Mineral nutrition of plants greatly depends on both environmental conditions, particularly of soils, and the genetic background of the plant itself. Being sessile, plants adopted a range of strategies for sensing and responding to nutrient availability to optimize development and growth, as well as to protect their metabolisms from heavy metal toxicity. Such mechanisms, together with the soil environment, meaning the soil microorganisms and their interaction with plant roots, have been extensively studied with the goal of exploiting them to reclaim polluted lands; this approach, defined phytoremediation, will be the subject of this review. The main aspects and innovations in this field are considered, in particular with respect to the selection of efficient plant genotypes, the application of improved cultural strategies, and the symbiotic interaction with soil microorganisms, to manage heavy metal polluted soils.

Plants, like all other organisms, have evolved different mechanisms to maintain physiological concentrations of essential metal ions and to minimize exposure to non-essential heavy metals. Some mechanisms are ubiquitous because they are also required for general metal homeostasis, and they minimize the damage caused by high concentrations of heavy metals in plants by detoxification, thereby conferring tolerance to heavy metal stress. Other mechanisms target individual metal ions (indeed some plants have more than one mechanism to prevent the accumulation of specific metals) and these processes may involve the exclusion of particular metals from the intracellular environment or the sequestration of toxic ions within compartments to isolate them from sensitive cellular components. As a first line of defense, many plants exposed to toxic concentrations of metal ions attempt to prevent or reduce uptake into root cells by restricting metal ions to the apoplast, binding them to the cell wall or to cellular exudates, or by inhibiting long distance transport. If this fails, metals already in the cell are addressed using a range of storage and detoxification strategies, including metal transport, chelation, trafficking, and sequestration into the vacuole. When these options are exhausted, plants activate oxidative stress defense mechanisms and the synthesis of stress-related proteins and signaling molecules, such as heat shock proteins, hormones, and reactive oxygen species.

Heavy metals are often present naturally in soils, but many human activities (e.g. mining, agriculture, sewage processing, the metal industry and automobiles) increase their prevalence in the environment resulting in concentrations that are toxic to animals and plants. Excess heavy metals affect plant physiology by inducing stress symptoms, but many plants have adapted to avoid the damaging effects of metal toxicity, using strategies such as metal chelation, transport and compartmentalization. Understanding the molecular basis of heavy metal tolerance in plants will facilitate the development of new strategies to create metal-tolerant crops, biofortified foods and plants suitable for the phytoremediation of contaminated sites.

Plants need many different metal elements for growth, development and reproduction, which must be mobilized from the soil matrix and absorbed by the roots as metal ions. Once taken up by the roots, metal ions are allocated to different parts of the plant by the vascular tissues. Metals are naturally present in the soil, but human activities, ranging from mining and agriculture to sewage processing and heavy industry, have increased the amount of metal pollution in the environment. Plants are challenged by environmental metal ion concentrations that fluctuate from low to high toxic levels, and have therefore evolved mechanisms to cope with such phenomena. In this review, we focus on recent data that provide insight into the molecular mechanisms of metal absorption and transport by plants, also considering the effect of metal deficiency and toxicity. We also highlight the positive effects of some non-essential metals on plant fitness.

The genetic engineering of plants to facilitate the reclamation of soils and waters contaminated with inorganic pollutants is a relatively new and evolving field, benefiting from the heterologous expression of genes that increase the capacity of plants to mobilize, stabilize and/or accumulate metals. The efficiency of phytoremediation relies on the mechanisms underlying metal accumulation and tolerance, such as metal uptake, translocation and detoxification. The transfer of genes involved in any of these processes into fast-growing, high-biomass crops may improve their reclamation potential. The successful phytoextraction of metals/metalloids and their accumulation in aerial organs have been achieved by expressing metal ligands or transporters, enzymes involved in sulfur metabolism, enzymes that alter the chemical form or redox state of metals/metalloids and even the components of primary metabolism. This review article considers the potential of genetic engineering as a strategy to improve the phytoremediation capacity of plants in the context of heavy metals and metalloids, using recent case studies to demonstrate the practical application of this approach in the field.

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Pseudomonas putida is a saprophytic bacterium with remarkable environmental adaptability and the capacity to tolerate high concentrations of heavy metals. The strain P. putida-Cd001 was isolated from soil contaminated with Cd, Zn and Pb. Membrane-associated and cytosolic proteomes were analyzed to identify proteins whose expression was modulated in response to 250 μM CdSO4. We identified 44 protein spots in the membrane and 21 in the cytosolic fraction differentially expressed in Cd-treated samples compared to untreated controls. Outer membrane porins from the OprD and OprI families were less abundant in bacteria exposed to Cd, whereas those from the OprF and OprL, OprH and OprB families were more abundant, reflecting the increased need to acquire energy sources, the need to maintain membrane integrity and the process of adaptation. Components of the efflux system, such as the CzcB subunit of the CBA system, were also induced by Cd. Analysis of the cytosolic proteome revealed that proteins involved in protein synthesis, degradation and folding were induced along with enzymes that combat oxidative stress, showing that the entire bacterial proteome is modulated by heavy metal exposure. This analysis provides new insights into the adaptation mechanisms used by P. putida-Cd001 to survive in Cd-polluted environments.

To succeed in life, living organisms have to adapt to the environmental issues to which they are subjected. Some plants, defined as hyperaccumulators, have adapted to metalliferous environments, acquiring the ability to tolerate and accommodate high amounts of toxic metal into their shoot, without showing symptoms of toxicity. The determinants for these traits and their mode of action have long been the subject of research, whose attention lately moved to the evolution of the hypertolerance and hyperaccumulation traits. Genetic evidence indicates that the evolution of both traits includes significant evolutionary events that result in species-wide tolerant and accumulating backgrounds. Different edaphic environments are responsible for subsequent refinement, by local adaptive processes, leading to specific strategies and various degrees of hypertolerance and hyperaccumulation, which characterize metallicolous from non-metallicolous ecotypes belonging to the same genetic unit. In this review, we overview the most updated concepts regarding the evolution of hyperaccumulation and hypertolerance, highlighting also the ecological context concerning the plant populations displaying this fascinating phenomenon.