Ecotoxicology and Environmental Safety 271 (2024) 115992

Contents lists available at ScienceDirect

Ecotoxicology and Environmental Safety

journal homepage: www.elsevier.com/locate/ecoenv

Review

Dynamic interplay of metal and metal oxide nanoparticles with plants:

Influencing factors, action mechanisms, and assessment of stimulatory and
inhibitory effects
Rabia Javed a, *, Bakhtawar Khan b, Uzma Sharafat a, Muhammad Bilal b, Lakshman Galagedara a,
Lord Abbey c, Mumtaz Cheema a, *
a
School of Science and the Environment, Memorial University of Newfoundland, NL, Canada
b
Department of Biotechnology, Quaid-i-Azam University, Islamabad 54320, Pakistan
c
Department of Plant, Food and Environmental Sciences, Faculty of Agriculture, Dalhousie University, NS, Canada

A R T I C L E I N F O A B S T R A C T

Edited by: Dr Muhammad Zia-ur-Rehman Nanoparticles (NPs) of metals and metal oxides have received increasing attention regarding their characteristic
behavior in plant systems. The fate and transport of metal NPs and metal oxide NPs in plants is of emerging
Keywords: concern for researchers because they ultimately become part of the food chain. The widespread use of metal-
Nanoparticles based NPs (MBNPs) in plants has revealed their beneficial and harmful effects. This review addresses the
Abiotic stress, Toxicity and detoxification
main factors affecting the uptake, translocation, absorption, bioavailability, toxicity, and accumulation of MBNPs
mechanisms
in different plant species. It appraises the mechanism of nanoparticle-plant interaction in detail and provides
Uptake and translocation
Elicitation of secondary metabolites understanding of the estimation strategies for the associated pros and cons with this interplay. Critical param­
eters of NPs include, but are not limited to, particle size and shape, surface chemistry, surface charge, concen­
tration, solubility, and exposure route. On exposure to MBNPs, the molecular, physiological, and biochemical
reactions of plants have been assessed. We have filled knowledge gaps and answered research questions
regarding the positive and negative effects of metal and metal oxide NPs on seed germination, callus induction,
growth and yield of plant, nutritional content, antioxidants, and enzymes. Besides, the phytotoxicity, cytotox­
icity, genotoxicity, and detoxification studies of MBNPs in plants have been outlined. Furthermore, the recent
developments and future perspectives of the two-way traffic of interplay of MBNPs and plants have been pro­
vided in this comprehensive review.

1. Introduction
Nanotechnology is emerging as the fifth technology of this century. It
is known for its extensive range of applications in many disciplines such
as agricultural science, material science, environment, and medicine
(Mohammad et al., 2022; Haleem et al., 2023). It is the study of mate­
rials’ size at a dimension ranging from 1–100 nm, their formulation,
characterization, and applications. At nanoscale, materials have specific
physical, chemical, optical, electrical, and mechanical properties
compared to their bulk form (Chhipa, 2017). Anthropogenic source is
the natural source to derive nanoparticles (NPs) and it can also be
manufactured in industries (Khan et al., 2019a). NPs can be classified
into organic as well as inorganic forms; organic NPs comprise fullerenes
and carbon-based NPs, however inorganic NPs are generally categorized
as metals and metal oxides (Iranbakhsh et al., 2021). Metal NPs (MNPs)
include copper (Cu), zinc (Zn), gold (Au), silver (Ag), iron (Fe), etc.,
while metal oxide NPs (MONPs) are the modified form of their respec­
tive MNPs and include copper oxide (CuO), zinc oxide (ZnO), iron oxide
(Fe2O3), silver oxide (AgO), titanium oxide (TiO2) NPs, etc. (Ealia and
Saravanakumar, 2017).
Stress tolerance and crop improvement can be managed by the
application of NPs. Because of their small size, NPs are very reactive that
makes them suitable for varying different plant’s biological functions
(Javed et al., 2023a) and improve plant growth and metabolism in a
species-specific manner, even under conditions of biotic and abiotic
stress conditions (Giraldo et al., 2014; Hanif et al., 2023a). To

* Corresponding authors.
E-mail addresses: rjaved@mun.ca (R. Javed), usharafat@mun.ca (U. Sharafat), lgalagedara@mun.ca (L. Galagedara), labbey@dal.ca (L. Abbey), macheema@
mun.ca (M. Cheema).

