Plant Physiology and Biochemistry 201 (2023) 107831

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Plant Physiology and Biochemistry

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

Multifactorial role of nanoparticles in alleviating environmental stresses for

sustainable crop production and protection
Biswajit Pramanik a, Puranjoy Sar a, **, Ruchi Bharti b, Rahul Kumar Gupta b,
Shampa Purkayastha c, Somya Sinha d, Sourav Chattaraj e, Debasis Mitra e, *
a
Department of Genetics and Plant Breeding, Palli Siksha Bhavana (Institute of Agriculture), Visva-Bharati, 731236, Sriniketan, West Bengal, India
b
Department of Agronomy, Palli Siksha Bhavana (Institute of Agriculture), Visva-Bharati, 731236, Sriniketan, West Bengal, India
c
Department of Genetics and Plant Breeding and Seed Science and Technology, Centurion University of Technology and Management, Paralekhamundi, 761211, Odisha,
India
d
Department of Biotechnology, Graphic Era (Deemed to be University), Dehradun, 248 002, Uttarakhand, India
e
Department of Microbiology, Raiganj University, Raiganj, 733134, Uttar Dinajpur, West Bengal, India

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

Handling Editor: Shivendra Sahi In the era of dire environmental fluctuations, plants undergo several stressors during their life span, which
severely impact their development and overall growth in negative aspects. Abiotic stress factors, especially
Keywords: moisture stress i.e shortage (drought) or excess (flooding), salinity, temperature divergence (i.e. heat and cold
Nanotechnology stress), heavy metal toxicity, etc. create osmotic and ionic imbalance inside the plant cells, which ultimately lead
Nanoparticles
to devastating crop yield, sometimes crop failure. Apart from the array of abiotic stresses, various biotic stress
Abiotic stress
caused by pathogens, insects, and nematodes also affect production. Therefore, to combat these major challenges
Biotic stress
Plant growth in order to increase production, several novel strategies have been adapted, among which the use of nano­
Sustainable agriculture particles (NPs) i.e. nanotechnology is becoming an emerging tool in various facets of the current agriculture
system, nowadays. This present review will elaborately depict the deployment and mechanisms of different NPs
to withstand these biotic and abiotic stresses, along with a brief overview and indication of the future research
works to be oriented based on the steps provided for future research in advance NPs application through the
sustainable way.

1. Introduction rigidity, reactivity and ability to absorption in visible light in order to

impart different colours. Since 1960s, NPs become popular element of
Agriculture is the primary pillar for the basic development of any research in the field of agriculture. It is a promising agent for plant
developing countries. But in current scenario, due to environmental growth, due to its wide array of uses in various inputs like fertilizer,
pollution and climate change, agriculture is facing severe adverse situ­ fungicides, weedicides, pesticides, etc. In the food industry, it helps in
ation to fulfill the demands of the enormously increasing global popu­ both maintaining the quality as well as locking the nutritive value of
lation (Pouratashi and Iravani, 2012). Nanoparticles (NPs), derived from food. Bottom-up and Top-down are the two approaches for synthesis of
a greek word, which means dwarf. These are the tiny particles with a size NPs (Wang and Xia, 2004). Rather, an eco-friendly green method has
range of 1–100 nm. It is generally composed of three layers, outer sur­ been developed to synthesize nanoparticles, which both increases the
face layer consists of metal ion, middle shell layer and the inner core, agricultural output without hampering the environment (Ghidan et al.,
although surfactants or polymers on the outer surface can also be found 2017; Mitra et al., 2023) and helps in reducing the harmful environ­
in the structure of NPs. Sometimes a single metallic core also forms the mental pollutants (Huang et al., 2015). Nanotechnology is multidisci­
entire structure of this molecule (Shin et al., 2016). NPs having unique plinary in action viz. nanotechnology, nanobiotechnology,
physical and chemical properties due to their smaller shapes and sizes, nanotoxicology, nanoponics. Even it has a potential application in the
which ultimately gives them a comparatively higher surface area, field of plant breeding and genetic transformation, as reported by Roco

* Corresponding author.
** Corresponding author.
E-mail addresses: [email protected] (P. Sar), [email protected], [email protected] (D. Mitra).

https://doi.org/10.1016/j.plaphy.2023.107831
Received 30 January 2023; Received in revised form 22 April 2023; Accepted 8 June 2023
Available online 29 June 2023
0981-9428/© 2023 Elsevier Masson SAS. All rights reserved.
B. Pramanik et al. Plant Physiology and Biochemistry 201 (2023) 107831

