Potential of Brassicaceae Burnett (Mustard Family Angiosperms) in Phytoremediation of Heavy Metals

International Journal of Scientific Research in Environmental Sciences, 2 (4), pp. 120-138, 2014 Available online at http://www.ijsrpub.
com/ijsres ISSN: 2322-4983; 2014 IJSRPUB http://dx.doi.org/10.12983/ijsres-2014-p0120-0138
Review Paper Potential of Brassicaceae Burnett (Mustard family; Angiosperms) in Phytoremediation of Heavy Metals
Ramesh Chandra Pantola, Afroz Alam*
Department of Bioscience and Biotechnology, Banasthali Vidyapith, Tonk- 304 022 (Rajasthan), India *Corresponding author Email: afrozalamsafvi@gmail.com
Received 26 January 2014; Accepted 19 March 2014
Abstract. Phytoremediation technology is using plants to clean up contaminated sites is a promising loom to restore the environment and ecosystem. Pollution of the environment with toxic organic and heavy metal pollutants is one of major problem facing developed and developing nations today. Heavy metals restrain biodegradation of chlorinated organics by interacting with enzymes directly concerned in biodegradation or those involved in common metabolism. Prophecy of heavy metal toxicity effects on organic pollutant biodegradation in co-contaminated soil is tricky since heavy metals may be present in a variety of chemical and physical forms. Even if much remains to be studied, phytoremediation will clearly play some role in the stabilization and remediation of many contaminated sites. The main factor driving the execution of phytoremediation projects are low costs with considerable improvements in site aesthetics and the potential for ecosystem restoration. In this review, the potential and various aspects of few Brassicaceae (Mustard family) members in phytoremediation have been discussed due to their well known natural tolerant against various environmental stresses including heavy metals. Keywords: Phytoremediation, Brassicaceae Burnett, Heavy metals
1. INTRODUCTION Phytoremediation is an assemblage of such technologies that use plants to reduce, remove, degrade, or immobilize environmental toxins, primarily those of anthropogenic origin, with the aim of restoring specific sites to a condition utilizable for private or public applications. The generic term Phytoremediation consists of the Greek prefix phyto (plant), attached to the Latin root remedian (to correct or remove an evil). The mechanism of Phytoremediation includes enhanced rhizosphere biodegradation (Ghosh and Singh, 2005). Many plants produce molecules called peptides that bind metals for storage in cell compartments. It is vital that the plants, which are selected for phytoremediation, must be suitable for agricultural practices and is able to produce enough biomass with maximum uptake of metallic compounds in order to be effective. The use of plants as a remeditive measure for extraction of contaminants from the environment or for lowering of their toxicity is defined as phytoremediation (Salt and Kramer, 2000). Metal hyper-accumulator plant species are able to accumulate at least 0.1% of the leaf dry weight in a heavy metal (Baker and Whiting, 2002). This is a low
cost and eco-friendly means of reclaiming heavy metal contaminated soils (Rajakaruna et al., 2006). As biological processes are ultimately solar-driven, phytoremediation is on average tenfold cheaper than engineering-based remediation methods (Marques et al., 2009). As with vegetation; they have to produce a large biomass and to have the ability to accumulate significant quantities of heavy metals in their stems (Kumar et al., 1995; Cunningham and Ow, 1996; Blaylock et al., 1997; McGrath, 1998). However, there are evidences that 400 hyper-accumulative plants are capable of extracting contaminants (Salt and Kramer, 2000). Most of them are unsuitable for phytoremediation, because they have a slow growth rate and small size. Metal toxicity issues in plants and soils are a significant problem throughout the world. The understanding of relationships between bioavailable metal fractions in the soil and plant responses to metals regarding toxicity in plants is mandatory for phytoremedian campaigns. So, the focus of present review is to elaborate the techniques, methods and procedures for end users who are using the plants for rehabilitation of metal contaminated soil.
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Pantola and Alam Potential of Brassicaceae Burnett (Mustard family; Angiosperms) in Phytoremediation of Heavy Metals
1.1. Methods of Phytoremediation 1.1.1. Phytoextraction The use of plants to remove contaminants from the environment and concentrate them in above ground plant tissue is known as phytoextraction. It is also known as phyto-accumulation, the uptake of contaminants by plant root and the translocation/accumulation of contaminants into the plant shoots and leaves. Research and development efforts focus on two areas of study: (1) remediation of contaminants such as Pb, As, Cr, Hg, and radio nuclides and (2) mining, or recovery, of inorganic compounds, mainly Ni and Cu is having intrinsic economic value. Phyto-extraction can be used in both water and soil environments. Phyto-extraction involves the removal of toxins, especially heavy metals and metalloids, by the roots of the plants with subsequent transport to aerial plant organs. Pollutants accumulated in stems and leaves are harvested with accumulating plants and removed from the site. Phyto-extraction can be divided into two categories: continuous and induced (Anderson et al., 1993). Continuous phytoextraction requires the use of plants that accumulate particularly high levels of the toxic contaminants throughout their life time (hyperaccumulators), while induced phytoextraction approaches enhance toxin accumulation at a single time point by addition of accelerants or chelators to the soil. In the case of heavy metals, chelators like EDTA assist in mobilization and subsequent accumulation of soil contaminants such as lead (Pb), cadmium (Cd), chromium(Cr), copper (Cu), nickel (Ni), and zinc (Zn) in Brassica juncea (Indian mustard) and Helianthus anuus (sunflower) (Prasad, 2011). The ability of other metal chelators such as CDTA, DTPA, EGTA, EDDHA, and NTA to enhance metal accumulation has also been assessed in various plant species (Aksoy et al., 2000). However, there may be risks associated with using certain chelators considering the high water solubility of some chelator toxin complexes which could result in movement of the complexes to deeper soil layers (Baker and Brooks, 1989) and potential ground water and estuarial contamination. 1.1.2. Phytodegradation Metabolism of contaminants within plant tissues are known as Phytodegradation. Phytodegradation, also called phyto-transformation, refers to the uptake of contaminants with the subsequent breakdown, mineralization, or metabolization by the plant itself through various internal enzymatic reactions and metabolic processes. Depending on factors such as the