https://doi.org/10.1016/j.ecoenv.2024.115992
Received 9 September 2023; Received in revised form 10 January 2024; Accepted 13 January 2024
Available online 22 January 2024
0147-6513/© 2024 The Authors. Published by Elsevier Inc. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-
nc-nd/4.0/).
R. Javed et al.
comprehend the possible benefits of MNPs and MONPs in agriculture,
the first step should be to analyze the penetration and transport of these
NPs in plants (Du et al., 2017). There are various ways in which NPs
interact with plants, such as direct application, accidental release, and
presence in soil as soil pollutants or atmosphere. When NPs reach the
soil and root zone, they interact with plants in a non-partial manner, and
hence influence the physiological and biochemical processes of plants
(Khanna et al., 2022).
The interaction of NPs with plants is significantly influenced by size,
shape, surface charge, surface modification, composition, and reactivity
of NPs (Parveen and Ledwani, 2022). Only a few studies have shown
that plant species, NPs characteristics, dose, and application method all
affect NPs impacts as well as NPs uptake, translocation, transformation,
and accumulation in plants. (Ebbs et al., 2016). Interestingly, MNPs and
MONPs improve the plant productivity and enhance agricultural inputs
by facilitating site-targeted delivery of nutrients and ensuring the min­
imal use of agricultural inputs. Additionally, it is crucial to encourage
plant adaptability to changing environmental factors like water scarcity,
temperature, freezing, water logging, salinity, and heavy metal
contamination without harming the ecosystem (Vermeulen et al., 2012;
Zhao et al., 2020).
Several positive effects of MNPs and MONPs to plants are reported,
but these NPs also display negative impacts on plants (Liu et al., 2020).
The induction of abiotic stress tolerance by metal-based NPs (MBNPs)
has been considered the basic reason for causing nanopollution or
nanotoxicity. To determine whether the MBNPs pose a threat to plants
and the environment, analytical data on their characterization, influ­
encing factors, and absorption and transport mechanisms are required to
study their impacts on various species of plant (Rastogi et al., 2017; Gao
et al., 2023). MNPs as well as MONPs produce a toxic amount of reactive
oxygen species (ROS) and as a result cause oxidative stress in plants
leading to phyto-, cyto-, and genotoxicity. The plant defense system is
triggered encompassing enzymatic and non-enzymatic antioxidants to
fight against the hazardous radical species and ROS (Ma et al., 2015;
Ranjan et al., 2021). Hence, the assessment of both the beneficial and
antagonistic effects of MBNPs on plant growth, physiological, morpho­
logical, molecular, and biochemical aspects is very important.
Fig. 1. Factors influencing the interplay of nanoparticles (NPs) and plants that are crucial in their stimulatory or inhibitory response.
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Ecotoxicology and Environmental Safety 271 (2024) 115992
Multiple reports exist on this topic, but we have compiled the most
recent data analyzing the interplay of MBNPs with plants. Moreover,
none of the previous review article has comprehensively detailed all
parameters of NPs affecting the relations of MBNPs with plants along
with the associated mechanisms and assessment methods for estimating
the stimulation or inhibition of MNPs- and MONPs-specific nano-plant
interaction. The intent of this review article is to deliver the existing
understanding of the MBNPs-plants interaction under changing sce­
narios. The areas that merit further research are pin-pointed and the
challenges faced by researchers for applying MBNPs to plants are
highlighted. Also, an outlook for future research has been provided.
2. Factors affecting interplay of nanoparticles with plants
Various factors influence the interaction of NPs with plants, and they
exhibit diverse relationships, yet they are all interconnected. The key
influential factors of NPs have been described below and are visually
represented in Fig. 1.
2.1. Size and Shape
The valuable and detrimental impacts of NPs on plants are mostly
dependent on their size. The size of NPs and surface to volume ratio are
correlated; the smaller the NP size, the higher the surface volume ratio,
and vice versa. The size of the NPs has a direct influence on their ability
to penetrate, absorb, and respond (Naz et al., 2020). According to
several reports, NPs between 5 and 20 nm in diameter can easily permit
through the plasma membrane. The cell wall’s pores extend to allow NPs
to enter if their size is more than 20 nm (Nhan et al., 2015). But
depending on the crops, certain NPs of different sizes produce various
effects. It has been shown that the uptake of Au NPs occurred through
both the cuticular layer and stomata when Au NPs of numerous sizes (3,
10, and 50 nm) were given to Triticum aestivum (wheat). Larger-sized
(50 nm) Au NPs had the opposite effect and reduced the exposed
plant’s dry biomass. Smaller-sized (3 nm, 10 nm) Au NPs enhanced
stomatal conductance and photosynthetic rate (Avellan et al., 2021).
Mesoporous silica NPs of 15, 100, and 200 nm produced distinct effects
R. Javed et al.
on pesticide loading, release, and transport in Cucumis sativa (cucumber)
in a comparative research. The larger-sized NPs were found to load more
readily than the smaller ones, which may be connected to the pore
volume and diameter of the NPs. The smaller-sized NPs, meantime, were
quickly engrossed by the roots and transported to the plant’s aerial
portions (Xu et al., 2021).
Shape of NPs plays a crucial role in nano-bio interactions; viz.
cellular uptake, internalization, circulation, distribution, and residency
duration inside the cells, also affecting the action mechanisms. Elon­
gated NPs can easily adhere to the cells compared to spherical NPs of the
similar size because of multiple binding sites for interaction with the
cellular receptors. However, spherical NPs are more efficient than
ellipsoidal NPs (Salatin et al., 2015). Hence, sharp shapes with increased
exposed surfaces exhibit higher reactivity because of their larger surface
area and smaller size (Ali et al., 2020). For example, Au NPs having
round-spherical shape with different sizes (10–18 nm) when directly
exposed to Arabidopsis thaliana negatively affected the phytotoxicity of
plants (Siegel et al., 2018). Besides, Ag NPs when applied to Arabidopsis
thaliana with different sizes and morphologies (47 ± 7 nm with trian­
gular shape, 8 ± 2 nm with spherical shape, and 45 ± 5 nm with deca­
hedral shape), decahedral shape of 45 ± 5 nm size influenced the root
growth promotion (RGP), while spherical shape of 8 ± 2 nm sized Ag
NPs displayed no RGP and increased the highest level of anthocyanin
collection. However, all three different sizes and shapes of Ag NPs
induced gene expression and root growth promotion (Syu et al., 2014).
2.2. Surface charge and surface chemistry
Charges of NPs could be modified by changing their physical struc­
ture and surface chemistry (Javed et al., 2016; Ali et al., 2020). Cationic,
anionic, and mixed-charged NPs can all form non-covalent and elec­
trostatic interactions with the cell walls of plants. Particularly, nega­
tively charged surfaces are linked to positively charged NPs, while
positively charged molecules on the surface are selectively bound by
negatively charged surfaces (Nangia and Sureshkumar, 2012). Accord­
ing to Lv et al. (2019), positively charged Au NPs of 12 nm size were
adsorbed to negatively charged mucilages released by Arabidopsis
thaliana roots, which prohibited the translocation of Au NPs into the root
tissues. However, negatively charged Au NPs did not bind to mucilage
and were able to enter the apoplast roots.
Coating/capping of NPs changes the surface chemistry by modifi­
cation and alters their reactivity (Cartwright et al., 2020). It also reduces
