Physiological responses to hypoxia and Manganese in Eucalyptus clones with differential tolerance to Vale do Rio Doce shoot dieback

Abstract

Vale do Rio Doce shoot dieback (VRDSD) is an anomaly whose cause seems to be associated with hypoxic conditions and their consequences (excess Mn and Fe) triggered by elevation of the water table in areas with poor drainage. Different plants have distinct survival strategies under this form of stress. The objective of this study was to understand the physiological responses involved in the differential tolerance of eucalyptus clones to VRDSD and their relationship to hypoxia and excess Mn. A hydroponic experiment was carried out using a 2 × 2 × 2 factorial arrangement, two eucalyptus clones with different levels of tolerance to VRDSD (sensitive Urograndis hybrid - 1213; and the tolerant Rio Claro hybrid - Eucalyptus grandis x unknown - 2719), two concentrations of O2 (8 and 4 mg L-1), and two Mn concentrations (1.39 and 300 mg L-1) in a randomized block design (RBD) with three replicates. Forty-day-old clones were maintained in Clark nutrient solution for 30 days. After this period, the treatments were applied for 11 days. Plant gaseous exchange shoot and root production, and the quantity of enzymes related to oxidative stress in leaves and roots were evaluated. In the tolerant clone, reactive oxygen species (ROS) were produced under hypoxic conditions, accompanied by reduction in production of dry matter, malondialdehyde (MDA), and in activity of the enzyme alcohol dehydrogenase (ADH). However, this clone had greater production of superoxide dismutase (SOD) under these conditions, an enzyme responsible for detoxification of ROS, which acts as part of the Low Oxygen Quiescence Syndrome (LOQS). In contrast, sensitive clones did not exhibit expressive reductions in growth or changes in the leaf/root ratio. These clones formed large quantities of adventitious roots and had high levels of MDA and ADH and low levels of SOD. Therefore, sensitive clones appear not to be prepared for detoxification of ROS and other toxic metabolites, but rather adopt morphological escape mechanisms, the Low Oxygen Escape Syndrome (LOES), in response to hypoxia. Thus, the period of soil waterlogging may cause the death of large numbers of roots in sensitive clones, limiting their ability to absorb water and nutrients and culminating in the death of these plants. Excess Mn seems to aggravate the damage caused by hypoxia, but it is not the causal agent of VRDSD.