concentration and composition, plant species, and soil conditions, contaminants may be able to pass through the rhizosphere only partially or negligibly impeded by phyto-sequestration and/or rhizodegradation. In this case, the contaminant may then be subject to biological processes occurring within the plant itself, assuming it is dissolved in the transpiration stream and can be phyto-extracted (Prasad, 2011). Plants catalyze several internal reactions by producing enzymes with various activities and functions specifically; oxygenases have been identified in plants that are able to address hydrocarbons such as aliphatic and aromatic compounds. Similarly, nitro-reductases are produced in some plants that can reduce and breakdown energetic compounds such as the explosives trinitrotoluene (TNT), 1, 3, 5 trinitroperhydro -1, 3, 5- triazine (RDX) and 1,3,5,7 tetranitro 1,3,5,7 tetrazocine (HMX High melting explosive) (Anonymous, 2009). Many of the plant enzymes may even be able to metabolize or mineralize several chemicals completely to carbon dioxide and water (McCutcheon and Schnoor, 2003). 1.1.3. Phytostabilization Production of chemical compounds by plant, are used to immobilize contaminants at the interface of roots and soil. Phytostabilization involves the establishment of a plant cover on the surface of the contaminated sites with the aim of reducing the mobility of contaminants within the vadose zone through accumulation by roots or immobilization within the rhizosphere, thereby reducing off-site contamination. The process includes transpiration and root growth that immobilizes contaminants by reducing leaching, controlling erosion, creating an aerobic environment in the root zone and adding organic matter to the substrate that binds the contaminant. Microbial activity associated with the plant roots may accelerate the degradation of organic contaminants such as pesticides and hydrocarbons to nontoxic forms. Phytostabilization can be enhanced by using soil amendments that immobilize metals combined with plant species that are tolerant of high levels of contaminants and low-fertility soils or tailings. Soil amendments used to enhance immobilization may need to be periodically reapplied to maintain their effectiveness (Nanthi et al., 2011). 1.1.4. Phytovolatilization Plants can also remove toxic substances, such as organics, from the soil through phyto-volatization. In this process, the soluble contaminants are taken up with water by the roots, transported to the leaves, and volatized into the atmosphere through the stomata.
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International Journal of Scientific Research in Environmental Sciences, 2 (4), pp. 120-138, 2014
The best example of this is the volatilization of mercury (Hg) by conversion to the elemental form in transgenic Arabidopsis and yellow poplars containing bacterial mercuric reductase. 1.1.5. Rhizofiltration Roots of plants are capable to decontaminate polluted waters and sewage by absorption or uptake of heavy metals. 1.2. Metals toxicity and tolerance in plants The effects of heavy metals on plants are related to physiological processes (Sharma and Agarwal, 2005). Because it can be easily measured, plant growth is commonly used as a general parameter to study the influence of stressors, with growth rate inhibition often being the most obvious plant reaction (Fodor, 2002; Hagemeyer, 2004). This is especially true of the root system, which is the first plant system to come into direct contact with toxic ions. Leaf chlorosis, disturbed water balance and reduced stomatal opening are characteristic effects of toxic Ni concentrations (Clemens, 2006), but they are also caused by many heavy metals and even occur more generally as a stress response. Fodor (2002) suggested an interesting stepwise model for the action of heavy metals in plants. Initially, there are interactions with other ionic components present at the locus of entry into the plant rhizosphere that subsequently have consequences for the metabolism. This is followed by an impact on the formation of reactive oxygen species (ROS) in the cell wall and an influence on the plasmalemma membrane system in stage I. At stage II, the metal ion reacts with all possible interaction partners within the cytoplasm, including proteins, other macromolecules and metabolites. Stage III is mainly related to the factors that influence homeostatic events, including water uptake, transport and transpiration. At this stage, symptoms start to develop, and they become visible at stage IV according to Fodors model. As an example, the chlorophyll and, usually to a smaller degree, carotenoid contents decrease, which have obvious consequences for photosynthesis and plant growth (Barcelo and Poschenrieder, 2004). The death of the plant cell occurs at stage V. This model has the advantage that visible effects are linked to metabolic events that are influenced by the metal ion of interest. The symptoms of metal toxicity are similar and the most investigated heavy metal, without any question, is cadmium. The most important effects of heavy metals/metalloids are as follows:
1- Oxidative stress because of oxidativeredox properties of many heavy metals and metalloids (DeVos et al., 1992; Supalkova et al., 2007) 2- Bonding of heavy metals/metalloids with the structures of proteins and other bioactive compounds because of their similarity to essential metals .Plants response to the presence of heavy metal/metalloid ions includes synthesis of plant thiol compounds, namely phytochelatins (Adam et al., 2005; Klejdus et al., 2004; Petrlova et al., 2006; Potesil et al., 2005; Rauser 1995; Supalkova et al., 2007, 2008; Vatamaniuk et al., 2000; Zehnalek et al., 2004a, b; Zitka et al., 2007). Some plants, called metallophytes, demonstrate tolerancehypertolerance to heavy metals/metalloids and in addition to hyperaccumulation of one or more metals/metalloids. These plants may have two important economic possibilities: phytomining-heavy metal extraction; phytoremediation that is metal accumulation from soil in plants. Definite mechanisms of hyper tolerance of metal are still unknown, but some genes, especially for metal homeostasis, and stress genes were identified to be responsible (Weber et al. 2004). To treat environmental problems, phytoremediation technologies can be used. The main advantage is that the cost of the phytoremediation is lower than that of traditional processes, both in situ and ex situ. In the case of remediation of environment polluted by organic compounds, they can be degraded. Moreover, there is also the possibility for the recovery and reuse of valuable metals. 1.3. Advantages phytoremediation and limitations of
Phytoremediation, has been reported to be an effective, in situ, non-intrusive, low-cost, aesthetically pleasing, ecologically benign, socially accepted technology to remediate polluted soils (Alkorta and Garbisu, 2001; Garbisu et al., 2002). It also helps prevent landscape destruction and enhances activity and diversity of soil microorganisms to maintain healthy ecosystems, which is consequently considered to be a more attractive alternative than methods that are currently in use for dealing with heavy metal contamination (Cunningham and Berti, 1993; Salt et al., 1995). Phytoremediation takes advantage of the unique, selective and naturally occurring uptake capabilities of plant root systems, together with the translocation, bioaccumulation and pollutant storage/ degradation abilities of the entire plant body. Several plant species both terrestrial and aquatic possess substantial hyper-accumulating power that can be used for phytoremediation from soil and water.