particle aggregation and enhances their dispersion and stability result­
ing in increased uptake and internalization by the plant cells (Ahmad
et al., 2023; Javed et al., 2023b). For instance, coated Fe3O4 NPs with
spermine having a spherical shape were applied to the soil grown Ros­
marinus officinalis (rosemary) that significantly increased resistance to
drought stress condition (Afrouz et al., 2023). It was reported that un­
coated CeO2 NPs applied to Lactuca sativa (lettuce) inhibited root
growth, reduced solubility, induced oxidative stress, as well as caused
root’s cell death. However, after coating the surface with organophos­
phate, the toxic effect of CeO2 NPs on lettuce was significantly weak­
ened. The modification of CeO2 NPs with organophosphate reduced the
release of Ce2+ ions resulting in lowering of toxicity (Zhao et al., 2021).
Recently, Hanif et al. (2023b) reported the alleviation of salinity stress
by the proline capped ZnO NPs in Coriandum sativum (coriander).
2.3. Solubility
The solubility of NPs is one of the most meaningful parameters, and
their solubility in solvents is facilitated by the release of their metal ions,
setting them apart from their bulk form. The metal ions release from NPs
plays a crucial role in defining both their solubility and toxicity (Naz
et al., 2020). It has been noted that the solubility of Ag NPs in freshwater
produced Ag+. These Ag+ reduced plant biomass, chlorophyll content,
and produced ROS in aquatic plant, Lemna minor under different
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Ecotoxicology and Environmental Safety 271 (2024) 115992
concentrations (10, 25, and 50 μg/mL) (Souza et al., 2021). Moreover,
solubility can also depend on particle size due to changes in surface area.
The greater surface area and decreased size of NPs facilitates dissolution
by maximizing the presence of exposed surface sites that can actively
participate in the dissolution/solubility mechanism (Hedberg et al.,
2019). CuO, NiO, and TiO2 NPs’ solubility in two different aqueous
media, namely water and Dulbecco’s modified Eagle’s medium
(DMEM), has been compared to that of the particles’ bulk forms. Due to
the development of Cu and Ni ion complexes, both CuO and NiO NPs
became more soluble in DMEM, whereas this tendency was the opposite
in water. In case of TiO2 NPs, higher solubility was observed in water
compared to DMEM (Avramescu et al., 2020).
2.4. Concentration
Concentration of NPs plays an important role in conferring beneficial
or harmful effects to plants. For instance, FeO NPs applied to Pisum
sativum (black-eyed peas) at 250–500 mg/L concentration increased
growth and chlorophyll contents. Similarly, when applied to Glycine max
(soybean) at the concentrations of 30–60 mg/kg, a higher amount of leaf
chlorophyll was observed (Ghafariyan et al., 2013). It was estimated
that the impact of Ag NPs on plant growth, i.e., whether it leads to in­
hibition or stimulation of growth depends on Ag NPs’ concentration. It
was noticed that the exposure to certain concentrations of Ag NPs can
nurture plant growth in comparison with the non-exposed plants.
However, both higher and lower concentrations adversely affected plant
growth (Almutairi and Alharbi, 2015). Apart from this, Fe2O3 NPs
applied to Oryza sativa (rice) crop with low dose (50 mg/L) improved
the growth of rice crop, while higher concentration (500 mg/L) reduced
growth and root length, surface area, diameter, and volume (Li et al.,
2021). In a recent study, Cu NPs (size: 25 nm) and ZnO NPs (size:
100 nm) revealed a positive effect on Lactuca sativa (lettuce) growth,
assessed by evaluating root elongation and biomass increase during
exposure for about 20 days to various concentrations (0.0 to 0.2 mg/L
and 0.2 to 20 mg/L) (Song et al., 2022).
2.5. Media, duration, and methods of exposure
Different media can be used for growing various plant species, for
instance, agar culture media, soil media, and aqueous media. Different
exposure media exhibit distinct behaviors, and prolonged durations of
exposure leads to increased accumulation and toxicity of NPs (Cox et al.,
2016). When Zea mays (maize) crop was exposed to CuO NPs in hy­
droponic media, it increased seed germination, root length, and leaf area
of maize crop. Besides, it increased photosynthetic pigments as well as
plant dry biomass (Rizwan et al., 2017). The mechanism of root and
shoot organ formation of Stevia rebaudiana (candy leaf) differ in solid
and liquid Murashige and Skoog (MS) culture media. The results showed
that liquid MS culture produced the best yield, while solid MS culture
produced highest amount of steviol glycosides (rebaudioside A and
stevioside) (Javed and Yücesan, 2022). The toxicity of Al2O3 NPs to Zea
mays (maize) in hydroponic and soil culture media was higher in hy­
droponic culture than in soil culture (Ahmed et al., 2022), which might
be due to the intrusion of natural organic matter as well as ion exchange
capacities in soil that were absent in hydroponic media.
Exposure duration of NPs plays pivotal role in their influence on
plants. Ahmed et al. (2022) compared the bioavailability and toxicity of
Al2O3 NPs to Zea mays (maize) crop applied for a duration of 20 and 40
days. The results revealed higher oxidative stress by the Al2O3 NPs and
Al+3 at 40 days than 20 days, and the antioxidant enzyme activities and
malondialdehyde (MDA) contents were found significantly lowered
after 40 days of exposure duration. In another study, CeO2 NPs were
applied to Phaseolus vulgaris (kidney beans) at different concentrations
(62.5–500 mg/L) in hydroponic culture for the duration of 1, 7, and 15
days. At 125 mg/L, kidney beans showed tolerance and effectively
defended the oxidative stress. However, at 500 mg/L for 15 days, the
R. Javed et al.
roots exhibited stress response revealed by increasing soluble proteins
and overpowering of antioxidant enzymatic activities (Majumdar et al.,
2014). Hence, both the concentration of NPs and exposure duration of
media showed effect in this study.
The routes/modes of NPs exposure to plant cells and tissues are also
important in directing their responsiveness. There are three primary
routes of exposure to NPs; direct injection of NPs into plant parts, foliar
spray of NPs, and direct injection of NPs into soil (Jogaiah et al., 2021).
However, roots could directly interact with NPs, enabling the absorption
of these NPs from the soil and subsequent transport to different tissues
within the plant. For example, CuO NPs of particle size < 7 nm applied
to Zea mays (maize) crops were easily absorbed by the roots and moved
to the aerial parts. Likewise, CeO2 NPs of size > 7 nm was taken up by
plants including Medicago sativa (alfalfa), Solanum lycopersicum (to­
mato), Cucumis sativus (cucumber), etc. (Ahmed et al., 2021a). On the
other hand, the foliar application seems even more efficient than soil
amendment. For instance, the foliar exposure to MgO NPs, ZnO NPs, and
CuO NPs resulted in increased seed yields for Gossypium hirsutum (cot­
ton) and fruit yield per tree for Punica granatum (pomegranate)
(El-Saadony et al., 2021).
2.6. Exposure, uptake, and translocation of nanoparticles in plants
The exposure, uptake, and transport of NPs in plants is determined by
both the physicochemical properties of NPs and the physiology of plants
(Schwab et al., 2016; Mohammadi Alagoz et al., 2022). Scanner electron
microscopy (SEM) and transmission electron microscopy (TEM) of root,
stem, and/or leaf tissues can be used to see how NPs are taken up,
absorbed, and transported in the plant tissues. There are few other
advanced techniques that can also be utilized for this purpose. For
instance, Stegemeier et al. (2017) employed µ-X ray fluorescence
(µ-XRF) to study the Ag NPs distribution in the root tip of Landoltia
puctata. Peng et al. (2017) used inductively coupled plasma optical
