Opinion
Plant nanotoxicology
Karl-Josef Dietz1 and Simone Herth2
1
Biochemistry and Physiology of Plants, Bielefeld University, 33615 Bielefeld, Germany
2
Thin Films and Nanostructures, Bielefeld University, 33615 Bielefeld, Germany
The anthropogenic release of nanoparticles (NPs) to the
environment poses a potential hazard to human health
and life. The interplay between NPs and biological pro-
cesses is receiving increasing attention. Plants expose
huge interfaces to the air and soil environment. Thus,
NPs are adsorbed to the plant surfaces, taken up through
nano- or micrometer-scale openings of plants and are
translocated within the plant body. Persistent NPs asso-
ciated with plants enter the human food chain. In this
Opinion, we document the occurrence and character of
NPs in the environment and evaluate the need for future
research on toxicological effects. Plant nanotoxicology is
introduced as a discipline that explores the effects and
toxicity mechanisms of NPs in plants, including transport,
surface interactions and material-specific responses.
Particulate matter
Particulate matter in the atmosphere is defined as parti-
cles between 0.1 and 10 mm in size. Approximately 25% of
all particulate matter is based on carbon and associated
with exhaust from all kinds of gas and coal engines [1]. The
total amount of particulate matter often varies between 20
and 30 mg/m3 in northern Europe [2] and daily values can
exceed 200 mg/m3 in some Asian cities [1], with 70% of the
major cities in the world registering values above 50 mg/m3
[3]. Scientific experts suggest a limit of 10 mg/m3 (no health
effects have been reported below this value), whereas
official guidelines favor a threshold between 20 and
25 mg/m3 [4]. Statistical data from more than 100 cities
in the USA [5] and Europe [6] suggest an increase in
mortality between 0.2% and 0.8% if particulate matter
increases by 10 mg/m3 [5]
Various metals are found in particulate fractions in
areas close to heavy metal industries, with abundant
vehicle traffic or along main wind directions [6,7]. Metals
that are particularly subject to biomonitoring include:
aluminium (Al), antimony (Sb), arsenic (As), lead (Pb),
cadmium (Cd), chromium (Cr), cobalt (Co), iron (Fe), copper
(Cu), molybdenum (Mo), nickel (Ni), mercury (Hg), vana-
dium (V) and zinc (Zn) as well as nitrogen (N) and sulfur
(S). These elements are divided into those most abundant
in the earth crust (Fe, Al und Cr), those characteristic for a
release by traffic (Sb, Cu and N), those in fertilizers or
pesticides (Cu, As and N), and those associated with hu-
man living (S). Platinum group elements [PGE; platinum
(Pt), palladium (Pd), ruthenium (Ru)] have not yet been
studied in detail, but will have to be added to the list in the
future, because their occurrence strongly depends on the
use of catalysts in vehicles [7]. The 1:5 ratio of Pt to Pd
Corresponding author: Dietz, K.-J. (karl-josef.dietz@uni-bielefeld.de).
582 1360-1385/$ – see front matter ß 2011 Elsevier Ltd. All rights reserved. doi:10.1016/j.tplants.2011.08.003 Trends in Plant Science, November 2011, Vol. 16, No. 11
found near streets is close to the ratio of these elements
used in catalysts and their concentrations decrease with
distance to the street [8–10]. Therefore, emission of PGEs
is mainly caused by traffic.
Much is known about the composition and location of
particulate matter, whereas it is difficult to get information
about the detailed size distribution. Particulate matter is
usually divided into particles with a 50% cut-off aerody-
namic diameter of less than 10 mm (PM10), less than
2.5 mm (PM2.5) and ultrafine particles (UP) less than
0.2 mm. Given that NPs are defined as particles with at
least two dimensions below 100 nm (i.e. 0.1 mm), only the
last category is interesting to the field of nanotoxicology.
NPs are generated during combustion processes and di-
verse naturally occurring abiotic and biotic chemical
processes. Their commercial production has increased
steadily, because they are used in many novel applications
(e.g. as catalysts, semiconductors, drug carriers and in
cosmetics) and the development of additional novel mate-
rials is likely. Thus, the particle load of the atmosphere,
aquatic environment, soil and, thus, of humans, plants and
animals will increase in the future [11]. Here, we discuss
the occurrence and character of NPs in the environment
and assess the need for future research on toxicological
effects, with specific emphasis on plants, including trans-
port processes, surface interactions and material-specific
responses. This important topic seems to be underexplored
at present if one considers the fact that plants represent by
far the largest interface between the environment and the
biosphere.
NP characteristics
Particles in the environment
The detailed size distribution and chemistry of particles in
the environment is difficult to examine, because their
concentrations are usually low. Fine particulate matter
is found in clean air at approximately 1 mg/m3 [12]. Alter-
natively, particles can be directly analyzed in the exhaust
of automotive engines [13]. Particle sizes of inorganic
materials, such as sulfate, ammonium, nitrate, chloride
and organic compounds range from 50 nm to 2 mm, with an
average size of approximately 100 nm [12]. Catalysts re-
lease small nanocrystalline Pt particles, which are at-
tached to aluminium oxide carrier particles with a
diameter that is generally bigger than 10 mm. It is also
assumed that a small fraction of Pt particles are oxidized,
therefore becoming soluble in water. In this state, the
particles are more hazardous to organic systems [14].
The maximum Pt concentration was found to be approxi-
mately 40 pg/m3 in urban sites with high traffic volume.
 Opinion Trends in Plant Science November 2011, Vol. 16, No. 11