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Phytoremediation is regarded as the cheapest and environmentally most friendly technology for cleaning up soil. The most widespread and most profitable technique is phytoextraction, used mainly for removing heavy metals and radioactive elements from the soil (Pulford and Watson, 2003). Initially, the scope of phytoremediation was limited, principally because of the low bioavailability of heavy metals and the low biomasses of plants. Moreover, the management of the plant matter obtained after phytoremediation was troublesome ((Pulford and Watson, 2003; Nouairi et al., 2006; Zhao and McGrath, 2009). There are many ways of improving the efficiency of this process, however. To enhance the accumulative potential of plants chelates can be used: these compounds substantially intensify the uptake and translocation of metals in plants in that they release metals from the soil and form soluble complexes with them, which are then transported by the xylem and deposited in the leaves. Uptake efficiency depends on the metals affinity for the chelate. The mobility of heavy metals in the soil can also be manipulated by altering its pH: a higher pH > 6.5 significantly reduces the quantity of readily soluble forms of metals in the soil and limits their uptake and accumulation by plants. By contrast, plants in an acidic environment can take up large amounts of these metals, even from soils that are only moderately polluted (Pulford and Watson, 2003).The success of
No. 1 2 3 4 5 6 Advantages Applicable to both inorganic and organic contaminants It can be applied in situ. Reduces the amount of waste going to landfills. Does not require expensive equipment or highly specialized personnel. Phytoremediation is cheaper than conventional remediation methods. Easy to implement and maintain. Plants are a cheap and renewable resource, easily available. Environmentally friendly, socially accepted. Less noisy than other remediation methods.
phytoremediation depends mainly on the choice of plant, which must obviously possess the ability to accumulate large amounts of heavy metals (hyperaccumulation). These plants also have to satisfy other criteria as: 1- The concentration of heavy metals in the shoots should be 50 to 100 times greater than in normal plants (Jabeen et al., 2009). 2- The bioaccumulation coefficient (the ratio of the concentration of a toxic substance in the tissues of an organism to its concentration in the living environment of that organism) must have a value greater than 1 (McGrath and Zhao, 2003). 3- Metal concentrations in the shoots should be higher than in the roots (Jabeen et al., 2009) 4- Fast growth and high accumulating biomass (Marchiol et al., 2004). 5- Easily grown as an agricultural crop and fully harvestable (Marchiol et al., 2004). The advantages and limitation of phytoremediation technology depends upon the factors as type and species of plant and metal nature against the subjected plant, the main advantages and limitation of phytoremediation technology are reported by number of the researchers (Pilon-Smits, 2005; Alkorta et al., 2004; Gardea-Torresdey et al., 2005; Evangelou et al., 2007). Table 1 shows the advantages and limitation of the technology.
Table 1: Advantages and limitations of phytoremediation technology

Limitations Not accessing elements below the root depth. Management of plant matter after phytoremediation. Low bioma. The bioavailability of the pollutants. Restricted to sites with low contaminant concentration. Introduction of inappropriate or invasive plant species should be avoided (non-native species may affect biodiversity). High concentrations of hazardous materials can be toxic to plans. Possibility for contaminants to enter food chain through animal and plant consumption.
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2. FAMILY BRASSICACEAE BURNETT The family Brassicaceae (Cruciferae) known as a mustard family. Brassicaceae together with additional 15 families classified under order-Brassicales. Other significant families are Capparaceae, which is sometimes included in Brassicaceae as sub family Capparoideae, Tropaeolaceae and Caricaceae, well known for edible fruits- papaya with interesting pharmacological properties (Breithaupt et al., 2003;
Nayak et al., 2007; Okeniyi et al., 2007; Amazu et al., 2009; Abdullah et al., 2011). Brassicaceae (mustard family) consists of usually hermaphroditic herbs, sometimes shrubs, with simple lobbed or divided, spiral, exstipulate leaves. Flowers are arranged in inflorescence represented usually by a raceme. Flowers are bisexual, usually actinomorphic, perianth is dichlamydeous, cruciate (cross-shaped). Androecium consists of six stamens (two outer are shorter than four inner), Gynoecium is syncarpous,
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consisting of two carpels. The fruit is a silique or silicle (Ehrendorfer et al., 2006). Till date, classification of Brassicaceae (338 genera with 3,709 species) is very complicated, still under the debate among taxonomists due to its typical morphology and anatomy (Crespo et al., 2000; Marhold et al., 2004; Johnston et al., 2005; Bremer et al., 2009; Haston et al., 2009; Huang et al., 2010; Qiu et al., 2010; AlShehbaz et al., 2006; Bailey et al., 2006; Koch and Mummenhoff, 2006; German et al., 2009; Endress, 2011; Ge et al., 2011). Brassicaceae has a worldwide distribution. There are many species of economical value, for example vegetable plants, plants used for obtaining of dyes, oil, etc. There are also many plants having ornamental value. The best known member of this family is Arabidopsis thaliana, plant noted as a model for molecular biology. Brassicaceae contains high number of species that are able of hyper-accumulation of heavy metals, especially nickel (Thlaspi and Alyssum), cadmium and zinc (Thlaspi caerulescens, Thlaspi praecox, Thlaspi goesingense and Arabidopsis halleri). About 25% of all known hyper-accumulators are members of this family (Rascio and Navari-Izzo, 2011; Marques et al., 2004; Doran and Nedelkoska, 1999). The members of the family Brassicaceae are prominent for their ability in accumulating the heavy metals in an extremely high degree (Broadley et al., 2001). Due to this fact, significant attention has been paid to the members of the Brassicaceae, which are well-known as plants having significant biomass and capacity to accumulate high quantities of heavy metals (Kumar et al., 1995; Dushenkov et al., 1995). It was found that plants with higher growth rates from the family Brassicaceae in their ability to tolerate and accumulate the metals, including Brassica juncea (L.) Czern., Brassica nigra Koch, Brassica campestris L., Brassica napus L., and Brassica oleracea L. (Kumar et al., 1995). Despite the fact, that all of the examined crops from the Brassicaceae do accumulate the metals, B. juncea shows the highest ability to accumulate and transport Cu, Cr, Cd, Ni, Pb, and Zn towards their stems. However, all of the species of the family Brassicaceae show equal ability to accumulate and transport the heavy metals towards their stems (Boye, 2002). It was shown that B. juncea indicates excellent ability in eliminating Pb from the soil solution and to accumulating (Liu et al., 2000: Clemente et al., 2005). In contrary, based on several research findings (Salt and Kramer, 2000; Kumar et al., 1995; Blaylock et al., 1997) it was possible to conclude that B. juncea (L.) Czern., possesses remarkable capacity to absorb and accumulate significant quantities of heavy metals, as Cd, Cu, Ni, Zn, Pb and Se and has a very high potential for phytoremediation of heavy metals