emission spectrometer (ICP-OES) and µ-XRF to detect CuO NPs in the
Oryza sativa tissues.
Soil amendment and foliar spray are two pathways for delivery and
movement of NPs from roots-stem and leaves-stem of plants through
xylem and phloem tissues, respectively (Du et al., 2017). In the case of
root exposure, NPs may enter the plant cells via cracks created during
the formation of lateral roots and later on via aquaporins, cell wall pores,
endocytosis, plasmodesmata, and/or ion transporters (Khan et al.,
2019a) (Fig. 2). Ion channels of plasma membrane called aquaporins,
which are up to 1 nm size, cause the entry of NPs of < 1 nm size making
Fig. 2. Exposure, Uptake, and Transport of nanoparticles (NPs) in plants involves foliar and soil entry routes. a) NPs are absorbed by stomata, and transfer from
leaves and stems to roots, b) NPs transfer through apoplastic pathway, c) NPs enter and transport by xylem and phloem, and transfer from roots to leaves or stems, d)
NPs transport through diffusion, endocytosis, carrier proteins, and channels (Gao et al., 2023).
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Ecotoxicology and Environmental Safety 271 (2024) 115992
them potential carriers for uptake by diffusion (Foroozandeh and Aziz,
2018; Yin et al., 2019). The NPs ranging in 5–10 nm size can enter
through the cell wall pores. The plasma membrane is disturbed by the
NPs causing a puncture which allows its entry into the cells (Chichiriccò
and Poma, 2015). Endocytosis is an entry method for NPs by invading
the cell membrane and moving in the form of intracellular vesicles after
being engulfed. Mostly 20–50 nm sized NPs can enter via endocytosis
(Sousa De Almeida et al., 2021). Ion transportors or carrier proteins also
play a significant role in the transport of NPs through root cells (Khan
et al., 2019a).
The translocation of NPs is done by the apoplastic or the symplastic
pathway. Apoplastic pathway is the one in which movement takes place
through cell wall without entering inside the cell, whereas if the
movement take place through cell cytoplasm, this is called symplastic
pathway. This movement gives an indication about the target site of
accumulation of NPs in plant tissues. Apoplastic transport takes place
when the NPs move in through root endodermis to the aerial parts of
plant (Buckley, 2015), whereas symplastic transport involves movement
through specialized structures known as plasmodesmata and sieve
plates. Plasmodesmata are specialized channels (40 nm size) composed
of complex intracellular cytoplasmic bridges. NPs up to 40 nm can be
transported by plasmodesmata, but it requires the NPs to be present in
the symplast for their movement (Khan et al., 2019a; Hong et al., 2021).
Entry of NPs to xylem results in their movement to the upper parts of
plant.
The foliar exposure causes entry through stomata, cuticle, lenticels,
and/or trichomes which then move downwards through phloem (Khan
et al., 2019b) (Fig. 3). Mostly, NPs the entry occurs via stomata. The
diffusion through foliar spray is more effective than soil amendment
because of the presence of nanosized pores in leaf plasmodesmata that
support efficacious nutrients delivery. The foliar part is protected by
cuticle. NPs cross the cuticle barrier and ultimately reach the target site.
Cuticle prevents entry of NPs of < 5 nm size (Ali et al., 2021). For their
entry, they may use hydrophilic or lipophilic path. The hydrophilic
uptake may be by diffusion or by polar aqueous pores in stomata, while
lipophilic diffusion occurs through cuticular waxes (Hong et al., 2021).
NPs of > 10 nm size cross through stomata. Endocytosis of NPs can also
occur in case of foliar exposure. If the NPs’ translocation is good enough
through phloem, the foliar application must be exploited (Avellan et al.,
2021). In a study, foliar spray of Se NPs mitigated the effect of drought
and improved yield in Punica granatum (pomegranate) (Zahedi et al.,
2021). Another study by Rossi et al. (2019) elucidated that the foliar
spray of ZnO NPs on Coffea arabica (coffee plant) enhanced the
R. Javed et al.
Fig. 3. Stimulatory and inhibitory pathways of metal nanoparticles (MNPs) and metal oxide nanoparticles (MONPs) after penetration and accumulation inside the
plants (Rastogi et al., 2017).
photosynthetic activity and increased biomass and quality of product.
2.7. Mechanism of action of nanoparticles in plants
The interaction of metallic and metallic oxide NPs with plants in­
fluences their growth and development, nutrient quality, oxidative
response, metabolism, gene expression, and other parameters
throughout their life cycle (Fiol et al., 2021). The NPs-plants interaction
is a two-way traffic and produces negative and positive responses in
plants (Fig. 3) that are described below:
2.8. Toxicity of plants by nanoparticles
MNPs and MONPs may injure plant tissues by the leaching of metal
ions or by interacting directly with them. Reduced biomass, decreased
seed germination, increase of oxidative stress, inhibition of photosyn­
thesis, disruption of chlorophyll synthesis, mitochondrial dysfunction,
DNA damage, chromosomal aberrations, inhibition of transcriptional
and translational machinery, and modifications in the ultrastructure of
cellular and subcellular organelles, are a few of the negative effects to
which these NPs have been associated (Rajput et al., 2020). ROS are the
products of aerobic respiration and are signaling molecules that are
produced by Fenton-type reactions in plants. These involve non-radical
species such as hydrogen peroxide (H2O2), singlet oxygen (1O2), and
radicals like hydroxyl (.OH) and superoxide (O.-2 ). The ROS are not toxic
if triggered in appropriate amount, but their over-production leads to
negative effects in plants. By releasing excessive ROS, MBNPs frequently
cause phytotoxicity. .OH having single unpaired electron has been found
to be the most lethal free radical (Yang et al., 2017). Over-production of
ROS allows it to interact with several biological elements and causes
various types of cell damage. It has the ability to change the electric
charge, bring about oxidative stress, and break apart peptide chains
damaging proteins, destroys cell membranes, and causes lipid peroxi­
dation (Kapoor et al., 2019; Dhiman et al., 2021). Lipid peroxidation
indicates cell membrane integrity which is damaged by ROS
over-production (Sadžak et al., 2020). Phytotoxicity occurs when ROS
amount surpasses the appropriate levels. The stages of plant growth are
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Ecotoxicology and Environmental Safety 271 (2024) 115992
adversely impacted by the harmful effects of NPs’ exposure on numerous
biochemical as well as physiological processes in various plant species.
Phytotoxicity is associated with the processes of cytogenotoxicity. The
cytotoxicity of NPs results in destruction of electron transport chain
(ETC) of mitochondria as well as impediment of photosystem I & II of
chloroplast affecting respiration and photosynthesis of plants. The
regulation and signaling of Ca2+ is also compromised. Mazumdar (2014)
studied the cytotoxicity of Ag NPs on Vigna radiata and Brassica cam­
pestris by which Ag NPs destroyed the vacuoles and other cellular or­
ganelles. Genotoxicity of NPs is the change in gene expression as a
response to any physical or chemical interaction. Genotoxicity can be
direct or indirect; the former involves physical interaction between NPs
and DNA, whereas, the latter might be due to the curtailment of DNA
repair (Jogaiah et al., 2021). NPs of < 50 nm size enter into the nucleus
via nuclear pores. It causes damage to DNA leading to impairment of
mRNA synthesis, inactivation of encoded proteins, and cell cycle. All
these phenomena impair cells’ normal functioning, ultimately resulting
in cell death (Zafar et al., 2023). Prakashi et al. (2014) studied the
genotoxic effects of TiO2 NPs on Allium cepa leading to DNA damage.