For a medium traffic volume, metallic fractions are rarely
above the detection limits [15]. The amount of particulate
matter less than 100 nm in size is approximately 3–8% of
the total amount smaller than 2.5 mm; however, this
changes throughout the year, with a higher concentration
in the summer [12]. In the summer, 50% of all particles are
based on carbon (organic and elemental) and 50% are
based on inorganic matter (mainly sulfate and ammoni-
um), whereas in the winter, these numbers change to 70%
and 30%, respectively [12].
Particles in toxicological studies
Studied particles can be divided into carbon-based, oxides,
metals and semiconductors. In principle, the number of
publications dealing with these major groups correlates
with their occurrence in the environment. However, car-
bon-related studies often focus on carbon nanotubes and
fullerenes [16], which are rarely emitted to the environ-
ment, and only a few studies have addressed the amor-
phous and nanocrystalline carbon black formed as a
byproduct in engines [17]. The physical and chemical
characteristics of NPs relevant for toxicity to biosystems
are: (i) average size; (ii) element composition; (iii) surface
area; (iv) porosity; (v) surface charge; (vi) hydrodynamic
diameter; (vii) aggregation propensity [18], and also (viii)
stability and (ix) coating with cellular or other constitu-
ents. Nanoparticles tend to aggregate, a behavior that
decisively affects their bioavailability, transport and reac-
tivity [19].
SiO2 and TiO2 are among the most commonly applied
NPs in industry, followed by other oxidic particles. Approx-
imately half of the large number of studies dealing with
TiO2 describe toxic effects, whereas the other half report
either no [20,21] or positive effects [22,23]. These contro-
versial results can be explained, if the concentrations,
different experimental setups and systems used are
taken into account. Test concentrations range from 1 to
1000 mg/ml, which have to be compared with a value of
approximately 1 mg/m3 total amount of naturally occur-
ring ultrafine matter in the environment [12]. Thus, these
studies address the rare situation where organisms might
be exposed to an acute and high dose of NPs [11].
Although metal NPs still constitute negligible fractions
in the ultrafine matter, they are often studied owing to
their expected biological interactions [11]. For health rea-
sons, most interesting are platinum group metals, which
are contained in traffic exhaust together with metals, such
as Fe, Co, Al, Pb, Cd, Zn, which either naturally occur in
particulate matter or are highly toxic at low concentra-
tions.
Semiconductors, such as CdSe [selenium (Se)], CdTe
[tellurium (Te)] and GaAs [gallium (Ga), arsenic (As)], are
used in chip and electronic industry. The total amount of
these materials emitted to the environment is supposed to
be quite low. However, in future, these elements can
accumulate and become important. Cd in particular is
known to be very toxic. Silver (Ag)-NPs are also of increas-
ing use particularly because of their bactericidal activity,
which is exploited in many industrial applications [24].
NP uptake in plant systems
NPs are adsorbed to plant surfaces and taken up through
natural nano- or micrometer-scale plant openings. Several
pathways exist or are predicted for NP association and
uptake in plants (Figure 1) and are discussed below. The
aboveground surface of plant shoots usually exceeds
ground area and facilitates the deposition of airborne
NPs on plant shoot surfaces. Leaf area indices (LAI) given
as ‘m2 leaf area/m2 ground area’ range from 0.1 to 0.18,
with minimum values for species from deserts and tundra