(Blaylock et al., 1997; Blaylock and Huang ,2000; Ebbs and Kochian, 1998; Haag-Kerwer et al., 1999). The available statistics indicate that B. juncea (L.) Czern., has an ability of removing 1.1550 kg Pb from an acre (Henry, 2000). Certain crops like B. oleracea L., are able to accumulate significant quantities of heavy metals (Kabata-Pendias, 2001) which may lead to a risk for the health of the people. Brassica is the most economically important genus in the Brassicaceae. Several species and types of Brassicas are significant oilseed crops, vegetables, forage crops, and are used in the production of condiments, such as mustard. Brassica species are widely used in the cuisine of many cultures and recognized as a valuable source of dietary fiber. Brassica vegetables contain little fat, and are sources of vitamins, minerals, and fiber. They also contain a large number of novel phytochemicals, some of which protect against carcinogenesis (Steinmetz and Potter, 1996). Hence, Brassicas are believed to be useful in the prevention of cancer. Among the Brassica crops, oilseeds have the highest economic value. The oilseed Brassicas are found within Brassica juncea, Brassica carinata, Brassica rapa (syn. Brassica campestris) and Brassica napus collectively, and are commonly called oilseed rape. When Brassica oils are low in aliphatic glucosinolates and erucic acid, the varieties are increasingly commonly referred to as canola, a more pleasant-sounding name. Canola, which is most often B. napus, has received much attention worldwide and may soon be the most popular oilseed crop. There are now also canola-quality B. rapa and B. juncea varieties. Canola oil is widely used in cooking since it is very low in saturated fat, making it appealing to health conscious consumers. Brassica nigra is mainly used as a mustard condiment in addition to oil. Vegetable Brassica are an important and highly diversified group of crops grown worldwide that belong mainly to the species Brassica oleracea, as well as B. rapa and B. napus. This group includes plants such as broccoli, Brussels sprouts, cabbage, cauliflower, collards, kale, kohlrabi, rutabaga, and turnip. While most research in Brassica crops has been performed on oilseed and vegetable biotypes, rapidcycling Brassica biotypes of various species have gained attention in recent years. These rapid cycling types were genetic selections having short life cycles of 20-60d and small sizes (Williams and Hill, 1986). Rapid cycling Brassicas are attractive model laboratory plants because of their small genome sizes, in some cases just 3 to 4 fold larger than Arabidopsis thaliana (Arumuganathan and Earle, 1991). Other desirable qualities are their high female fertility, rapid seed maturation, and absence of seed dormancy.
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In addition to Brassica crops and rapid-cycling models, the genus contains several weedy species, of which the most important is B. rapa. Weedy B. rapa is considered one of the two most important weeds in the world (the other is wild rice) that is closely related to and sexually compatible with prominent row crops (Holm et al., 1997). Therefore, there is considerable interest in better understanding its genetics and genomics because of concern of introgression from transgenic canola to weedy biotypes (Halfhill et al., 2003; Stewart et al., 2003). B. rapa is also receiving attention as an important candidate for those interested in weed genomics to better understand the molecular basis of weediness as a syndrome (Basu et al., 2004). Considerable research has been conducted in tissue culture, transformation and molecular breeding of the Brassicas. Transformation in Brassica has been reviewed in detailed by researchers (Earle et al., 1996; Earle and Knauf, 1999). While information is available on genetic transformation in Brassica, there is no review on the other aspects of recent progress in cellular and molecular biology of the genus. In this review, we present pertinent research with an eye towards improving Brassica biotechnology. This includes organogenesis, somatic embryogenesis, microspore culture and doubled haploids, somatic cell fusion, molecular markers for genetic fidelity of in vitro-grown plants, marker-assisted selection, and transformation. We have reviewed transformation only with respect to desirable traits engineered since all the other aspects of transformation have been recently covered as indicated above. 2.1. Role of Brassicaceae in phytoremediation The continuous application of large amounts of fertilizers and other soil amendments to agricultural land has raised concern regarding the possible accumulation of elevated levels of their trace element constituents and potential harm to the environment (Colbourn and Thornton, 1978; Ma and Rao, 1997; Raven and Leoppert, 1997). Furthermore, increasing amounts of urban and industrial wastes (Haines and Pocock, 1980; Parry et al., 1981; Culbard et al., 1983; Gibson and Farmer, 1983) which may contain significant quantities of heavy metals are being disposed on the agricultural lands (Raven and Leoppert, 1997). Severe heavy metal contamination in soil may cause a variety of problems, including the reduction of yield and metal toxicity of plant, animals and humans. The decontamination of these soils by engineering methods is high costing project (Baker et al., 1991; Salt et al., 1995). Over the couple of decades there has been an increasing interest in developing a plant based technology to remediate
heavy metal contaminated soils (Chaney, 1983; Cunningham and Berti, 1993; Baker et al., 1994; Raskin et al., 1994). Phyto extraction is the use of plants to remove heavy metals from contaminated soils. The concept of using plants to clean up contaminated environments is not new. About 300 years ago, plants were proposed for use in the treatment of waste water. At the end of the 19th century, Thlaspi caerulescens and Viola calaminaria were the first plant species documented to accumulate high levels of metals in leaves (Hartman, 1975). The idea of using plants to extract metals from contaminated soil was reintroduced and developed by Utsunamyia (1980) and Chaney (1983), and the first field trial on Zn and Cd phytoextraction was conducted by Baker et al., (1991). Since plant cultivation and harvesting are relatively inexpensive processes as compared to traditional engineering practices that rely on intensive soil manipulation. Phyto-extraction may provide an attractive alternative for the clean up of heavy metal contaminated soils. The goal of heavy metal phytoextraction is to reduce metal levels in the soil up to the acceptable levels with