2.9. Detoxification of nanoparticles by plants
The toxicity of NPs is reversed by detoxification of ROS produced by
them through oxidative stress. As a result of over-production of ROS, the
defensive machinery of plants is also activated. This produces antioxi­
dants that may be non-enzymatic (thiols, ascorbic acid (AA), glutathione
(GSH), phenols, flavonoids) or enzymatic (catalase (CAT), peroxidase
(POD), superoxide peroxidase (SOD), ascorbic acid peroxidase (APX),
glutathione S-transferase (GST), glutathione reductase (GR), glutathione
peroxidase (GPX), guaiacol peroxidase (GPOX)) signaled by the detox­
icity pathways (Yang et al., 2017). The antioxidant activities and en­
zymes production work to balance the radical species that scavenge
excess free radicals or transform them to non-toxic forms. It functions to
prevent nuclear, mitochondrial, and chloroplast damage. AA is the most
crucial lower molecular weight antioxidant that acts as a first line of
defense against oxidants by scavenging OH and O.-2 . GSH is the
non-protein low molecular weight thiol that plays a central part in
R. Javed et al.
intercellular antioxidative defense. In addition, carotenoids and to­
copherols are the non-enzymatic antioxidant agents that are vital for the
defense system of plants. Regarding the role of enzymatic antioxidants
in detoxicity of ROS, SOD is a very efficient intracellular metal enzyme,
that catalyzes more toxic O.-2 to less toxic H2O2 and O2. CAT catalyzes
less toxic H2O2 to non-toxic H2O and O2. POD and APX perform similar
functions. However, APX is more effective in the conversion of H2O2 to
H2O (Chichiriccò and Poma, 2015; Ma et al., 2015;Khan et al., 2019a).
All the antioxidants are mostly produced in mitochondria, chloroplasts,
peroxisomes, vacuoles, and endoplasmic reticulum (ER) and act to
enhance tolerance in plants against the toxicity of MBNPs by mitigating
oxidative stress. It eventually promotes growth indices, development,
and yield of plants through the rise of phytonutrients and antioxidants in
the plant system (Du et al., 2017; Ali et al., 2021). The homeostasis of
biological macromolecules & micromolecules is balanced and crop
quality is enhanced in this way.
2.10. Assessment of beneficial and adverse effects of metal-based NPs on
plants
The rapid growth of agro-nanotechnology in the past two decades
has raised concerns as regard of the potential positive and negative ef­
fects of NPs (Rizwan et al., 2017). Literature states that most of the
research work on the effect of MNPs and MONPs on plants showed their
toxic effects, while few studies also reported their beneficial part in the
way of improving growth parameters and plant productivity (Landa
et al., 2016; Wang et al., 2016). The studies have shown that the use of
non-toxic concentrations could promote seed germination, improve
growth of plant, and also increase yield. In addition, MBNPs can be used
to protect plants from environmental influences, for instance, salt or
drought stress, and to reduce heavy metal accumulation and toxicity.
MBNPs can act as a source of micronutrients (such as ZnO, and Fe- and
Mn-based NPs), which increase resilience and help plants adapt to stress
conditions (Landa, 2021). Moreover, activation of the antioxidant sys­
tems and the formation of secondary metabolites are the common re­
sponses of plants under MNPs and MONPs toxicity (Li et al., 2017; Mosa
et al., 2018). In the following sub-sections, estimation of beneficial and
detrimental effects of MBNPs are reported at the morphological,
biochemical, physiological, and molecular levels in important food
crops (Fig. 4).
Fig. 4. Methods for estimation of positive and negative effects of metal-based
nanoparticles (MBNPs) on plants.
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Ecotoxicology and Environmental Safety 271 (2024) 115992
2.11. Examples of stimulatory and inhibitory effects of metal-based NPs
on plants
Different types of metal and metal oxide NPs having distinct physi­
cochemical properties when applied by distinguishing exposure routes
under different concentrations and duration produce either positive or
negative effects, and/or both (Table 1). These effects are essential to be
studied in detail in regards to varying parameters of NPs and their
application.
2.12. Seed germination
The first step in elucidating the toxicity of MBNPs in plant growth is
seed germination. Numerous studies are reported on the impact of MNPs
and MBNPs on the germination of plant seeds, resulting in both stimu­
latory and inhibitory effects. In a study, seven different concentrations
(0.05, 0.1, 0.5, 1, 1.5, 2, and 2.5 mg/mL) of Ag NPs were used to study
its effect on three plants, i.e., Zea mays (corn), Citrullus lanatus (water­
melon), and Cucurbita pepo (zucchini). Results showed that Ag NPs
enhanced rate of seed germination of three plants with significant
enhancement observed in watermelon and zucchini plants at 2 and
2.5 mg/mL respectively, as compared to untreated seeds (Almutairi and
Alharbi, 2015). On the contrary, rice seed germination (Oryza sativa L.
cv. KDML 105) declined with an increase in size and concentration of Ag
NPs. The inhibitory effect was reported to be due to decreased pene­
tration and transportation of Ag NPs through the plant tissues (Thue­
sombat et al., 2014). In a study, increasing concentrations (10, 25, 50,
100, 250, 500, 1000, 2000 mg/L) of Cu and CuO NPs were assessed on
Hordeum vulgare (barley) seedlings for their stimulatory or inhibitory
effects. Results depicted that seed germination and seedling growth
parameters were enhanced by low concentrations of both Cu and CuO
NPs while concentrations over 500 mg/L exhibited inhibiting effects
(Kadri et al., 2022). In another study, seed germination in Brassica nigra
(mustard) seeds was decreased with increasing concentrations (200,
400, and 600 mg/kg) of ZnO NPs embedded in the soil (Rehman et al.,
2020). It can be concluded that seed germination can be ameliorated or
mitigated by employing MBNPs on different growing media of crop
species depending on various parameters.
2.13. Morphology and physiology of plants
The health of plants is determined by their morphological and
physiological parameters, such as stem and root length, leaf area and
number, biomass, chlorophyll content, photosynthesis, etc. In the liter­
ature, the effects of MBNPs on the plant growth at various growth stages
are documented in many crops. In an investigation, the effect of TiO2
NPs (0, 50, 100, and 200 mg/L) was performed on agronomic traits of
Dracocephalum moldavica (Moldavian balm) grown under various
salinity levels (0, 50, and 100 mM NaCl). Results reported that TiO2 NPs’
application at 100 mg/L under all salinity levels improved all agronomic
traits compared to the growth of plants under salinity without TiO2 NPs
treatment (Gohari et al., 2020). In a comparative study, the toxicity of
ZnO and TiO2 NPs was reconnoitered on in vitro grown seedlings of the
Vigna angularis (red bean) plant. Seedlings were exposed to ZnO and
TiO2 NPs suspensions (0–200 μg/mL) for two weeks in hydroponic
media. Results indicated that ZnO NPs significantly inhibited the phys­
iological parameters of plants in a dose-dependent manner due to their
high solubility and accumulation in root tissues, while TiO2 NPs had a
beneficial effect on the plant physiology with increasing concentration,
resulting in enhanced growth. Moreover, ZnO NPs considerably reduced
chlorophyll content due to oxidative stress while no detrimental effect
was found in TiO2 NPs-treated plant seedlings (Jahan et al., 2018). In an
in vivo study of ZnO and CuO NPs on Brassica nigra, ZnO (41 nm) and
CuO (47 nm) NPs were applied to the synthetic soil at 200, 400, and
600 mg/kg and 12.5, 25, and 50 mg/kg concentration, respectively.
Results concluded that both NPs significantly enhanced growth profile
R. Javed et al. Ecotoxicology and Environmental Safety 271 (2024) 115992

