RESUMEN
Arsenic (As) contaminates the food chain and decreases agricultural production through impairing plants, particularly due to oxidative stress. To better understand the As tolerance mechanisms, two contrasting tobacco genotypes: As-sensitive Nicotiana sylvestris and As-tolerant N.tabacum, cv. 'Wisconsin' were analyzed. The most meaningful differences were found in the carbohydrate status, neglected so far in the As context. In the tolerant genotype, contrary to the sensitive one, net photosynthesis rates and saccharide levels were unaffected by As exposure. Importantly, the total antioxidant capacity was far stronger in the As-tolerant genotype, based on higher antioxidants levels (e.g., phenolics, ascorbate, glutathione) and activities and/or appropriate localizations of antioxidative enzymes, manifested as reverse root/shoot activities in the selected genotypes. Accordingly, malondialdehyde levels, a lipid peroxidation marker, increased only in sensitive tobacco, indicating efficient membrane protection in As-tolerant species. We bring new evidence of the orchestrated action of a broad spectrum of both antioxidant enzymes and molecules essential for As stress coping. For the first time, we propose robust carbohydrate metabolism based on undisturbed photosynthesis to be crucial not only for subsidizing C and energy for defense but also for participating in direct reactive oxygen species (ROS) quenching. The collected data and suggestions can serve as a basis for the selection of plant As phytoremediators or for targeted breeding of tolerant crops.
RESUMEN
MAIN CONCLUSION: This review provides insights into As toxicity in plants with focus on photosynthesis and sugar metabolism as important arsenic targets and simultaneously defence tools against accompanying oxidative stress. Heavy metal contamination is a great problem all over the world. Arsenic, a metalloid occurring naturally in the Earth's crust, also massively spreads out in the environment by human activities. Its accumulation in crops poses a severe health risk to humans and animals. Besides the restriction of human-caused contamination, there are two basic ways how to cope with the problem: first, to limit arsenic accumulation in harvestable parts of the crops; second, to make use of some arsenic hyperaccumulating plants for phytoremediation of contaminated soils and waters. Progress in the use of both strategies depends strongly on the level of our knowledge on the physiological and morphological processes resulting from arsenic exposure. Arsenic uptake is mediated preferentially by P and Si transporters and its accumulation substantially impairs plant metabolism at numerous levels including damages through oxidative stress. Rice is a predominantly studied crop where substantial progress has been made in understanding of the mechanisms of arsenic uptake, distribution, and detoxification, though many questions still remain. Full exploitation of plant potential for soil and water phytoremediations also requires deep understanding of the plant response to this toxic metalloid. The aim of this review is to summarize data regarding the effect of arsenic on plant physiology with a focus on mechanisms providing increased arsenic tolerance and/or hyperaccumulation. The emphasis is placed on the topic unjustifiably neglected in the previous reviews - i.e., carbohydrate metabolism, tightly connected to photosynthesis, and beside others involved in plant ability to cope with arsenic-induced oxidative and nitrosative stresses.