in a reasonable time frame (Raskin et al., 1994; Nanda-Kumar et al., 1995; Huang et al., 1997). The process of phytoextraction generally requires the translocation of heavy metals to the easily harvestable shoots. A few plant species are able to survive and reproduce on soils heavily contaminated with Zn, Cu, Pb, Cd, Ni, Cr and As. Such species are divided into two main groups. The first group called pseudo metallophytes, which grow on both contaminated and non-contaminated soils, and second group called as absolute metallophytes, that grow only on metalcontaminated and naturally metal rich soil (Baker, 1987). Depending on plant species, metal tolerance may result from two basic strategies: metal exclusion and metal accumulation (Baker, 1981; Baker and Walker, 1990). The exclusion strategy, comprising avoidance of metal uptake and restriction of metal transport to the shoots (De Vos et al., 1991), is usually used by pseudo-metallophytes. The accumulation strategy caused high uptake of metal and storage in vacuoles to prevent metal toxicity. The extreme level of metal tolerance in vascular plants is called hyper-accumulation. Hyper-accumulators are defined as higher plant species whose shoots contain 100mg Cd Kg-1,1000mg Ni, Pb and Cu Kg-1 or 10,000mg Zn and Mn Kg-1 (dry weight) when grown in metal-rich soils (Baker and Brooks, 1989; Baker et al. 2000). Crops with both a high metal uptake capacity and a high biomass production are needed to extract metals from soils with in a reasonable time frame (Ebbs and Kochian, 1997). According to Brooks et al. (1977), metal hyper accumulation is a rare phenomenon that occurs in some plants called
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hyper accumulators. However, hyper-accumulators are often described as slow growing and low biomass plants (Dushenkov et al., 1995; Nanda-Kumar et al., 1995; Ebbs et al., 1997; Rouhi 1997). The potential of some crop plants from Brassicaceae for phytoremediation has been extensively studied (Baker et al., 1994; Brown et al., 1995; Dushenkov et al., 1995; Huang and Cunningham, 1996; Ebbs and Kochian, 1997; Ebbs et al., 1997) and it was demonstrated that some efficient shoot accumulators of the genus Brassica contained up to 3.5% on a dry weight basis of heavy metals (Nanda-Kumar et al., 1995). Over 400 plant species have been identified as natural metal hyper-accumulators representing about 0.2% of all angiosperms. Unfortunately, most of these plants are characterized by slow growth and limited biomass production. Because of these limitations such plants cannot be used to remove certain heavy metals from soil. For instance, Pb phytoremediation technology can only be feasible if systems can be developed to employ high biomass plants, which are capable of accumulating more than 1% Pb in shoots and produce more than 20 t of biomass ha1 yr1 (McGrath and Zhao, 2003).Based on the review of literature it can be stated that the most frequently cited species in phytoremediation studies are Brassica juncea (L.) Czern., followed by Helianthus annuus L., Brassica napus L. and Zea mays L. The greater interest in Brassicaceae derives from the fact that research on these species started earlier, together with the interesting concentrations they provide, especially for Brassica juncea (L.) Czern. (Vamerali et al., 2010). Among the plants of the Brassica species, the Brassica juneca deserve special attention because its relevance to the process of phytoexctration of heavy metals from soil was confirmed in many experiments. It has been found that B. juncea exhibits a high capacity to accumulate Cd- mainly in the shoots, where Cd level was recorded at level of 1450 g Cd/g dry wt. This is three times more than reported Brassica napus (555 g/g dry wt) (Nouairi et al., 2006). In addition, this plant exhibit a high removal efficiency of other metals such as Pb (28% reduction) and Se (reduced between 1348%; Salt et al., 1998). In addition, this plant is more effective at removing Zn from soil than Thlaspi caerulescens, a known hyper-accumulator of zinc. This is due to the fact, that B. juneca produces ten-times more biomass than Thlaspi cearullescens (Gisbert et al., 2006). However, Brassica juneca (L.) Czern., needs to be harvested shortly after the plant becomes mature, which causes problems of disposal of obtained biomass. When these plants are dried, they easily crumble and flake off, greatly reducing the yield obtained, and the rest of the plant residues are a source of secondary emissions of
toxic substances. Quite a large biomass and the lack of difficulties after harvesting are advantages described for different types of cabbage. In the case of Chinese cabbage, the high cumulative capacity of lead was observed within the limits of 5010 to 4,620 mg/kg dry wt. During testing capacity of phyoextraction of Zn, Cu and Pb for three Brassica crop species: B. oleracea L., B. carinata A. Br. and B. juneca (L.) Czern., the highest concentration of Zn (381 mg/kg dry wt.) and Cu (8,34 mg/kg dry wt.) were recorded in the shoots of B. oleracea L. The Pb concentrations of all Brassica species were more or less constant over the tested range of soil Pb concentrations, with lower values than the other metals. The low bioaccumulation of lead is due to its extreme insolubility and not generally being available for plant uptake in the normal range of soil pH (Gisbert et al., 2006). The high potential of plants from the family Brassicaceae exhibit for bioaccumulation of heavy metals along with management of plant matter after phytoremediation process means that phytoremediation could become one of the most important technologies for cleaning the components from the environment. In recent years, interest in natural methods of plant protection against various pests has grown. 2.2. Metal tolerance in the Brassicaceae The 93 documented species of metal-hyperaccumulating Brassicaceae provide substantial opportunity to study the physiological and genetic mechanisms behind metal tolerance and hyperaccumulation as well as the ecological implications of these mechanisms. Some of the most well-studied genera of hyperaccumulators in this family include Arabidopsis, Brassica, Alyssum, Noccaea (formerly Thlaspi), Stanleya, and Streptanthus (Bhargava et al., 2012; Boyd et al., 2009; Freeman et al., 2010; Vamerali et al., 2010; Verbruggen et al., 2009). The most studied metal- hyperaccumulating taxa of Brassicaceae are Noccaea caerulescens (formerly Thlaspi caerulescens) is, perhaps, the most wellstudied metal-hyperaccumulator (Milner and Kochian, 2008), accumulating up to 36,900 g g-1 Zn and 1800 g g-1 Cd without signs of toxicity (Bhargava et al., 2012). Because, N. caerulescens, like most other model taxa in the Brassicaceae-grows easily in the lab, it has been extensively studied, revealing several mechanisms for metal uptake, transport, and localization (Cosio et al., 2004). However, its small biomass limits its potential as a candidate for phytoremediation (Bhargava et al., 2012). Arabidopsis thaliana, although not a naturally metalaccumulating species, is a popular model organism for plant-based research (Bevan and Walsh, 2005).