Table 1
The effects of metal and metal oxide NPs on different crop plants.
Nanoparticles Size of Plant Concentration Exposure Application Beneficial Effect Adverse Effect Reference
(NPs) NPs duration method

Metal NPs
Ag NPs 74 nm Citrus 25, 50, 75, Exogenously Foliar spray Boost chlorophyll and Decrease in proline (Raza et al.,
reticulate 100 mg/L applied twice at carotenoid content, content 2023)
(Kinnow) 14-d intervals Increase SOD, POD, CAT,
before the phenolic and flavonoid
flowering stage content
Ag NPs 50- Pennisetum 0, 10, 20, NPs treatment Soil media 20 mM Ag NPs improve Salt stress restrict (Khan et al.,
100 nm glaucum 30 mM for 20 d water + proline content, growth, Decline in 2020)
(Pearl millet) NPs, NaCl treatment Improve antioxidant fresh/dry weight,
0, 120, for additional 10 enzymes activities Decrease Na+, Na+/K+
150 mM NaCl d Enhance H2O2 and MDA ratio
content Increase K+
Ag NPs 40 nm Vicia faba 0, 10, 50, 6h Seed priming Total soluble sugars Total soluble sugars (Abdel-Aziz
(Broad bean) 100 ppm increase at 100 ppm decrease in 10 and and Rizwan,
50 ppm, 2019)
Adversely affect
chloroplast
ultrastructure
Ag NPs Zataria 100, 150 mg/L 14 d MS media Elevate callus induction, (Mosavat
multiflora thymol and calvacrol et al., 2019)
content
Ag NPs 20 nm Pisum sativum 1000 and 15 d Hoagland Stimulate SOD and APX Inhibit GR and (Tripathi
(Garden pea) 3000 µM nutrient activity dehydroascorbate et al., 2017)
medium reductase (DHAR)
activity, Decline in
growth parameters
Cu NPs 47.47 nm Triticum 0, 25, 50, 30 d Soil mixture At 25, 50 mg/kg Decrease ROS and Cu (Noman et al.,
aestivum 100 mg/kg Cu NPs augment plant translocation to roots 2020)
(Wheat) growth, biomass, and and shoots
cellular antioxidants
Cu NPs 33 nm Cajanus cajan 20 ppm Colloidal Soil mixture Prominent increase in (Shende et al.,
(Pigeon pea) suspension of the length of root and 2017)
NPs given for 4- shoot as well as biomass
w at the interval
of 5-d
Cu NPs 100- Triticum Cu NPs: 0, 3, 5, Greenhouse Increase in chlorophyll (Ahmed et al.,
Ag NPs 1000 nm aestivum 7 mg/L experiment stability index and leaf K 2021b)
≤ 10 nm (Wheat) Ag NPs: 0, 10, content in plants
20, 30 mg/L
Zn NPs < 40 nm Spinacia 0.1%, 0.2%, 12 h for priming Seed Reduce H2O2, MDA, (Zafar et al.,
oleracea 0.3% Foliar priming, anthocyanin content, 2022)
(spinach) application after Foliar spray soluble proteins,
30 d chlorophyll content,
ascorbic acid, sugars,
TPC
Metal Oxide NPs
ZnO NPs Spinacia 0.0, 50, 100, 24 h In vivo Increase germination (Aly et al.,
oleracea 200 ppm method percentage, chlorophyll, 2023)
(Spinach) carotenoid, and proline
content,
Improve nutritional
value
ZnO NPs < 50 nm Hordeum 300 and 7d In vitro Increase in activity of (Azarin et al.,
vulgare 2000 mg/L method SOD, CAT, glutathione S- 2022)
(Barley) transferase (GST) and GR
ZnO NPs 24 nm Brassica 0, 50, 100, 200, 2h Seed priming Enhance seed (Awan et al.,
oleracea 400, 800, germination, root/shoot 2021)
(Broccoli) 1000 µg/L length,
increase chlorophyll,
proline, sugar, and
phenolic content
ZnO NPs 18 nm Triticum 1.7 mg Zn/kg 2 wk In vivo Act as source of (Dimkpa
aestivum method micronutrient and et al., 2020)
(Winter reduce drought stress
wheat)
ZnO NPs 34 nm Stevia 0, 0.1, 1.0, 10, 4 wk In vivo At 1 mg/L, significantly At 1000 mg/L, (Javed et al.,
rebaudiana 100, 1000 mg/ method enhanced steviol decrease production of 2017)
Bertoni L glycosides, increased secondary metabolites
(Honey Leaf) shoot formation, DPPH and anti-oxidant
scavenging activity, TAC activities.
(total antioxidant Decline in
capacity), TRP (reducing physiological
parameters
(continued on next page)

