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

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 microM for Cu and 500 microM for Al and Zn, corresponding to about 30 microM Cu2+, 140 microM Al3+ and 290 microM Zn2+. Simultaneous with the increase in organic acid anion exudation, sulfate and the nutrient cations K+, Ca2+ and Mg2+ 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 microM and above, at Zn concentrations of 500 microM and above, and at an Al concentration of 1000 microM, and callose formation, which was observed solely in response to Al concentrations of 500 microM 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.