Cuticle, epidermis

Bark, other surfaces

Long distance translocation (xylem/phloem)

Stomata
Shoot
Cell-to-cell

Hydathodes
apoplast

Stigma

Other entry sites

Nanoparticles

Root tips
Cell-to-cell
apoplast

Rhizodermis/cortex

Lateral root junctions

Root
Wounding
TRENDS in Plant Science

Figure 1. Pathways of nanoparticle association, uptake and translocation in plants. The line thickness tentatively correlates with the assumed significance of the pathways.
The schematic addresses shoots (green box) and roots (yellow box), movement in the tissue and transfer to vessels for long-distance transport (connecting blue box).
Broken lines indicate the assumption of very low rates of transport.

583
Opinion
to maximum values of 47.0 in coniferous tree stands. Only
14% of records have LAI greater than 8.0 [25] and average
LAIs are between 4.0 and 7.0 in temperate climates, thus
exposing significant areas to NP adsorption. Typical shoot
surfaces are lipophilic, because they are covered with
cuticular waxes. In addition, epicuticular structures, such
as tubes, platelets and papillae, frequently decrease wet-
ability (i.e. the Lotus effect; [26]) but probably increase
deposition of NPs in epicuticular cavities. Many shoot
organs are covered with trichomes [27], which are also
likely to increase NP deposition. In an attempt to investi-
gate shoot deposition, young maize (Zea mays) plants were
exposed for 20 min to 400 mg aerosol CeO2-NPs (which had
a normal size distribution of 37 nm) [28]. NPs were found
associated with the plants (approximately 50 mg/g fresh
weight) and could not be removed by surface washing,
suggesting a tight association with the plants. However,
the study did not quantitatively distinguish between NPs
adsorbed to the aerosol-treated leaf surface and NPs struc-
turally incorporated into the leaves.
NP uptake into the plant body can use different paths.
Uptake rates will depend on the size and surface properties
of the NPs. The outer surfaces of leaves, fruits and epider-
mal cells are mostly covered by cuticle. Very small and
lipophilic NPs can be incorporated into apolar fluid areas of
the cuticle, which contains apolar and polar pathways for
uptake. Permeation properties differ between cuticles on
the epidermal cells and trichomes or stomata [29]. Physical
and, thus, biomechanical properties of the cuticle also
change with temperature. The physical interaction and
penetration of NPs into the cuticle have not been analyzed.
Larger NPs can penetrate through cuticle-free areas, such
as hydathodes, the stigma of flowers and stomates. Aper-
tures of opened stomates reach values of 10 mm in diameter
in mesophytic plants, such as Vicia faba leaves exposed to
high light [30]. Penetration of NPs into leaves via stomates
has been shown after deposition of airborne CeO2-NPs.
However, newly developed leaves during a subsequent
phase of continued growth did not contain Ce above the
detection limit of 0.4 ng/g fresh weight, excluding any
significant long-distance translocation of CeO2 through
the phloem [28]. Likewise, no Ce was detected in above-
ground organs upon irrigation with CeO2-NPs for 14 d [28].
NPs that enter the intercellular gas space of leaves by
passage through the stomates are deposited on the cell wall
of the substomatal cavity or neighboring cells.
Below ground, organs such as roots and tubers develop
suberin layers as an interface with the soil environment.
Suberin is deposited as single layer or multi-layered la-
mella on the inner surface of the cell wall [29]. Most
primary roots that develop in soil have a suberinized
exodermis as well as a suberinized endodermis. The exo-
dermis prevents apoplastic bypass flow of solutes and
water from the soil to the central cylinder, as revealed
by application of the fluorescent dye 3-hydroxy-5,8,10-pyr-
ene trisulfonate (PTS), which has a weight of 508.4 g/mol
and is mobile in the apoplast, but does not pass the
exodermis [31]. However, in the more basal root zone of
lateral root development, the newly formed lateral roots
break through the cortex and apoplastic bypass is rendered
possible. Thus, at the site of lateral root formation, NPs
584
Trends in Plant Science November 2011, Vol. 16, No. 11
could enter the xylem via the cortex and the central cylin-
der. The ratio of apoplastic bypass flow and symplastic flow
increases with increasing transpiration [32]. However, the
consequences of these relationships on uptake and trans-
location of NPs is unknown.
ZnO-NPs (diameter 20 nm) administered to ryegrass in
the range 8–1000 mg/L in hydroponics were mostly associ-
ated with the root surface [33]. At the highest concentra-
tion, some ZnO particles were localized in the endodermis
and stelar cells and the root showed severe damage and
collapsed cells. How these findings relate to more natural
concentration scenarios of ZnO-NPs remains unclear. Lu-
minescent NaYbEr4:ytterbium (Yb), erbium (Er)-NPs (di-
ameter 45 nm) administered to soil grown Phalaenopsis
spp. and Arabidopsis thaliana plants as 1% irrigation
solutions also appeared in shoot sections. These particles
emit luminescence between 650 and 710 nm after photon
upconversion, if excited at 974 nm [34]. Luminescence was
observed in leaves and flower stalks. Initially, the NPs
accumulated in the velamen of the aerial orchid roots of
Phalaenopsis, but after 6 days, these NPs had reached the
central cylinder and had spread throughout the plant.
These results show that, despite the size exclusion limits
of the cell wall discussed below, at least in some plants
even large NPs are translocated over long distances after a