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Arabidopsis thaliana's genome is mapped (Weigel and Mott, 2009) and its sequence is very similar to its metal- accumulating congener A. halleri (Becher et al., 2004; Meyer and Verbruggen, 2012; Weber et al., 2004). For this reason, both A. thaliana and A. halleri are commonly used to study the genetic basis for metal tolerance and hyperaccumulation (Bevan and Walsh, 2005; Cho et al., 2003; Chaffai and Koyama, 2011; Courbot et al., 2007; Hanikenne et al., 2008). A common condiment crop in North America and Europe, Brassica juncea (Indian mustard) is a popular choice for phytoremediation (Lim et al., 2004; Neilson and Rajakaruna, 2012). Although not a hyperaccumulator, with the ability to accumulate Cd, Zn, Se, and Pb and a biomass at least 10-fold greater than that of N. caerulescens, B. juncea has been used with success in several phytoremediation studies and trials (Bhargava et al., 2012; Szczyglowska et al., 2011; Warwick, 2011). The molecular mechanisms responsible for selenium (Se) tolerance and hyperaccumulation have been investigated in the Se hyperaccumulator Stanleya pinnata by comparing it with its Se-tolerant congener, S. albescens, using a combination of physiological, structural, genomic, and biochemical approaches (Freeman et al., 2010). Additionally, the ecological functions and implications of Se hyperaccumulation in Stanleya and other plants (El Mehdawi and PilonSmits, 2012) and the potential for Se phytoremediation have also been investigated (Banuelos, 2001). 2.3. Metal tolerance in the Brassica Many hyper-accumulators belong to the Brassica family. Once it was suspected that known hyperaccumulators were not suited for phytoextraction, researchers looked to other high biomass accumulating members of the Brassicaceae for plants which accumulated large quantities of toxic metals (Dushenkov et al., 1995; Kumar et al., 1995). Kumar et al. (1995) tested many fast growing Brassicas for their ability to tolerate and accumulate metals, including Indian mustard (B. juncea), black mustard (Brassica nigra Koch), turnip (Brassica campestris L.), rape (Brassica napus L.), and kale (Brassica oleracea L). Although all Brassicas accumulated metal, B. juncea showed a strong ability to accumulate and translocate Cu, Cr VI, Cd, Ni, Pb, and Zn to the shoots. Kumar et al. (1995) also investigated possible genetic variation of different B. juncea accessions in hope of finding some that had more phytoextraction potential than others (Table 2). Indian mustard is an oilseed Brassica crop for which cultivation extends from India through western Egypt and Central Asia to Europe (Nishi, 1980).
According to Prakash (1980), the oldest reference to B. juncea in Sanskrit literature is by the name Rajika, and carbonized seeds of this species have been found in the ancient sites of the Harappan civilization (2300-1750 B.C.). Despite the efforts of historians and researchers, the precise origin of this crop remains an enigma. Perhaps the most likely place or places of origin are those regions where its parents, B. nigra and B. campestris, overlap in their distribution. Possible centers of origin include Africa (Zeven and Zhukovsky, 1975), China (Chen et al., 1995), the Middle East, Southwest Asia, and India (Sauer, 1993). Indian mustard is eaten as a leafy vegetable in China but is grown in India primarily for its oilcontaining seeds (Prakash, 1980), which serve as a source of cooking oil and spice (Nishi, 1980; Krzymaski, 1997). Indian mustard is capable of producing 18 tons of biomass per hectare per crop (Kumar et al., 1995). Plants perform very well in nutrient solution culture, progressing from the fourleaf stage to fully grown plants (up to 50 g shoot fresh mass) in as little as 21 days (personal observations). Although short-day conditions (<12 hrs light) result in a more compact growth, shorter height, and limited leaf production (Bhaskar and Vora, 1994), biomass accumulation is greater than under long-day conditions that is 9 to 10 hrs light optimal (Neelam et al., 1994). Long-day conditions promote early flowering (Bhaskar and Vora, 1994) but are not required for flower development. These plants have indeterminate growth and continue to branch from the nodes and to accumulate biomass after flower and siliquae (seed pod) development. The recommended fertility rate for maximum growth of B. juncea under un-contaminated conditions is 75 to120 kg N ha-1 and 30 to 50 kg P2O5 per hectare (Gurjar and Chauhan, 1997; Thakral et al., 1995; Tomar et al., 1997). Zaurov et al., (1999) reported that biomass accumulation of B. juncea was greatest when plants in soil are supplied with 200 kg N, 100 kg P2O5, and 66 kg K2O per hectare. However, Cd concentration in the tissue was greatest when no N was supplied. Indian mustard is given considerable attention by present day researchers, geneticists, and plant breeders in particular, because of its unique polyploid genome. Brassica juncea is an allotetraploid, a plant with a genome composed of the complete diploid genomes of parents, B. nigra and B. campestris. In modern breeding programs, selection of B. juncea is based on a wide variety of characters. Improving oil and meal quality by eliminating nutritionally undesirable erucic acid or by modifying the fatty-acid composition of oil is an important objective for some plant breeders (Banga, 1997). Other selections are based on insect (diamondback moth), Plutella xylostella L.
