Cadmium toxicity and tolerance in vascular plants

Heavy metals constitute a very heterogeneous group of elements widely varied in their chemical properties and biological functions. Heavy metals are kept under environmental pollutant category due to their toxic effects on plants, animals and human being. Heavy metal contamination of soil results from anthropogenic as well as natural activities. Anthropogenic activities such as mining, smelting operation and agriculture have locally increased the levels of heavy metals such as Cd, Co, Cr, Pb, As and Ni in soil up to dangerous levels. Heavy metals are persistent in nature, therefore get accumulated in soils and plants. Heavy metals interfere with physiological activities of plants such as photosynthesis, gaseous exchange and nutrient absorption, and cause reductions in plant growth, dry matter accumulation and yield. Heavy metals also interfere with the levels of antioxidants in plants, and reduce the nutritive value of the produce. Dietary intake of many heavy metals through consumption of plants has long term detrimental effects on human health.

Biological effects of heavy metals: an overview.

This study focuses on the characterization of four bacterial isolates from heavy metal-polluted rhizosphere in order to examine their plant growth promoting (PGP) activity. The PGP activity on the canola ( Brassica napus ) of the strains which showed cadmium resistance and multiple PGP traits was assessed in the presence and in the absence of Cd 2+ . The strains, Pseudomonas tolaasii ACC23, Pseudomonas fluorescens ACC9, Alcaligenes sp. ZN4 and Mycobacterium sp. ACC14 showed 1-aminocyclopropane-1-carboxylate deaminase (ACCD) activity. They also synthesized ACCD enzyme in vitro when 0.4 mM Cd 2+ was added to the growth medium. The presence of the metal, however, reduced the ACCD activity in Alcaligenes sp. ZN4 and Mycobacterium sp. ACC14, while it did not affect the ACCD activity of P. tolaasii ACC23 and P. fluorescens ACC9. ACC9 and ACC23 produced indole acetic acid (IAA) and siderophores, while ACC14 produced only IAA. IAA and siderophores were produced more actively under Cd-stress. Root elongation assays conducted on B. napus under gnotobiotic conditions demonstrated increases (from 34% up to 97%) in root elongation of inoculated canola seedlings compared to the control plants. Subsequently, the effect of inoculation with these strains on growth and uptake of Cd 2+ in roots and shoots of canola was studied in pot experiments using Cd-free and Cd-treated (15 μg Cd 2+  g −1  dw) soil. Inoculation with P. tolaasii ACC23, P. fluorescens ACC9 and Mycobacterium sp. ACC14 promoted the growth of plants at concentrations of 0 and 15 μg Cd 2+  g −1  soil. The maximum growth was observed in the plants inoculated with P. tolaasii ACC23. The strains did not influence the specific accumulation of cadmium in the root and shoot systems, but all increased the plant biomass and consequently the total cadmium accumulation. The present observations showed that the bacterial strains used in this study protect the plants against the inhibitory effects of cadmium, probably due to the production of IAA, siderophores and ACCD activity.

Improvement of Brassica napus growth under cadmium stress by cadmium-resistant rhizobacteria

Heavy metal toxicity is one of the major abiotic stresses leading to hazardous health effects in animals and plants. Because of their high reactivity they can directly influence growth, senescence and energy synthesis processes. In this review a new indirect mechanism of heavy metal action is proposed. This mechanism is connected with the generation of reactive oxygen species (especially H2O2) and jasmonate and ethylene signaling pathways and shows that toxicity symptoms observed in plants may result from direct heavy metal influence as well as the activity of some signaling molecules induced by the stress action.

Signaling responses in plants to heavy metal stress

Heavy metals are natural non-biodegradable constituents of the Earth's crust that accumulate and persist indefinitely in the ecosystem as a result of human activities. Since the industrial revolution, the concentration of cadmium, arsenic, lead, mercury and zinc, amongst others, have increasingly contaminated soil and water resources, leading to significant yield losses in plants. These issues have become an important concern of scientific interest. Understanding the molecular and physiological responses of plants to heavy metal stress is critical in order to maximize their productivity. Recent research has extended our view of how plant hormones can regulate and integrate growth responses to various environmental cues in order to sustain life. In the present review we discuss current knowledge about the role of the plant growth hormones abscisic acid, auxin, brassinosteroid and ethylene in signaling pathways, defense mechanisms and alleviation of heavy metal toxicity.

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Interactions between plant hormones and heavy metals responses.

Six day old rice seedlings (Oryza sativa L. cv. Bahia) were grown for 5 or 10 days in a nutrient solution with either Cd (0.01, 0.1 mmol/l) or Ni (0.1, 0.5 mmol/l). Both Cd and Ni reduced the length of shoots and roots depending on the concentration and type of ion tested. On the other hand, the dry weight to fresh weight ratio was increased by heavy metal treatments, especially in the aerial part of 0.5 mmol/l Ni treated plants. The application of 0.1 mmol/l Cd and 0.5 mmol/l Ni to the seedlings produced an inhibition of the transport of carbohydrate reserves from the seeds from which plants were developing, to the rest of the plant. Net photosynthesis was also inhibited in treated plants. However, the total carbohydrate content in the shoots of these plants was higher than in controls. Thus, the starch, soluble sugars and sucrose content in the shoots of 0.5 mmol/l Ni treated plants was respectively up to 2.6, 2.8 and 4 times greater compared to controls. The distribution of assimilates between organs was also affected by the treatment: the carbohydrate content increased in the stem and second leaf but it was not affected or decreased in the root and third leaf. Although less evident, the effect of Cd on carbohydrate distribution and content was similar to that of Ni. The possible mechanisms involved in the abnormal carbohydrate accumulation and distribution are discussed.

Influence of cadmium and nickel on growth, net photosynthesis and carbohydrate distribution in rice plants.

The effect of external applications of gibberellins (GA3) and abscisic acid (ABA) on the growth, carbohydrate content, and net photosynthesis of heavy metal-stressed rice plants (Oryza sativa cv. Bahia) was investigated. Treatment with cadmium (0.1 mm) and nickel (0.5 mm) inhibited rice growth and stimulated carbohydrate accumulation, especially in seeds from which seedlings were developing, stems, and first leaves. The addition of GA3 (14 μm) to the rice culture solution together with Cd or Ni partially reversed the effects of heavy metals, stimulating growth as well as mobilization of carbohydrate reserves in seeds from which seedlings had developed. GA3 increased the sugar content in roots and second and third leaves and also modified the carbohydrate distribution pattern compared with heavy metal-treated plants. In contrast to GA3, ABA (19 μm) supplied to rice cultures potentiated the effect of heavy metals, inhibiting the growth of young leaves and the translocation of storage products from source to sink organs. In addition, sugars were accumulated in roots and second leaf but not in the third leaf, the extension in length of which was also inhibited by the treatment. Net photosynthesis rates recovered transitorily in Cd-treated plants after the addition of hormones. The possible relationship between growth and carbohydrate distribution, as well as the involvement of hormones, in the response of plant to heavy metal stress is discussed.

J. L. Moya

Papers

Influence of cadmium and nickel on growth, net photosynthesis and carbohydrate distribution in rice plants.

Heavy metal-hormone interactions in rice plants: Effects on growth, net photosynthesis, and carbohydrate distribution