7
R. Javed et al. Ecotoxicology and Environmental Safety 271 (2024) 115992

Table 1 (continued )
Nanoparticles Size of Plant Concentration Exposure Application Beneficial Effect Adverse Effect Reference
(NPs) NPs duration method

power), phenolic and

flavonoid content.
CuO NPs 47 nm Trigonella 0, 2.5, 5, 15 d Tissue Increase DPPH activity, Copper acetate, PVP (Ain et al.,
PEG-capped 27 nm foenum- 10 mg/l culture TAC, TRP, TPC and TFC and PEG decrease seed 2018)
CuO NPs 27 nm graecum L technique in roots. germination, length
PVP-capped (Fenugreek) and weight parameters
CuO NPs
Fe2O3 NPs Triticum 10, 50, 21 d Hydroponic Decrease in growth, (Arikan et al.,
aestivum 100 mg/L Hoagland water content, loss of 2022)
(Wheat) solution photosynthetic
efficiency
Fe3O4 NPs 15-20 nm Hordeum 50, 100, 3 wk Hydroponic Improve germination (Tombuloglu
γ-Fe2O3 8-10 nm vulgare 200 mg/L medium rate, biomass, iron et al., 2022)
(Barley) content
Fe2O3 NPs 20-40 nm Triticum 500 mg/L 3 wk Hydroponic NPs act as a source of Fe, (Al-Amri
aestivum medium Enhance synthesis of et al., 2020)
(Wheat) chlorophyll
Fe-O NPs 35-45 nm Helianthus 1 or 2% 5 wk Soil spiked Increase in CAT, SOD, Decrease in (Mohammadi
annuus with NPs APX, and ascorbate accumulation of Cr et al., 2020)
(Sunflower)
SiO2 Graphene- Ammi visnaga 0, 15, 25 mg/L 3 wk Callus GO-SiO2 NPs increase (Golkar et al.,
oxide SiO2 L. (Apiaceae) Culture TPC, CAT, GPX and APX 2023)
NPs Enhanced H2O2, TFC and
FRAP at 25 mg/L SiO2
and 15 mg/L SiO2
Mn2O3 NPs 50 nm Capsicum 0.1, 0.5, 4-6 h Seeds soaked Increase root length and (Ye et al.,
annuum 1.0 mg/L in NPs act as source of Mn, i.e., 2020)
(Jalapenno suspension micronutrient
pepper)
Mn2O3 NPs 30 nm Atropa 25 mg/L 45 d Solid media Increase growth as well (Tian et al.,
belladonna as biomass and 2018)
(Deadly secondary metabolism
nightshade) (production of phenolic
compounds and
alkaloids)
Uncoated CeO2 19 nm Glycine max 100 mg/kg 3 wk Soil media Accelerate soybean (Cao et al.,
NPs: PVP 10 nm (Soybean) growth and rate of 2017)
coated CeO2 photosynthesis
NPs:
TiO2 NPs 6.5 nm Zea mays 100 mg/L 14 d Hoagland Activation of glutathione (Lian et al.,
(Maize) solution S-transferase, SOD, and 2020)
amino acids synthesis
TiO2 NPs Oryza sativa 5, 10, 20, 4 wk Foliar spray Enhance activity of APX, (Rizwan et al.,
(Rice) 30 mg/L CAT, and POD, Decrease 2019)
electrolyte leakage

POD: Peroxidase; SOD: Superoxide dismutase; CAT: Catalase; APX: Ascorbate peroxidase; MDA: Malondialdehyde; GPX: Glutathione peroxidase; GST: Glutathione S-
transferase; GR: Glutathione reductase; DHAR: Dehydroascorbate reductase.

of the plant while decreasing its primary root length in a
concentration-dependent manner (Zafar et al., 2020). In a recent study,
the effect of cerium oxide NPs (CeO2, 50 mg/L), and cerium oxide sal­
icylic acid NPs (CeO2: SA; 100 µM + 50 mg/L) along with manure
addition (0%, 10%, and 20% of v/v of soil) was studied on growth and
the physiological responses of Aloe vera. The combination of CeO2:
SA-NPs with 20% manure treatment had the highest growth index,
indicating that the foliar CeO2: SA-NPs along with the co-application of
soil-based cow manure stimulated the physiological responses and
growth of Aloe vera (Hassanpouraghdam et al., 2022). It can be
concluded that although MNPs and MONPs have both beneficial and
detrimental effects on plant’s growth and morphogenesis, the response
could be different according to the applied dose, exposure time, plant
species, and other experimental conditions.
2.14. In Vitro regeneration and induction of callus
A recent study reported increased phenolic and glycosidic content in
Vigna radiata (mung bean) when in vitro callus cultures of mungbean
were exposed to ZnO and CuO NPs at 0.5 mg/L concentration on MS
growth medium (Iqbal et al., 2022). Results of in vitro cultures of
Medicago sativa (alfalfa) exposure to 0.8 ppm CuO, ZnO, and CaO NPs
with and without salt stress claimed that CaO NPs significantly
enhanced callus induction and antioxidant enzymes activities as well as
H2O2 content in plants without salt stress. However, the best response in
protein rate and MDA activity was detected in callus treated with CuO
NPs along with 50 mM NaCl. It might be due to the effect of NPs that
diminished the detrimental impact of NaCl stress by eliminating stress
severity in the callus cells (Simsek et al., 2021). In another study, Fagonia
indica grown in an in vitro condition was supplemented with iron doped
ZnO NPs in a concentration range of 15.62–250 µg/mL. The results
depicted that Fe-ZnO-NPs displayed higher levels of phenolic, flavonoid,
and antioxidant activity at 125, 62.5, and 125 µg/mL, respectively,
attributed to the elicitation effect of Fe-ZnO-NPs (Khan et al., 2021). In
an investigation, leaf explants of Chrysanthemum cultured on the MS
medium augmented with 20 nm Ag NPs at concentrations of 0, 50, and
100 mg/L. Results found that application of 100 mg/L NPs suppressed
the formation of the adventitious shoots and shoots regeneration,
biosynthesis of metabolites, and modified the activity of the antioxidant
enzymatic defense system (Tymoszuk and Kulus, 2022). Hence, both the
in vitro regeneration and callus induction were estimated to be either
positively or negatively affected by different MBNPs depending on
numerous parameters.

