Update

Research Focus

Transgenic plants for phytoremediation: helping

Phytoremediation is the use of plants to clean up From polluted soils to ‘toxic plants’
environmental pollution. However, detoxification of In comparison with other clean-up technologies,
organic pollutants by plants is often slow, leading to phytoremediation has potentially many advantages, in-
the accumulation of toxic compounds that could be later cluding low installation and maintenance costs, less dis-
released into the environment. A recent publication by ruption of the environment and other beneficial side
Doty and colleagues describes the development of trans- effects such as carbon sequestration and biofuel pro-
genic poplars (Populus) overexpressing a mammalian duction [5,6]. However, phytoremediation also suffers
cytochrome P450, a family of enzymes commonly from several limitations, among which the most commonly
involved in the metabolism of toxic compounds. The evoked are the slow rate of removal, incomplete metab-
engineered plants showed enhanced performance with olism and potential increase in bioavailability of toxic
regards to the metabolism of trichloroethylene and the contaminants [1,3]. Indeed, in the absence of significant
removal of a range of other toxic volatile organic pollu- detoxification, parent compounds and toxic metabolites
tants, including vinyl chloride, carbon tetrachloride, can accumulate inside plant tissues and eventually return
chloroform and benzene. This work suggests that trans- to the soil or volatilize into the atmosphere.
genic plants might be able to contribute to the wider and The recognition that plants can transform xenobiotic
safer application of phytoremediation. compounds emerged in the 1940s, when plants were shown
to metabolise pesticides [7]. Since then, the development of
genomics, proteomics and metabolomics has exposed the
Introduction: phytoremediation – plants to clean up plant metabolism of many xenobiotic compounds [1].
pollution Plants often use pathways and enzymes similar to those
Plants are autotrophic organisms capable of using sunlight of mammals, which led to the ‘green liver’ concept
and carbon dioxide as sources of energy and carbon. How- (Figure 2) [7]. However, being autotrophic organisms,
ever, plants rely on the root system to take up water and plants do not actually use organic compounds for their
other nutrients, such as nitrogen and minerals, from soil and energy and carbon metabolism. As a consequence, they
groundwater. As a side effect, plants also absorb a diversity usually lack the catabolic enzymes necessary to achieve
of natural and man-made toxic compounds for which they full mineralization of organic molecules, potentially result-
have developed diverse detoxification mechanisms [1]. Pol- ing in the accumulation of toxic metabolites [1]. Hence, the
lutant-degrading enzymes in plants probably originate from idea to enhance plant biodegradation by genetic transform-
natural defense systems against the variety of allelochem- ation was developed, following a strategy similar to that
icals released by competing organisms, including microbes, used to develop transgenic crops [1,5].
insects and other plants [2]. From this viewpoint, plants can
be seen as natural, solar-powered pump-and-treat systems Transgenic plants for phytoremediation
for cleaning up contaminated environments, leading to the Typically, transgenic plants exhibiting new or improved
concept of phytoremediation [3]. First developed for the phenotypes are engineered by the overexpression and/or
removal of heavy metals from soil, the technology has since introduction of genes from other organisms, such as bac-
proven to be efficient for the treatment of organic com- teria or mammals. Being heterotrophs, bacteria and mam-
pounds, including chlorinated solvents, polyaromatic hydro- mals possess the enzymatic machinery necessary to
carbons and explosives [3,4]. Beyond the removal of achieve a complete mineralization of organic molecules;
contaminants from soil, phytoremediation involves differ- bacterial and mammalian degradative enzymes can there-
ent processes, such as enzymatic degradation, that poten- fore be used to complement the metabolic capabilities of
tially lead to contaminant detoxification (Figure 1) [3–5]. plants [1]. Historically, transgenic plants for phytoreme-
However, despite great promise, rather slow removal rates diation were first developed in an effort to improve heavy
and potential accumulation of toxic compounds within metal tolerance; for example, tobacco plants (Nicotiana
plants might have limited the application of phytoremedia- tabacum) expressing a yeast metallothionein gene for
tion [1]. In a recent study, Doty et al. [6] have developed higher tolerance to cadmium, or Arabidopsis thaliana over-
transgenic poplars with an enhanced uptake and metab- expressing a mercuric ion reductase gene for higher toler-
olism of toxic volatile pollutants. In doing so, they have ance to mercury [8,9]. The first attempts to transform
delivered a technology that is likely to lead to the wider plants for phytoremediation of organic compounds tar-
application of phytoremediation in the field. geted explosives and halogenated organic compounds in
Corresponding author: Van Aken, B. ([email protected]). tobacco plants [10,11].
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Figure 1. Phytoremediation involves several processes: pollutants in soil and groundwater can be taken up inside plant tissues (phytoextraction) or adsorbed to the roots
(rhizofiltration); pollutants inside plant tissues can be transformed by plant enzymes (phytotransformation) or can volatilize into the atmosphere (phytovolatilization);
pollutants in soil can be degraded by microbes in the root zone (rhizosphere bioremediation) or incorporated in soil material (phytostabilization) [3–5].