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(Andrahennadi and Gillott, 1998) and disease resistance (blackleg), Leptosphaeria maculans (Desm.) Ces.; Pang and Halloran, 1996) and various temperature adaptations (Banga, 1997). Only recently has there been an interest in selecting Indian mustard lines based on their ability to tolerate and accumulate heavy metals. Several accessions of B. juncea have been identified as moderate accumulators of metallic elements and are maintained by the USDA-ARS Plant Introduction Station at Iowa State University, Ames, Iowa. The benefit of using B. juncea seed from the plant introduction station is that the genetic integrity
of the accessions is preserved through appropriate breeding techniques. Experiments that utilize these seeds have more precision than those conducted with seeds from commercially available sources. Precision is also greater, because future researchers can obtain the same accessions for their experiments. The USDA-ARS Plant Introduction Station maintains a world-wide collection of B. juncea accessions that are known metal-accumulators, and the seeds are distributed to public and private research institutions at no cost.
Table 2: Promising species of Brassica for the phytoextraction of various metals

No. 1 2 3 4 Metal Cd Cr (VI) 137 Cs Cu Plant species Brassica juncea (L.) Czern. Brassica juncea (L.) Czern. Brassica juncea (L.) Czern. Brassica oleracea L. Brassica juncea (L.) Czern. Reference Kumar et al., 1995 Kumar et al., 1995 Lasat et al., 1998; Lasat et al., 1997 Lasat et al., 1997 Kumar et al., 1995 Pantola et al., 2013 Purakayastha et al., 2008 Kumar et al.,1995 Purakayastha et al., 2008 Kumar et al., 1995 Kumar et al., 1995 Kumar et al., 1995; Blaylock et al., 1997; Begonia et al., 1998 Kumar et al., 1995; Herrero et al., 2003 Kumar et al.,1995 Banuelos et al., 1997 Huang et al., 1998 Huang et al., 1997 Huang et al., 1997 Ebbs and Kochian, 1998; Kumar et al., 1995; Salt et al., 1995; Ebbs and Kochian, 1997; Purakayastha et al., 2008 Ebbs and Kochian, 1997; Herrero et al., 2003. Ebbs and Kochian, 1997
5 6
Ni Pb
Brassica juncea (L.) Czern. Brassica campetris L. Brassica carinata A. Br. Brassica juncea (L.) Czern. Brassica napus L. Brassica nigra (L.) Koch. Brassica napus L. Brassica chinensis L. Brassica juncea (L.) Czern. Brassica narinosa L. Brassica juncea (L.) Czern.
7 8
Se U
9 Zn
Brassica napus L. Brassica rapa L.
2.4. Role of chelators for accumulation of metals in Brassica There are different strategies for enhancement of accumulation properties of members of the Brassicaceae. The first approach consists in supplementation of soil by additives, compounds that are able to increase uptake of heavy metals. There are many compounds, which were tested of possible chelators of heavy metals. However, their usage has many disadvantages including potential toxicity with reduction of biomass production and hazard for the environment due to mobilization of heavy metals. On the other hand, heavy-metals accumulating plants may
represent potential risk because of their possible entry the food chain (Table 3). The second approach is based on construction of transgenic plants. This method is based on an introducing of foreign genes, which are connected with uptake, transport and accumulation of heavy metals. A yeast cadmium factor 1 (YCF1), a member of the ATP-binding cassette (ABC) transporters, is localized at the vacuolar membrane in Saccharomyces cerevisiae. YCF1 gene introduced to the transgenic Brassica juncea plants showed 1.3 to 1.6 fold tolerances to cadmium ions compared to wild plants (Bhuiyan et al., 2011).
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Table 3: Chelators used in members of the Brassicaceae for phytoremediation

No. 1 2 3 4 5 6 7 8 9 10 Chelator EDTA, EDDS, histidine EDTA EDTA, DTPA Oxalic acid, citric acid EDTA Nitrilotriacetate Citric acid [S,S]-ethylenediamine disuccinate Thiol-rich chelators EDTA, EDDS, DTPA NH4Cl, casein, citric acid EDTA Heavy metals Cu, Pg, Fe, Zn Cd, Cr, Ni Cr, Ni Cu, Cd, Pb, Zn Cd Pb) As, Hg Cu Cd, Cu, Ni, Zn Pb Plant species Brassica juncea Thlaspi caerulascens Brassica juncea Brassica napus Brassica juncea Brassica juncea Thlaspi goesingense Arabidopsis halleri Brassica rapa v. pekinensis Brassica chinensis Brassica juncea Reference Karczewska et al., 2009 Munn et al., 2008 Hsiao et al., 2007 Turan and Bringu, 2007 Quartacci et al., 2005 Finzgar et al., 2005, 2006 Meagher and Heaton, 2005 Kos and Lestan, 2004 Gramss et al., 2004 Blaylock et al., 1997
3. TRANSLOCATION OF METAL IN PLANTS Metal uptake by plants is regulated by the electrochemical potential gradient for each metal ion that exists across the plasma membrane of root cells (Welch, 1995). The effects of metals on the rate of movement and composition of the xylem and phloem sap may impact on plant response to metal toxicity. Translocation effects include the relative proportions of metals in roots vs. shoots, potential sites of toxic action of metals and the translocation of other nutrients within the plant. Within a plant the two major transport mechanisms for metals are via the xylem and phloem. Transport of metal ions within the xylem is essentially driven by mass upward flow of water created by the transpiration stream (Kochian, 1991; Welch, 1995). Water transpiration rate has a large effect on macronutrient translocation rate, however at low supply, processes such as xylem loading and unloading and transfer between xylem and phloem have been shown to be more important for the rate of nutrient supply (Welch, 1995). There is little to suggest that the case would be different for metals, and hence, under conditions of excess metal it is likely that the rate of transpiration would dominate metal movement in the xylem sap. Transport of metals within the phloem is thought to occur via the positive hydrostatic pressure gradient developed from the loading of sucrose into the phloem from mature actively photosynthesizing leaves and unloading of sucrose into the sink tissues such as rapidly growing tissues, apical root zones and reproductive organs (MacRobbie, 1971; Hocking, 1980; Welch, 1995). As in the xylem, the pH, redox potential, ionic strength and organic constituents of the phloem sap will determine the loading, transport and unloading of metals in the phloem (Welch, 1995).