long period of time [34].
Uptake of NPs into plants is likely to be achieved upon
wounding as consequence of belowground herbivores and
mechanical injuries. Such pathways, in addition to lateral
root junctions, are known entry routes for bacteria [35] and
certainly are open for efficient particle uptake.
Uptake and translocation
NPs must traverse the cell wall before entering the intact
plant cell protoplast. Solute exclusion techniques provide
information on limiting pore sizes in cell walls. Based on
such experimental data, the maximum pore size of plant
cell walls is usually in the range of a few nanometers; for
example, 3.5–3.8 nm in root hairs and 4.5–5.2 nm in pali-
sade parenchyma cells [36]. Similar pore sizes are obtained
with chromatographic methods using ethanol-extracted
cell walls. Exclusion pore sizes are affected by pH, divalent
ions and boric acid [37], which suggests that only NPs less
than 5 nm in diameter will be able to traverse the cell wall
of undamaged cells efficiently. Because all the measure-
ments on cell pore exclusion were performed with bulk
materials, it cannot be excluded that cell wall defects or
discontinuities enable single NPs to reach and cross
the plasma membrane as described in [34]. Alizarin red
S-labeled TiO2-NPs of approximately 3 nm revealed a
tissue- and cell-specific distribution in Arabidopsis seed-
lings, traversed the cell wall and were found in epidermal
cells, for example, and at the subcellular level in different
compartments, including vacuoles [38].
Pits in the peripheral primary walls of xylem vessels
allow for perfusion of free dyes between adjacent vessels. In
maize, NPs smaller than 4.9 nm seem to perfuse to a minor
extent, whereas transfer of particles greater than 20 nm in
diameter is impeded. The authors of [39] estimated the size
exclusion limit to be 3.5 nm, which is in the range of other
primary cell walls compiled in the same study. Pit sizes in
 Opinion
xylem vessels of other species, such as Abies nordmanni-
ana, Thuja plicata, Gingko biloba and Eucalyptus regnans,
are reported to be much larger, with values in the mm range
[39]. Movement between vessels in such species should be
easily possible once NPs reach the xylem, because these
vessels constitute open conduits for particle movement
that are used by bacteria to spread within plants. Fluo-
Spheres polystyrene microspheres (diameter 1.0 mm) fed to
excised vine leaves moved from petioles, via petiole–leaf
blade junctions, into the 1st and 2nd order veins of the leaf
blade [40]. However, this connectivity can significantly
vary between species and pit membranes might restrict
free movement of larger NPs through the xylem in other
species. Pumpkin (Cucurbita maxima) plants exposed to
0.5 g/L Fe3O4-NPs (diameter 20 nm) in hydroponics medi-
um translocated 1.3% of the total amount to the leaves, as
determined by vibrating sample magnetometry. Most par-
ticles were associated with the pumpkin root surface. Lima
bean plants failed to show any magnetic signal, so did not
accumulate any particles. This difference reveals signifi-
cant variation in particle uptake and translocation among
species [41]. In the same study, pumpkin was shown to
take up F3O4-NPs from sand culture, whereas no uptake
was seen in soil culture. Thus, not unexpectedly, soil
functions as an efficient nanofilter, preventing NPs from
accessing and entering roots. Cell-to-cell translocation
depends on the size exclusion limit of plasmodesmata
Plasmodesmos
cell
Vacuole
phyll
Meso
Cell wall
Epidermis
Key: > 50 nm
8 – 20 nm
< 5 nm
Leaf surface
Figure 2. Distribution of nanoparticles (NPs) of three different size classes in shoots at the organ, tissue and cell level. All NPs associate with the shoot surface, partly enter
the intercellular space and associate with the outer apoplast, whereas only small NPs efficiently traverse cell walls and can enter the protoplast. The schematic depicts the
entire shoot, leaves and a cross-section of a leaf, showing epidermis, mesophyll and stomatal cells.
Trends in Plant Science November 2011, Vol. 16, No. 11
between adjacent cells. GFP (27 kDa) moves from compan-
ion cells to sieve tubes and from cell to cell [42]. A globular
protein of 30 kDa is approximately 3 nm in size, suggesting
that NPs smaller than 3 nm might also traffic between cells
and through the phloem. Carbon-coated magnetic NPs
injected into pumpkin shoots moved from the site of appli-
cation across, for example, the stem parenchyma tissue
and in xylem vessels [43]. Upon spraying, NPs also
appeared inside the epidermal cells of the petiole. Overall,
the efficiency of NP translocation was low. Figure 2 sum-
marizes the association, uptake and distribution of NPs of
different size in shoots.
Toxicology of NPs
Uptake, translocation and toxicity of NPs to humans,
animals and, in a few cases, to plants have been studied
in diverse experimental setups. The accumulation, persis-
tence and impact of NPs on plant metabolism and devel-
opment depend on the size, concentration and chemistry
of NPs, as well as the chemical milieu of the subcellular
sites to which the NPs are deposited. NPs can be toxic to
plant tissues, owing to chemical or physical effects. Thus,
NPs act as catalysts and interactors or, upon dissolution,
as soluble metal ions. Physical toxicity is linked to associ-
ation with cell structures or mechanical clogging. NPs
interact with biological systems by five main modes: (i)
chemical effects as metal ions in solution upon dissolution;
Gas space
Stoma
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Opinion Trends in Plant Science November 2011, Vol. 16, No. 11

