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Full Journal - Journal of Cell and Molecular Biology - Haliç Üniversitesi

Full Journal - Journal of Cell and Molecular Biology - Haliç Üniversitesi

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46 Gülriz Bayçu<br />

Cd Contamination in the environment <strong>and</strong> toxic<br />

effects on plants<br />

Heavy metals such as cadmium (Cd), chromium (Cr),<br />

copper (Cu), mercury (Hg), lead (Pb), aluminium<br />

(Al), nickel (Ni), etc., are important environmental<br />

pollutants particularly in areas where there is high<br />

antropogenic pressure (Abrahamson et al.,1992;<br />

Sanità di Toppi <strong>and</strong> Gabbrielli, 1999). This group <strong>of</strong><br />

elements is toxic to all organisms at varying<br />

concentrations (Speiser et al., 1992). Cd is a toxic<br />

element that normally occurs in very low<br />

concentrations in soils (Wagner, 1993). However, it is<br />

released into the environment by urban traffic, metalworking<br />

industries, heating systems, power stations,<br />

cement factories, waste incinerators <strong>and</strong> phosphate<br />

fertilizers (Sanità di Toppi <strong>and</strong> Gabbrielli, 1999). In<br />

the areas that have been subjected to mining, the<br />

concentration can be high, varying from 100-600 mg<br />

kg -1 dry weight. In addition, following the application<br />

<strong>of</strong> sewage sludge to agricultural l<strong>and</strong>, Cd can<br />

accumulate in the top soil (Lombi et al., 2000). In<br />

humans, Cd is suspected carcinogen it displace Ca or<br />

Zn in proteins <strong>and</strong> can cause oxidative stress (Steffens<br />

<strong>and</strong> Bagchi, 1995).<br />

Photosynthetic organisms are the principal entry<br />

point <strong>of</strong> metals into the food chain leading to animals<br />

<strong>and</strong> man (Rauser, 1990). Heavy metals such as Cu<br />

<strong>and</strong> Zn are essential for normal plant growth, but<br />

elevated concentrations can result in growth<br />

inhibition <strong>and</strong> toxicity symptoms. Non-essential<br />

metals like Cd <strong>and</strong> Pb are potentially highly toxic <strong>and</strong><br />

show toxic effects even at very low concentrations<br />

(Hall, 2002). However, a number <strong>of</strong> plants (termed<br />

hyperaccumulators) that grow on metalliferous soils,<br />

are able to translocate Cd from the roots <strong>and</strong><br />

accumulate it in high concentrations in the shoots<br />

(Chardonnens et al., 1998). It has been suggested that<br />

such plants would be <strong>of</strong> considerable value in the<br />

remediation <strong>of</strong> soils that are heavily contaminated<br />

with heavy metals (Zhu et al., 1999).<br />

Several mechanisms are known to allow plants<br />

<strong>and</strong> other organisms to tolerate the presence <strong>of</strong> toxic<br />

non-essential metal ions inside the cell. Physiological<br />

studies indicate that heavy metal tolerance is one <strong>of</strong><br />

the prerequisites <strong>of</strong> heavy metal hyperaccumulation<br />

in plants (Raskin et al.,1997). Wide information is<br />

available on ecological impact <strong>of</strong> Cd, especially as<br />

concerns its motility in soil <strong>and</strong> uptake by plants<br />

(Leita et al., 1991). Although Cd is not an essential<br />

nutrient for plants, the metal ion is taken up rapidly by<br />

the roots <strong>and</strong> on most occasions causes inhibition <strong>of</strong><br />

growth (Leita et al., 1991). It has been demonstrated<br />

that Cd is not only easily available to plants from soil<br />

or other substrates, but also it is toxic to them at much<br />

lower concentrations than other heavy metals like Zn,<br />

Pb or Cu. The degree to which higher plants are able<br />

to take up Cd depends on its bioavailability,<br />

modulated by the presence <strong>of</strong> organic matter, pH,<br />

redox potential, temperature <strong>and</strong> concentrations <strong>of</strong><br />

other elements (Sanità di Toppi <strong>and</strong> Gabbrielli,<br />

1999). Cd is believed to penetrate the root through the<br />

cortical tissue <strong>and</strong> it is readily taken up <strong>and</strong><br />

transported within the plant through an apoplastic<br />

<strong>and</strong>/or a symplastic pathway (Page et al., 1981; Salt et<br />

al., 1997). Phytotoxicity has been observed to be<br />

dependent upon plant species as well as Cd<br />

concentration in substrate (Leita et al., 1991). Cd<br />

interferes with plant metabolism <strong>and</strong> in a very general<br />

way, causes leaf roll, chlorosis, growth <strong>and</strong> yield<br />

reduction (Leita et al., 1991). Inhibition <strong>of</strong><br />

ribonuclease <strong>and</strong> nitrate reductase activity, interaction<br />

with the water balance, decrease in chlorophyll<br />

content, inhibition <strong>of</strong> stomatal opening, reduction <strong>of</strong><br />

normal H + /K + exchange, production <strong>of</strong> oxidative<br />

stress <strong>and</strong> enhanced lipid peroxidation are the most<br />

important phytotoxic effects <strong>of</strong> Cd (Sanità di Toppi<br />

<strong>and</strong> Gabbrielli, 1999).<br />

Heavy metal stress <strong>and</strong> detoxification mechanisms<br />

All living cells are confronted with the dilemma that<br />

on one side they need certain amounts <strong>of</strong> free heavy<br />

metal ions (such as Zn, Cu , Ni , etc.) for their normal<br />

metabolic function, <strong>and</strong> on the other side they have to<br />

protect themselves from an intracellular excess <strong>of</strong><br />

these metal ions which would lead to cell death. This<br />

dilemma can only be overcome by a stringent<br />

regulation <strong>of</strong> free metal ion concentrations within the<br />

cells (Tomsett <strong>and</strong> Thurman, 1988; Gekeler et al.,<br />

1989).<br />

In general, resistance to excessively available<br />

chemical elements can be based on avoidance, i.e.<br />

exclusion <strong>of</strong> the element from the body, or on<br />

tolerance, i.e. the ability to survive, grow <strong>and</strong><br />

reproduce with the element present at elevated<br />

concentrations in the body (Levitt, 1980; Ernst et al.,

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