Trends in
Plant Science
Spotlight
Silica nanoparticles:
the rising star in plant
disease protection
Nidhi Kandhol, 1
Vijay Pratap Singh, 2
José Peralta-Videa , 3
Francisco J. Corpas , 4,* and
Durgesh Kumar Tripathi 1,*
When applied exogenously, silica
(Si) can have a beneficial impact
on plants under biotic stress con-
ditions, as revealed by its recent
application in the form of nanoparti-
cles (SiO2NPs) to induce pathogen
resistance (El-Shetehy et al.). This
opens up a new window of research
into combating the devastating
effects of biotic stresses.
Nanotechnology, a rapidly developing
field of research, can be applied in
many areas, including agriculture. The
use of nanoparticles (NPs), combined
with the controlled release of com-
pounds such as nitric oxide (NO) and a
diverse range of metals, has been the
subject of numerous studies [1–3]. The
application of silica (Si) is known to
have beneficial effects on various abiotic ingly, it has confirmed the effectiveness of
stresses, in many cases by affecting
antioxidant systems to alleviate oxida-
tive damage associated with these
stresses [4]. A recent study indicated
that SiO2NPs are capable of inducing
an immune response, called systemic
acquired resistance (SAR) [5], in plants; study provides clear evidence of interac-
this is similar to the response trig-
gered following an attack by some
pathogens [6]. This finding points to
a new role for SiO 2 NPs as effective,
sustainable, and economical biostimulants SAR is characterized by a basal defense (1600 mg SiO2 l–1) and Si(OH)4 (1000 mg
in plants.
SiO2NPs endow arabidopsis
(Arabidopsis thaliana) plants with
pathogen resistance by activating
SAR
In order to establish a balance between
increasing food demand and a compara-
tively low food supply, while maintaining
food security, it is of paramount impor-
tance to develop new tools combatting
plant diseases and raising agricultural pro-
ductivity. Up to now, genetic engineering
has been the most effective method used
to enhance disease tolerance, as well as
the nutrient profile of plants under patho-
gen attack [7]. However, this technique is
complex, time-consuming, has a high
economic cost, and requires specific con-
ditions to achieve the desired results. The
assessment of suitability and toxicity of
nanotechnological applications to plants
under different conditions has recently
caught the attention of researchers. So
far, contentious observations have been
made, trying to understand the mode of
action of SiO2NPs in mediating stress
forbearance to plants [8], but a recent
approach by El-Shetehy et al. [5] has
provided new insight. El-Shetehy et al.
describe triggering SAR through the
dose-dependent treatment of leaves with
SiO2NPs. This has led to an investigation
of the capacity of SiO2NPs to boost the
resistance of Arabidopsis thaliana to the
bacterial pathogen Pseudomonas syringae
at both a local and systemic level. Accord-
SiO2NPs in activating SAR. Thus, a com-
parative analysis of the mode of action of
SiO2NPs and orthosilicic acid, Si(OH)4,
was carried out (Figure 1), while SiO2NPs
were also tested for controlled delivery of
nanoporous silicon (Si) to the plant. This
tions between SiO2NPs and plant leaves,
ranging from entry to distribution in the
apoplast at the subcellular level.
However, at higher concentrations, SiO2NPs
mechanism induced by either pathogen SiO2 l –1) are less effective at activating
attacks or the application of resistance-
inducing compounds such as benzo-
thiadiazole, which activate signal trans-
duction pathways leading to signal
transmission throughout the plant [9].
Phytohormone salicylic acid (SA) also
contributes to SAR by activation of
pathogenesis-related (PR) genes [9]. SAR
also involves a natural signaling cascade
to control plant pathogens and pests,
composed of SA, reactive oxygen species
(ROS), and NO [6]. To provide a mechanis-
tic insight into the processes involved
in SiO2NP-induced triggering of SAR,
bacterial growth on arabidopsis leaves
was measured to quantify local defense
responses to a virulent strain of P. syringae
under control, SiO2NP, and Si(OH)4 treat-
ment regimes. SiO2NPs were able to
reduce the number of bacteria eightfold
within 24 h as compared with the control
treatment, while treatment with Si(OH)4
also resulted in the induction of SAR
against the bacterial pathogen. In addi-
tion, the triggering of SAR by both
SiO2NPs and Si(OH)4 is dose-dependent.
In the case of SiO2NPs, maximum bacterial
inhibition (>90%) was observed at a con-
centration of 100 mg SiO2 l–1 as compared
with control plants treated with only HEPES
buffer or MgCl2. qPCR analysis of the
OPRF gene, encoding the bacterial outer
membrane protein F (OmpF), also provided
a good idea of bacterial biomass and
asserted the results of bacterial colony
counting in line with previous similar work
[10]. The armoring effect of SiO2NPs and
Si(OH)4 is not due to a direct toxic effect
on bacterial growth, as neither damaged
bacterial growth when cultured in vitro
with virulent P. syringae. This demon-
strates that SiO2NPs make plants patho-
genically resistant by activating defense
responses, rather than by inhibiting or
killing the pathogen.
Trends in Plant Science, January 2022, Vol. 27, No. 1 7
 Trends in Plant Science