Box 1. Basic characteristics of nanoparticles and their

significance for biological processes
Reactive Damage
NPs are characterized by their elemental and/or molecular composi- oxygen species to structures
tion and their surface area (Figure I) [11]. The chemistry defines the (nanotubes)
production
reactivity, whereas the surface area defines the effect size. In
addition, NP surface groups can covalently or non-covalently bind to
cell constituents, such as proteins. The size of the particles Catalytic Size-
determines their penetration depth into the tissue. It is known that
Redox activities of dependent Clogging
particles less than 5 nm translocate readily through pores of the cell
catalyst large mechanical
wall with a diameter of 5–8 nm [53]. Particles up to 20 nm can move
between cells through plasmodesmata [53]. Particles can also be surfaces Plant interactions
taken up by cells via endocytosis [64]. The surface chemistry is a nano-
major determinant of the ability of a molecule to penetrate a cell; for toxicology Lipophilic
example, polyethylenglycol-functionalized particles are able to enter Dissolution
cells through the cell membrane.
interactions
(ionic form)
Affinity-based (cuticle
interactions membrane
Nanoparticles Feature Effect Covalent protein)
on …
binding
Size Penetration (-S-protein, Au
Binding by functional
DNA)
groups (-OH, =O,
protein)
Surface area Effect amplitude
TRENDS in Plant Science

Figure 3. Categorization of nanoparticle-dependent toxicity mechanisms. Different

