TcYSL3, a member of the YSL gene family from the hyper‐accumulator Thlaspi caerulescens, encodes a nicotianamine‐Ni/Fe transporter

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Abstract
Summary: The two main features of plant hyper‐accumulator species are the massive translocation of heavy metal ions to the aerial parts and their tolerance to such high metal concentrations. Recently, several lines of evidence have indicated a role for nicotianamine (NA) in metal homeostasis, through the chelation and transport of NA–metal complexes. The function of transport of NA–metal chelates, required for the loading and unloading of vessels, has been assigned to the Yellow Stripe 1 (YSL)‐Like family of proteins. We have characterized three YSL genes in Thlaspi caerulescens in the context of hyper‐accumulation. The three YSL genes are expressed at high rates compared with their Arabidopsis thaliana homologs but with distinct patterns. While TcYSL7 was highly expressed in the flowers, TcYSL5 was more highly expressed in the shoots, and the expression of TcYSL3 was equivalent in all the organs tested. In situ hybridizations have shown that TcYSL7 and TcYSL5 are expressed around the vasculature of the shoots and in the central cylinder in the roots. The exposure to heavy metals (Zn, Cd, Ni) does not affect the high and constitutive expression of the TcYSL genes. Finally, we have demonstrated by mutant yeast complementation and uptake measurements that TcYSL3 is an Fe/Ni–NA influx transporter. This work provides therefore molecular, histological and biochemical evidence supporting a role for YSL transporters in the overall scheme of NA and NA–metal, particularly NA–Ni, circulation in a metal hyper‐accumulator plant.