8
R. Javed et al.
2.15. Nutritional quality
Recently, Yan et al. (2022) treated Oryza sativa (rice) plant to a
relatively low dose (1 and 10 mg/kg soil) of Ag NPs and found that
exposure to 1 mg/kg Ag NPs reduced Fe by 24.7% and Mo content by
31.8% in grains, whereas 10 mg/kg Ag NPs reduced K content by 19.4%
in grains. This report indicated that the exposure to Ag NPs has a
damaging effect on nutritional quality of the crops, especially the min­
eral nutrients. In an in vivo study, the impact of ZnO NPs on Oryza sativa
(rice) nutritional quality of grain and nutrient uptake along with other
parameters was evaluated at two concentrations (25 and 100 mg/kg).
The results showed that adding ZnO NPs at a concentration of 25 mg/kg
increased the P, N, and K content in the rice grain yield, and the Zn
concentration also increased to 13.5–39.4%. The results revealed that
ZnO NPs serve as high-performance fertilizers in rice production (Yang
et al., 2021). A study reported that under greenhouse conditions, Allium
fistulosum (green onion) was grown in soil that had been amended with
varying concentrations (75, 150, 300, and 600 mg/kg) of CuO NPs, bulk
CuO, and CuSO4. Plants exposed to 150 mg/kg concentration of CuO
NPs increased the allicin and other essential elements such as Mg, Fe, Ca
and Ni, consequently upgraded root nutrient quality compared to bulk,
CuSO4, and control (Wang et al., 2020). In a comparative study, the
influence of Cu and CuO NPs was analyzed on hydroponically grown
Brassica campestris (mustard) plant under Cd stress. Single Cu NPs
(20 mg/L), CuO NPs (20 mg/L), Cd (2 mg/L), and combined Cd+Cu NPs
(2 +20 mg/L), Cd+CuO NPs (2 +20 mg/L) treatments were supple­
mented to Hoagland nutrient solution. Results displayed that CuO
NPs+Cd upheld the uptake of Mn and Na elements by 24.3% and 28.3%,
respectively, while Cu NPs+Cd suppressed the uptake of Mn, Ca, Mg, K,
Fe, and Zn elements in leaves of plant (Wang et al., 2022).
2.16. Production of secondary metabolites
The usage of NPs as an elicitor of plant secondary metabolites has
been a common practice. Compared to primary metabolites which are
involved in plant’s growth and development, the secondary metabolites
play central roles in plants’ sustainability and their adaptation to the
environment (Erb and Kliebenstein, 2020). NPs, after entering into plant
cells control the breakdown of ROS and initiate downstream signaling
actions which are involved in primary and secondary metabolism. In
response to different biotic as well as abiotic stresses, plants increase the
synthesis of secondary metabolites which include terpenes, phenolics,
flavonoids, and alkaloids (Sun et al., 2020; Javed et al., 2022). Fig. 5
Fig. 5. Mechanism of activation of secondary metabolism by nanoparticles (NPs) stress.
9
Ecotoxicology and Environmental Safety 271 (2024) 115992
shows the possible mechanism of stimulation of secondary metabolites
in plants upon NPs’ exposure.
There are numerous studies of the improvement of secondary me­
tabolites in plants by MBNPs. In a study, an untargeted metabolic
approach was used to study the effect of Ag, Cu, Au, CuO, CeO2, ZnO,
and TiO2 NPs on Hypericum perforatum in a cell suspension culture.
Overall, MNPs enhanced secondary metabolites more than MONPs.
However, Ag and CuO NPs induced greater alterations in secondary
metabolism. Findings suggested that secondary metabolites elicitation
in plants vary among different types of MBNPs (Kruszka et al., 2022).
Moreover, Li et al. (2021) indicated that foliar application of ZnO
(50 nm) and SiO2 (20 nm) NPs at a concentration of 50 and 100 mg/L,
and 20 and 60 mg/L, respectively on Cucumis sativus (cucumber) seed­
lings improved the metabolomic profile of plants by enhancing the TFC
in the leaves, thus potentially regulating the stress resistance in the
plants. Conclusively, MNPs and MONPs stimulated the secondary
metabolism in various crops by an up-regulation of signaling pathways.
Besides, hairy root culture is a technique adapted to increase sec­
ondary metabolites production in medicinal plants. Hairy roots mimic
the phytochemical constitutions of parent plants having an additional
benefit of higher productivity and rate of growth (Halder et al., 2019).
Hairy roots are generated from Agrobacterium rhizogenes, hence these are
transformed roots that are involved in metabolic engineering of regu­
latory genes in the pathway of production of bioactive compounds. Their
properties like free of contamination and being genetically stable makes
them potent biotechnological system for the production of secondary
metabolites (Biswas et al., 2023). Metal-based NPs act as elicitors for
accumulation of therapeutic and pharmaceutical compounds in different
medicinal plants on small and large scale. Table 2 gives few recent ex­
amples of metal-based nanoelicitors for boosting hairy root culture and
enhanced production of secondary metabolites from highly valuable
medicinal plants.
2.17. Antioxidants (Enzymatic and Non-Enzymatic)
Oxidative stress caused by MBNPs is one of the environmental factors
that increase or decrease plant growth and functioning (Adeel et al.,
2020). Naturally, there is a balance of ROS production in plant struc­
tures such as chloroplast, mitochondria, and peroxisomes. These are
continually removed enzymatically and non-enzymatically by a complex
defense system in plants (Hussain et al., 2016; Roychoudhury and Tri­
pathi, 2019). A study reported the application of Ag NPs on the Camelina
sativa plant at a concentration of 0.5, 1, 2, 3, and 4 g/L via in vivo
R. Javed et al.
Table 2
The effects of metal-based NPs on different medicinal plants for induction of hairy roots and production of secondary metabolites.
Nanoparticles Optimum conditions Medicinal plant
(NPs)
Ag NPs Hyoscyamus muticus
Ag NPs 30 mg/L, 7 d Salvia miltiorrhiza
Ag NPs 21 d Cucumis anguria
SiO2 NPs Dracocephalum
kotschyi
Ag-SiO2 NPs 20 d Artemisia annua
Fe, ZnO NPs Trigonella foenum-
graecum
ZnO NPs 100 mg/L, 24 h Hyoscyamus
reticulatus
FeO NPs Hyoscyamin: 900 mg/L, 24 h Hyoscyamus
Scopolamine: 450 mg/L, 48 h reticulatus
SOD: Superoxide dismutase; CAT: Catalase; APX: Ascorbate peroxidase; MDA: Malondialdehyde; TPC: Total phenolic content; TFC: Total flavonoid content.
method. Oxidative stress induced by Ag NPs was indicated by an in­
crease in the accumulation of MDA, H2O2, and methylglyoxal, as well as
up-regulated the activity of antioxidant enzymes, for instance, SOD,
APX, CAT, and GR and glyoxalase (GLO) system enzymes (Mirmoeini
et al., 2021). A relative study displayed higher efficiency of ZnO NPs as
compared to bulk ZnO in Portulaca oleracea (Purselane) where seeds
were treated with four concentrations (0, 10, 100, and 500 mg/L) of
both ZnO NPs and bulk ZnO. Findings depicted that the application of
ZnO NPs up-regulated the activity of POD enzymes more compared to
bulk ZnO in the leaves where the highest activity was detected at
500 mg/L of ZnO NPs. Thus, it was manifested that MBNPs affect the
activity of POD more efficiently than its bulk counterparts. In contrast,
the activity of CAT enzymes decreased in ZnO NPs-treated plants while
increased in bulk ZnO (Iziy et al., 2019). In another experiment,
simultaneous application of salinity (0, 50, and 100 mM NaCl) and
Fe2O3 NPs (20 nm) at a concentration of 0, 30, 60, and 90 ppm was
probed on the Dracocephalum moldavica (Moldavian balm) under
greenhouse conditions. Results presented that under salinity stress, NPs
at 60 ppm significantly increased the activity of GPX, APX, CAT, and
glutathione reductase (GR) in the shoot and root of the plants
concluding the importance of non-enzymatic system in plants so as to
counteract the noxious effect of salinity stress (Moradbeygi et al., 2020).
Thus, enzymatic as well as non-enzymatic antioxidants either increased
or decreased depending on the conditions of other stresses interfering
with the NPs’ stress. However, most of the studies represented amelio­
ration of enzymes and antioxidants acting as defense factories.
2.18. Alterations at molecular level
MNPs and MONPs could induce genotoxicity in plants that impair
the genetic information within the cells causing mutations. Such mu­
tations in plants are monitored via randomly amplified polymorphic
DNA (RAPD) analysis which could detect variations in genome profiles
of plants. According to AlQuraidi et al. (2019), 200, 400, and 800 mg/L
of Cu NPs concentration when exposed to hydroponically grown Cor­
iandrum sativum (coriander) plant resulted in genotoxicity at 400 and
800 mg/L concentration evaluated by the RAPD technique. In a study
reported by Kokina et al. (2020), Medicago falcate (medicine yellow) was
grown hydroponically with Fe3O4 NPs added at a concentration of 1, 2,
and 4 mg/L for about five weeks. This study’s RAPD analysis confirmed
that Fe3O4 NPs demonstrated genotoxic effect on plants that resulted in
genomic DNA modifications. Moreover, the increase in miR159c
expression indicated that these NPs could be used for alleviating plant
resistance against pathogenic fungi. In aother study, the genotoxic po­
tentialities of phytofabricated ZnO NPs were investigated in the leaf
extract of Ipomea obscura (morning glory) plant at a concentration of 0.2,
Ecotoxicology and Environmental Safety 271 (2024) 115992
Secondary metabolite Effect Reference
Hyoscyamine, Scopolamine Antimicrobial activity (Abdelkawy et al.,
2023)
Tanshinone Enhancement of SOD, CAT, (Ma et al., 2020)
APX
Hydroxycinnami, Antimicrobial, antioxidant, (Chung et al., 2018)
Hydroxybenzoic acids anticancer activity
Rosmarinic acid, Xanthomicro, Increased expression of pal and (Nourozi et al., 2019)
Cirsimaritin ras
Artimisinin Increase of MDA and CAT (Zhang et al., 2013)
Trigonelline Increase of TPC and TFC (Tariverdizadeh et al.,
2021)
Tropane alkaloid Highest h6h gene expression (Asl et al., 2018)
Hyoscyamin, Scopolamine Enhancement of antioxidant (Moharrami et al.,
enzymes 2017)
0.4, 0.6, 0.8, and 1 mg/mL. Results declared a decrease in the mitotic
index in a concentration-dependent manner, indicating the potential
inhibitory effect against cell division, eventually leading to apoptosis
(Murali et al., 2021). Comparatively, upon revelation to Al2O3 and ZnO
NPs, the gene expression of selected aquaporins (HvTip1;1 and
HvPip1;1), also transcription factors from Hordeum vulgare (barley) was
examined. The gene expression profiling showed that genes differen­
tially regulated as aquaporin genes were greater in leaves compared to
control, while a gradual decrease was observed in roots upon Al2O3 NPs
exposure. Moreover, the effect of ZnO NPs on gene expression level was
distinctive from that of Al2O3 NPs (Akdemir, 2021).
3. Conclusion and future outlooks
Nano-agri revolution and development of “smart plants” is a must to
ameliorate the efficiency of crops. Despite numerous studies docu­
mented in this field, it is still in its emerging stages and facing various
challenges and shortcomings. A comprehensive literature review of
previous and current studies about the interplay between metal-based
nanoparticles (MBNPs) and plants was thoroughly conducted. The re­
view revealed a series of physiological, biochemical, and molecular
events involved in this interplay. The photosynthesis, growth parame­
ters, antioxidants, etc., were either stimulated or mitigated on exposure
to MBNPs. There are several knowledge gaps related to this topic that
must be addressed to achieve a thorough and deep understanding of this
interaction, particularly in terms of ROS-dependent damage. The
following approaches will help enhance favorable effects and reduce
hazardous effects, ultimately ensuring environmental safety and
security:
• The key factors influencing the interaction between MBNPs and plant
should be managed to the non-significant level in order to avoid
toxicity and ensure the safety of these NPs. Of utmost significance is
the regulation of metal ion release and surface morphology of both
MNPs and MONPs as they play a role in their toxic potential.
• Majority of the recent research have been conducted under the
controlled conditions, such as growth chambers and greenhouses.
Therefore, future research is required to examine the application of
MBNPs on plants under field conditions. This will enable a proper
assessment of the benefits and risks associated with their use.
• To enhance the understanding of the effects of MBNPs on plants,
comparative studies should be designed to investigate the differences
in plant exposure through foliar spray or soil amendment of MNPs
and MONPs. Moreover, life cycle and transgenerational studies
should be performed to study their long-term effects.
10
R. Javed et al.
• Research must be conducted within the framework of risk assessment
concerning the application of MBNPs’ to plants and their potential
release into the food chain, affecting animals, humans, and the
environment. In brief, trophic transfer of MNPs and MONPs should
be elucidated in detail.
Funding
We greatly acknowledge the support of the Government of Canada’s
New Frontiers in Research Fund [NFRF-2020-00452] in supporting this
study.
CRediT authorship contribution statement
Bilal Muhammad: Writing – original draft, Visualization, Software,
Methodology, Formal analysis, Data curation. Galagedara Lakshman:
Writing – review & editing, Validation, Supervision, Project adminis­
tration, Investigation, Formal analysis. Abbey Lord: Writing – review &
editing, Validation, Formal analysis. Cheema Mumtaz: Writing – re­
view & editing, Supervision, Resources, Project administration, Funding
acquisition. Javed Rabia: Writing – review & editing, Writing – original
draft, Visualization, Validation, Software, Project administration,
Methodology, Investigation, Formal analysis, Data curation, Conceptu­
alization. Khan Bakhtawar: Writing – original draft, Visualization,
Software, Methodology, Formal analysis, Data curation. Sharafat
Uzma: Writing – original draft, Methodology, Formal analysis, Data
curation.
Declaration of Competing Interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influence
the work reported in this paper.
Data availability
No data was used for the research described in the article.
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