However, unlike xylem cells, phloem cells are alive and metabolically active. Hence, metabolic reactions within the phloem have the potential to make the phloem sap more responsive to changes in the internal plant environment than the xylem sap (Welch, 1995). The mode of phytoremediation, the effect of the metals on the plants, the ability of the plant to extract metals from soil (Bioconcentration Factor - BCF), and the ability of the plants to move the metals to the aerial parts of the plants (Translocation Factor - TF) can be evaluated for the different metals. To evaluate the potential of plants for phytoextraction, the translocation factor (TF) is used. This ratio is an indication of the ability of the plant to translocate metals from the roots to the aerial parts of the plant (Marchiol et al., 2004). The bioconcentration factor (BCF) and translocation factor (TF) are represented by the ratio as given below: BCF TF = =
[Metal g g -1 DW](root/shoot/leaf) [Metal mg L-1 ]nutrient solution [Metal g g -1 DW](shoot/leaf) [Metal g g -1 DW]root
4. CONCLUSION The Industrial revolution and anthropogenic activities are accountable for generating large amounts of nonbiodegradable compounds. These hazardous compounds cause public annoyance. These can be separated into organic and inorganic compounds, in the case of inorganic compounds, heavy metals, such as lead, zinc, cadmium, copper, nickel, chromium and radioactive elements, such as uranium are more usual. Most of them are released into the environment, causing severe destruction to the ecosystem and, being non-biodegradable, cause adverse effects on human
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health due to the unavoidable process of bioaccumulation. In most of cases, the only way to restore the environment is removing the pollutants. Hence, previous methods used to remediate the environment were based on the physico-chemical theory, like volatilization, verification, excavation, soil washing, soil incineration, chemical extraction, solidification, and landfills. Although, these methods are effective in numerous interventions, they are often expensive, invasive, and are not suitable for large-area application. In this context, the concept phytoextraction proposed by Chaney (1983) at first time, and subsequently rapidly sparking the interest of many researchers in order to make it a popular field. In the light of developments in scientific and technical knowledge it will be more appropriate, to establish the techno-commercial awareness among the people about de-bottle-necking of ongoing methodologies for enhancement of scavenging potential of the plants. The large scale field application can be achieved by developing the pre-defined protocols with advantages and limitations of method and plant species used for the remediation of heavy metal contaminated sites. Over the past decade, researchers have sought to perfect this remediation technology. The majority of phytoextraction research has focused on finding the ideal metal-accumulating plants and the means by which metals can be liberated from the soil for root uptake. At present, Indian mustard (B. juncea) along with few other potential members of family Brassicaceae considered among the most viable candidates for the phytoextraction of a number of metals including Cd, Cr (IV), Cs, Cu, Ni, Pb, U, and Zn. Proper plant nutrition has the potential to be an effective remedy for low cost agronomic practice to enhance the phytoextraction of heavy metals by plants of this particular family of flowering plants (Angiosperms). 5. ACKNOWLEDGEMENTS The authors are grateful to Prof. Aditya Shastri, Vice Chancellor, Banasthali Vidyapith and Prof. Vinay Sharma, Dean, Faculty of Science and Technology, Banasthali University, Rajasthan for their encouragement and support. REFERENCES Abdullah M, Chai PS, Loh CY, Chong MY, Quay HW, Vidyadaran S, Seman Z, Kandiah M, Seow HF (2011). Carica papaya increases regulatory T cells and reduces IFN-gamma(+)CD4(+) T cells in healthy human subjects. Mol. Nutr. Food Res., 55:803806.
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Ramesh Chandra Pantola is a Ph. D scholar in Department of Biosciences and Biotechnology, Banasthali University, Rajasthan, India. His current research is focuses on phytoremedial potentials of Brassica. To date, he has published several scientific articles in reputed journals related to pharmaceutical microbiology, stress biology and fate of heavy metals on plants.
Afroz Alam, Ph. D., has vast experience of teaching and research in various aspects of plant science including stress physiology. He has over 50 research publications in prestigious International and National Journals and a 4 text books with reputed publication houses to his credit. He is life member of various associations of Plant Sciences. Presently, he is working as Associate Professor in the Department of Bioscience and Biotechnology, Banasthali University, Rajasthan. He is currently supervising seven Ph. D. students in Plant Science.
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Potential of Brassicaceae Burnett (Mustard Family Angiosperms) in Phytoremediation of Heavy Metals

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Potential of Brassicaceae Burnett (Mustard Family Angiosperms) in Phytoremediation of Heavy Metals

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International Journal of Scientific Research in Environmental Sciences, 2 (4), pp. 120-138, 2014 Available online at http://www.ijsrpub.

com/ijsres ISSN: 2322-4983; 2014 IJSRPUB http://dx.doi.org/10.12983/ijsres-2014-p0120-0138

Table 1: Advantages and limitations of phytoremediation technology

Table 2: Promising species of Brassica for the phytoextraction of various metals

Brassica napus L. Brassica rapa L.

Table 3: Chelators used in members of the Brassicaceae for phytoremediation

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