Au TiO2 PE Chemistry Persistence
Binding
OH
SH
Surface chemistry Reactivity
TRENDS in Plant Science
Figure I. Relationship between NP properties and biological effects.
Abbreviations: Au, gold; PE, polyethylene glycol.
(ii) mechanical effects owing to hard spheres and defined
interfaces; (iii) catalytic effects on surfaces; (iv) surface
effects owing to binding of proteins to the surface, either
by non-covalent or covalent mechanisms or oxidative
effects; and (v) changes in the chemical environment
(pH). For modes (ii–v) see also Box 1 [mode (i) is not covered
because it is not specific to NPs].
Chemical effects
Toxic metals, such as Cu, Cd, mercury (Hg), Ni and Zn,
bind to cell components including proteins, for example to
sulfhydryl, carboxyl or imidazole groups, and modify their
activities (Figure 3). This interference with cellular pro-
cesses often causes redox imbalances and oxidative stress
in metal-exposed plants [44,45]. Some metals, such as Cu
and Fe, transfer electrons to O2 as acceptors to form O2– or
to H2O2 to form the extremely reactive OH radical. These
reactions enhance oxidative stress in the affected cells.
Elemental particles of base metals often dissolve readily.
Oxidic particles dissolve as metal ions in solution with
highly varying rates and can thereby exert their toxic
action as ions. More noble metals, such as Ag, dissolve
slowly. NP concentrations in the atmosphere and soil are
usually low and most of them associate with the plant
surfaces. Therefore, the toxicity effect through this path-
way can be considered small, except for massive intoxica-
tion and local effects, owing to the release of large amounts
of ions at one specific site.
shades of the same color indicate related mechanisms subsumed to: (i) size-
dependent mechanical interactions; (ii) catalytic activities of large surfaces; and (iii)
affinity-based interactions. Abbreviation: Au, gold.
Mechanical effects
Mechanical effects depend on particle size and not on
particle chemistry (Figure 3). They include the filling of
pores, which are then unavailable for other transport
processes. An example is the inhibition of water transport
in the presence of high NP concentrations in the soil [46].
Catalytic effects
Many metals, including Ag, Pt, Pd, gold (Au), Fe and Co,
catalyze chemical reactions, such as reduction–oxidation
reactions (Figure 3). Metal ions at cell-compatible low
concentrations are usually bound to constitutive or induc-
ible chelators, including phytochelatins, organic acids,
metallothionein and ferritin [47] or are compartmentalized
by transport processes [48]. These mechanisms are inef-
fective if these metals are introduced as particles into cells
or cell compartments. However, depending on the accessi-
ble surface and cellular milieu, the catalytic activity of the
metal ions can be preserved and, as a result, modified
molecules, such as oxidized proteins, can accumulate. Such
mechanisms can explain the strong toxicity of Cu-NPs on
mice in vivo [49].
Surface effects
Particles with an oxidic surface often form a layer of OH–
groups at the surface, which are negatively charged and
thereby attract positively charged side groups of proteins
[50]. Once the proteins are bound to the particles, they are
either no longer available for their function or function
with lower efficiency. A more extreme effect occurs if side
groups of proteins bind covalently to the particles, such as
cysteines to Au surfaces, because this type of binding is
permanent. In a proteomic screen, 11 different engineered
nanomaterials revealed some common affinity to human