However, although tobacco and A. thaliana are transfer to plants [12]. However, A. tumefaciens-mediated
good laboratory models, their small stature might not be transformation of forest trees is notoriously challenging,
suitable for field applications. Hence, there is particular which explains why there have been only a few reports
interest in the genetic transformation of poplar trees about the genetic modification of poplar plants [13]. Gull-
(Populus sp.), which are fast growing plants with high ner et al. [14] developed the first transgenic poplars for
biomass – ideal attributes for phytoremediation. Plant phytoremediation. Their transgenic line was designed to
transformation is usually performed using the ‘natural treat chloroacetanilide herbicides by the overexpression of
genetic engineer’ Agrobacterium tumefaciens, a plant a gamma-glutamylcysteine synthetase, an enzyme
pathogen that has become the favorite vector for gene involved in glutathione synthesis.

Figure 2. The three phases of the ‘green liver’ model. Hypothetical pathway representing the metabolism of trichoroethylene (TCE) in plant tissues. Phase I, activation of
TCE by oxidation to trichloroethanol; phase II, conjugation with a plant molecule; phase III, sequestration of the conjugate into the cell wall or within the vacuole [7].

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Poplar trees overexpressing a mammalian the first report about genetic engineering of plants for
cytochrome P450 phytoremediation applications, it constitutes a milestone
Cytochrome P450s constitute a large enzyme superfamily in the field for several reasons: first, it is one of the very few
commonly involved in the metabolism of toxic compounds. studies describing the successful development of trans-
In 2000, Doty et al. [11] described the development of genic poplars, which is technically challenging [13]; second,
transgenic tobacco plants expressing a human cytochrome the technology is efficient for the treatment of several
P450 and capable of metabolizing trichloroethylene (TCE) important organic pollutants likely to be found in mixture
640-fold faster than wild type plants. The same group later in the environment; and third, it constitutes the achieve-
reported the introduction of a rabbit cytochrome P450 in ment of a pioneer work initiated by the same group a
transgenic hairy root cultures of Atropa belladonna, which decade ago. With federal regulations limiting the use of
also exhibited a faster metabolism of TCE [15]. In the transgenic forest trees, further developments of phytore-
current study, Doty et al. [6] described the genetic trans- mediation are likely to involve genetic use restriction
formation of hybrid poplar plants (Populus tremula  technologies (GURTs) for controlling the dispersion of
Populus alba) overexpressing mammalian cytochrome transgenes in the environment [17]. As for transgenic
P450 2E1 (CYP2E1). The engineered trees were capable crops, risks inherent to genetically modified organisms
of the enhanced metabolism of five volatile toxic have to be minimized and balanced with the increasing
compounds: TCE, vinyl chloride, carbon tetrachloride, needs of an ever-expanding human population.
chloroform and benzene. Among the different transgenic
clones tested, the most efficient one, line 78, expressed
References
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toxic compounds would volatilize or otherwise contaminate
0167-7799/$ – see front matter ß 2008 Elsevier Ltd. All rights reserved.
the food chain. Although the study by Doty et al. [6] is not doi:10.1016/j.tibtech.2008.02.001 Available online 18 March 2008

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