release of Si(OH)4 after entering through

the stomata and their dispersal in the
spongy mesophyll. However, this protec-
tion is majorly mediated by the activation
of SA-dependent defense reactions by
intact particles that play a prominent role
in plant immunity [9]. Neither SiO2NPs
nor Si(OH)4 produce any local or systemic
disease resistance in SA induction-deficient
2 (sid2), an arabidopsis mutant deficient
in SA biosynthesis. The gene expression
of the SA-responsive marker genes,
AtPR-1 and AtPR-5, encoding PR-1 and
PR-5 proteins, has also been observed
to increase in response to SiO2NPs and
Si(OH)4, an increase that is comparatively
lower in the case of SiO2NPs. These analy-
ses confirm that SA-dependent pathways
are involved in SAR activation by SiO2NPs
and Si(OH)4.
Trends in Plant Science

Figure 1. Systemic acquired resistance (SAR) induced in Arabidopsis thaliana by SiO2 nanoparticles Given the evidence provided by this study
(NPs) and Si(OH)4 via salicylic acid (SA) signaling for fostering plant defense against Pseudomonas [5], SiO2NPs are clearly safer for plants
syringae. Slowly released Si(OH)4 from SiO2NPs (some cling between guard cells) into cells induces local than the direct application of Si(OH)4.
defense by triggering SA signaling; signals move to distant organs of plants and stimulate SAR for subsequent
Also, as the NPs remaining in the extracel-
defense. Directly applied Si(OH)4 also shows similar SAR activation, but excess SA signaling at higher
concentrations, which leads to less effective SAR. Low concentration: 100 mg SiO2 l−1; high concentration: lular air spaces of the spongy mesophyll
1000 mg SiO2 l−1. (Drawn as per the results described by El-Shetehy et al., 2021 [5]). do not risk being washed away from leaf
surfaces, they can function more effec-
tively; further in-depth research into the
SAR and even increase bacterial infec- showed that NPs measuring ~50–70 nm physical effects of NPs on leaves needs
tion, which is more severe in the case are capable of entering the leaf only to be carried out. As SAR is mainly regu-
of Si(OH)4, possibly due to increased through the stomata, refuting the earlier lated by SA and PR genes [9], defense-
oxidative stress. This is confirmed by the speculations that NPs are capable of pene- related genes in other plants of agronomical
much higher gene expression levels of trating the cuticle [12]. SiO2NP uptake by importance, containing different pathogens,
AtHSP17.4C1, a cytosolic class I small the cell wall, as well as cell-to-cell translo- will also need to be studied. The role of
heat shock protein (sHSP) induced by cation via plasmodesmata, has been partially closed stomata and NP interactions
abiotic, biotic, and oxidative stresses found to be obstructed due to the size of with the spongy mesophyll in initiating
[11], in the case of Si(OH)4 treatment NPs [12]. SiO 2 NPs, which are dis- SA-related responses should also be
at concentrations of 100 and 1000 mg persed in the extracellular air spaces investigated. Beneficial NPs as a tool to
SiO2 l–1 but not in the case of SiO2NPs, of the spongy mesophyll without pen- enhance disease resistance hold the
which led to an insignificant increase etrating any cell walls, have also been potential to bypass the risks associated
in gene expression. But, impairment of observed to change in the guard cells with genetic modification of plants, such
SAR by SiO2NPs, only under very high of leaves, thus preventing full stomatal as unexpected genetic variations in the
concentrations, might be credited to closure. Even after 2 days of treatment, next generation and undesired inter-
the superfluous release of Si(OH) 4 from numerous intact SiO2NPs remain present action with environment if applied after
SiO2NPs resulting in oxidative stress or in the stomata, suggesting that their disso- passing complete risk evaluation. The
redundant stomatal clogging by SiO2NPs lution in plants is slow. favorable role exhibited here by SiO2
that ultimately disrupts evapotranspira- NPs does not approve their successful
tion. Electron microscope analysis of The overall protective effect of SiO2NPs commercialization. Possible risks in-
treatment with SiO2NPs after 2 days can be partly attributed to their slow volved must be considered before the

