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We studied the effects of a 2-day exposure to aluminum (Al), copper (Cu) and zinc (Zn) stress on root exudates and root apices of aseptically grown poplar ( Populus tremula  L.) cuttings. Aluminum induced root exudation of oxalate and citrate, Cu induced root exudation of oxalate, malate and formate, and Zn induced root exudation of formate. The threshold treatment concentrations were 100 μM for Cu and 500 μM for Al and Zn, corresponding to about 30 μM Cu 2+ , 140 μM Al 3+  and 290 μM Zn 2+ . Simultaneous with the increase in organic acid anion exudation, sulfate and the nutrient cations K + , Ca 2+  and Mg 2+  were released into the solutions. Significant positive correlations between the organic acid anions and the cations indicate possible co-release. Toxicity symptoms of the poplar roots included browning of the root apices, which occurred at Cu concentrations of 50 μM and above, at Zn concentrations of 500 μM and above, and at an Al concentration of 1000 μM, and callose formation, which was observed solely in response to Al concentrations of 500 μM or higher. The results indicate that the composition of the exuded organic acid anions from poplar roots and the toxicity symptoms are specific to each of the applied heavy metals.

Exudation of organic acid anions from poplar roots after exposure to Al, Cu and Zn