conditions. These features no longer assist the lower leaves when transfer cells are necrotized in both minor and main veins of NaCl-treated plants. As transfer cells are the only degenerating phloem constituent, our observations emphasize their role in controlling nutrient (in particular, Na(+)) fluxes associated with the stress response.