8 Trends in Plant Science, January 2022, Vol. 27, No. 1

Trends in Plant Science

3. Mahakham, W. et al. (2017) Nanopriming technology for

use of SiO2NPs as biostimulants, pesti- Declaration of interests enhancing germination and starch metabolism of aged
cides, herbicides, or fertilizers; in partic- No interests are declared. rice seeds using phytosynthesized silver nanoparticles.
Sci. Rep. 7, 8263
ular, possible effects on human health 4. Tripathi, D.K. et al. (2021) Silicon crosstalk with reactive
and on microorganisms in soil–plant 1Amity Institute of Organic Agriculture, Amity University Uttar oxygen species, phytohormones and other signaling mol-
Pradesh, I 2 Block, 5th Floor, AUUP Campus Sector-125, ecules. J. Hazard. Mater. 408, 124820
systems require investigation. Neverthe- Noida 201313, India 5. El-Shetehy, M. et al. (2021) Silica nanoparticles enhance dis-
less, this research has outlined many 2Plant Physiology Laboratory, Department of Botany, C.M.P. ease resistance in Arabidopsis plants. Nat. Nanotechnol. 16,
Degree College, A Constituent Post Graduate College of 344–353
pointers as to how SiO 2 NPs can sus- University of Allahabad, Prayagraj 211002, India 6. Gao, H. et al. (2021) Signals in systemic acquired resis-
tainably protect plants against patho- 3Department of Chemistry and Biochemistry, The University of tance of plants against microbial pathogens. Mol. Biol.
Texas at El Paso, 500 West University Avenue, El Paso, Rep. 48, 3747–3759
gens through the controlled induction of TX 79968, USA 7. Kumar, S. et al. (2020) Nanovehicles for plant modifica-
SAR, with a minimum amount of adverse 4Group of Antioxidants, Free Radicals and Nitric Oxide in tions towards pest-and disease-resistance traits. Trends
Biotechnology, Food and Agriculture, Department of Plant Sci. 25, 198–212
effects. Biochemistry and Cell and Molecular Biology of Plants, Estación 8. Mukarram, M. et al. (2021) Silicon nanoparticles elicit an
Experimental del Zaidín, Consejo Superior de Investigaciones increase in lemongrass (Cymbopogonflexuosus (Steud.)
Científicas (CSIC), Profesor Albareda 1, 18008, Granada, Spain Wats) agronomic parameters with a higher essential oil
Acknowledgments yield. J. Hazard. Mater. 412, 125254
D.K.T. is thankful to Science and Engineering 9. Durrant, W.E. and Dong, X. (2004) Systemic acquired re-
*Correspondence:
Research Board, New Delhi for providing research javier.corpas@eez.csic.es (F.J. Corpas) and sistance. Annu. Rev. Phytopathol. 42, 185–209
dktripathiau@gmail.com (D.K. Tripathi). 10. Ross, A. and Somssich, I.E. (2016) A DNA-based real-time
grants (SRG/2020/002309) to carry out this work.
PCR assay for robust growth quantification of the bacterial
F.J.C.’s research is supported by a European https://doi.org/10.1016/j.tplants.2021.10.007 pathogen Pseudomonas syringae on Arabidopsis thaliana.
Regional Development Fund cofinanced grant from Plant Methods 12, 1–8
© 2021 Elsevier Ltd. All rights reserved. 11. Sewelam, N. et al. (2019) The AtHSP17.4C1 gene expres-
the Ministry of Economy and Competitiveness
sion is mediated by diverse signals that link biotic and abiotic
(PID 2019-103924 GB-I00), the Plan Andaluz de stress factors with ROS and can be a useful molecular
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