RESUMEN
Most angiosperm genomes contain several genes encoding metallothionein (MT) proteins that can bind metals including copper (Cu) and zinc (Zn). Metallothionein genes are highly expressed under various conditions but there is limited information about their function. We have studied Arabidopsis mutants that are deficient in multiple MTs to learn about the functions of MTs in plants. T-DNA insertions were identified in four of the five Arabidopsis MT genes expressed in vegetative tissues. These were crossed to produce plants deficient in four MTs (mt1a/mt2a/mt2b/mt3). The concentration of Cu was lower in seeds but higher in old leaves of the quad-MT mutant compared to wild-type plants. Experiments with stable isotopes showed that Cu in seeds came from two sources: directly from roots and via remobilization from other organs. Mobilization of Cu out of senescing leaves was disrupted in MT-deficient plants. Tolerance to Cu, Zn and paraquat was unaffected by MT deficiency but these plants were slightly more sensitive to cadmium (Cd). The quad-MT mutant showed no change in resistance to a number of microbial pathogens, or in the progression of leaf senescence. Although these MTs are not required to complete the plant's life cycle, MTs are important for Cu homeostasis and distribution in Arabidopsis.