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proteins, such as peroxiredoxin 1, annexin A2 and ribo-
somal proteins [51]. Such preferential binding is assumed
to also occur in vivo and can induce specific toxicity effects.
Similar to many abiotic and biotic stressors, the most
common general stress symptom of NP toxicity appears
to be the development of oxidative stress by enhanced
production of reactive oxygen species and peroxidative
processes [52]. Oxidative stress in response to NPs has
been observed in diverse animal cell and tissue systems,
but remains largely unexplored in plants.
NP toxicity in plants
Most published studies on NP toxicity in plants have
addressed easily scored parameters, such as germination
rate and growth-related features. Positive, negative or no
effects are reviewed in [53] and we summarize some exam-
ples here.
Colloidal suspensions of TiO2-NPs (30 nm in diameter)
and bentonite, a natural clay, caused the cell wall pore size
of maize roots to decrease from 6.6 nm to approximately
3 nm and inhibited hydraulic conductivity and, therefore,
transpiration of intact plants [46], while the growth rate
was inhibited by approximately 10%. The inhibitory effect
was reversed upon transfer to NP-free growth medium,
indicating mechanical and nontoxic modes of action of
these NPs [46]. NPs administered to the roots by irrigation
were only found in the soil and in aggregates attached to
the roots [54]. The lack of strong growth inhibition by clay
and TiO2 was explained by the fact that intact root systems
are not limited in water flow to the shoot under normal
conditions. Only removal of 80% of the roots inhibited
transpiration of wheat (Triticum aestivum) [55]. The de-
velopment of excessive root systems might represent an
evolutionary adaptation to potential root clogging in soils
that naturally contain particulate minerals [46]. Similar to
the results in this study, very high NP concentrations often
lead to significant effects on plants. For example, zucchini
(Cucurbita pepo) seed germination and root elongation
were not significantly different in the presence of
1000 mg/L ZnO added as NPs or as bulk powder, whereas
Cu- and Ag-NPs inhibited root growth [56]. Dissolved Ag or
Cu ions could only partly explain the observed NP toxicity.
Different dose-response dependencies were seen for root
growth of radish (Raphanus sativus), rape (Brassica
napus) and ryegrass (Secale cereale). Inhibitory concentra-
tions 50% (IC50) ranged from 20 to 50 mg/L for Zn- and
ZnO-NPs [57]. Ag-NPs had genotoxic effects on Allium cepa
root tip cells, as indicated by chromosome aberrations [58].
At 40 mg/L, gum arabic-coated Ag-NPs completely inhib-
ited root hair formation, triggered deformation of roots and
caused cell collapse, whereas the same amount of dissolved
AgNO3 was inconspicuous [59]. These studies indicate that
some plant developmental stages and structures are par-
ticularly sensitive to NPs, in this case the growing root
apex of the grass Lolium. Likewise, pollen germination and
tube growth were more than one order of magnitude more
sensitive to Pd-NPs than to equivalent concentrations of
dissolved PdCl2 [60]. A table summarizing some reported
effects of NPs on plants is presented in [59]. Taken togeth-
er, these studies indicate NP-specific mechanisms of toxic-
ity that are dependent on NP chemistry and size.
Trends in Plant Science November 2011, Vol. 16, No. 11
A need for future research
Past and future research on plant nanotoxicology should be
placed in context of current risk assessments associated
with NPs, their use, distribution and release. In addition,
plant nanotoxicological research tailored to this demand
(US Environmental Protection Agency, cited in [41]) must
focus in general on: (i) characterization of NP properties; (ii)
analysis of transformation and interaction processes; (iii)
elucidation of environmental fate and transport; (iv) assess-
ment of human exposure; (v) study of ecological effects; (vi)
investigation of human health effects; and (vii) determina-
tion of the NP life cycle. As described in this Opinion,
interactions between NPs and plants affect transformation,
environmental fate and life time of NPs and can have
significant implications for human health, either positive
as atmospheric clearance or negative as carriers into the
human diet. An understanding of the ecological effects of
increasing NP emissions must include plant nanotoxicology.
At present, it is premature to draw conclusions about the
effect of naturally occurring levels of NPs on plants in the
field and about the role of plants in relation to NP-related
effects within ecosystems. The situation might change if
large amounts of NPs are administered to crops as novel
delivery tool for fertilizers, herbicides and pesticides, as
suggested in [61]. Future research projects should specifi-
cally focus on understanding nanotoxicity in plants (see also
[53]). Prerequisites for insightful experiments are compre-
hensive details about the experimentally used NPs, their
size, surface area and chemistry [11,62]. Deposition, uptake,
cell-to-cell and long-distance translocation of NPs adminis-
tered at reasonable concentrations need to be put into
context of plant structures, such as the epicuticle, stomates,
plasmodesmata and xylem. An important point is to assess
NP persistence in plants depending on NP size and chemis-
try, similar to studies in animal cell systems [63]. Up to now,
most studies in plants indirectly drew conclusions on the
overall stability of NPs by considering the long-distance
translocation of elements within the plants. Because the
apoplast is the first site of NP deposition within the plant
tissue, a major emphasis should be the interaction of NPs
with apoplast structures, interference with apoplast func-
tions and NP persistence in the specific chemical environ-
ment of the apoplast, including the cell wall. The search for
molecular mechanisms of toxicity needs to address NP
association with cell constituents, such as plasma mem-
brane proteins, as possible primary sites of molecular
interactions. Investigations of plant species- and geno-
type-specific differences in NP sensitivity must be intensi-
fied. As a novel approach to distinguish ion toxicity from
NP-specific effects, it is suggested to compare closely related
species with known differences in metal tolerance. Direct
observation of NPs on plant surfaces and in plant tissues has
become possible by new methods using quantum dots and
novel microscopy technologies and should be used for in-
depth studies on the localization of particles in plants.
Finally, plants grown in their natural habitat or in the field
need to be assessed for particle accumulation to judge the
role of plants in the life cycle of NPs and the risk of intro-
ducing NPs into the food chain. This topic must also address
soil–plant interactions, although this route appears less
significant for plant nanotoxicology.
587
Opinion
Acknowledgments
We gratefully acknowledge stimulating discussion with Lukas Schreiber.
This review is part of our research efforts within the SFB613, project D12.
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