(2003). The application of nanosensor helps in reducing the agro­ nano-based coatings. Nowadays, for different nanoparticles different
chemical residue in the field. Further, diversified facets of nano­ slandered leaching methods according to the regulations of ISO
biotechnology have been deployed for early stage disease diagnosis, 2812:2018 and ISO 12457–3:2004 are used to taste paints. For further
escalated nutrient acquisition ability of plants, gene transfer to develop analysis of leached substances, several performing liquid chromatog­
disease- or insect-resistant varieties, increased self-life of food products, raphy (HPLC) combined with UV detection, liquid
nanodevice-mediated genome editing and many more. It also has a wide chromatography-mass spectrometry (LC/MS), and LC/MS/MS were
spectrum of applicability in agriculture from production, storage, pro­ used with high reproducibility (Brunelli et al., 2021). Hence, proper
cessing to transport. In conventional era, pest attack diminishes nearly quantification of nanoparticles is crucial in the post-application process.
1/3rd of the crop production each year (Tudi et al., 2021). Hence, to The exposure to different leaching tests confirms the effects of exposed
overcome this issue, nanotechnology has emerged as a potent innova­ nanoparticles in terms of environmental safety and health hazards
tion to bring forth a revolutionary approach in this field without (Zhang et al., 2019).
compromising food security. It became the modern agricultural practice NPs also help to combat several biotic and abiotic stress conditions
for plant health management. The knowledge of plant and nanoparticles by upregulating different stress related genes. In last few years, NPs
interaction opens a new arena for crop improvement (Mittal et al., 2020; drew much attention towards study of waterlogging stress condition
Kandhol et al., 2022a; Parveen et al., 2022). (Tariq et al., 2021). Different NPs have different effects on
NPs possess strong reactivity due to its greater surface to volume morpho-physiology of plant organs, which efficiently increase the stress
ratio, and their physiochemical properties highlighted the vitality to test tolerance capacity, eventually (Hassanisaadi et al., 2022). NPs, made up
extant of their toxicity inside plant and human bodies, before exper­ of either organic or inorganic substances, helps to activate certain an­
imenting them into the market (Vishwakarma et al., 2018; Usman et al., tioxidants in order to protect the plants from oxidation during adverse
2020). Therefore, a controlled delivery system to alter the physiology of climatic condition (Chandel et al., 2022). Basically, NPs having versatile
plants to some extent is needed along with proper knowledge about applications to mitigate stress condition. The application of NPs offers
plant-NPs interaction (Fraceto et al., 2016). The interaction of NPs with several positive changes including regulation of gene expression, chlo­
cell wall creates pores, which facilitate their penetration and trans­ rophyll content, enhances the water uptake, controls root hydraulic
portation via apoplastic and symplastic pathway (Carlson et al., 2008). conductance, detoxification of reactive oxygen species and ultimately
On the other way, negatively charged cell wall promotes ion exchange, increases the adaptation of cultivated plants against stressful condition
which helps in penetration of cationic NPs faster than anionic one (Das and Das, 2019). Hence, the present review predominantly focusses
(Meychik et al., 2005). Notably, dicots have more cuticles than monocot, on the implementation of various NPs on withstanding various stress
which eventually ease the way of NPs uptake. Also, bacterial and conditions (both biotic as well as abiotic) along with a small interference
mycorrhizal association in rhizosphere aid their acquisition (Feng et al., about the application strategies and mechanisms followed. The molec­
2013; Vala et al., 2018). Hence, its implementation in the field of agri­ ular, and cellular levels of biological research concepts and agricultural
culture is a new frontier which will enhance the quality of life as well as application uses are made possible by the convergence of nanotech­
makes farming more scientific and target oriented. That’s why, a proper nology science. The advancement of nanobiotechnology through current
research system with significant government policies is required for and upcoming research efforts may eventually result in the creation of
extension knowledge in this new area of nanotechnology application in novel biomolecular, nano-biosensor, nano-fertilizer, and biodegradable
agriculture. nano-fiber technologies. The overview role and applications of nano­
The development of nanoparticles as well as their delivery through particles are attempted to be explained in this review, which begins with
specific methods inside plants is crucial. The current delivery system has nanotechnology’s modes of action, biochemical and molecular in­
several loopholes as it is applicable to a specific host, causes toxicity to terventions, biosynthesis strategies, the various benefits in mitigating
tissues, and has low transformation efficiencies (Demirer et al., 2019). plant stress, and ends with its use as artificially intelligent amalgamation
Although a nanoparticle can enter inside a plant through both active and in plant stress nanotechnology. In summary, this in-depth research has
passive transport, using the nanoparticles as the nanocarriers, especially thoroughly studied the application of a variety of NPs towards tye
in plants is a challenging task because of the low rate of endocytosis in management of plant stress together with a novel touch of machine
plants, the presence of cell walls and their different chemical composi­ learning and plant genetic engineering. Additionally, this thorough
tion (Torney et al., 2007; Silva et al., 2010; Chang et al., 2013). Recently investigation unearths a more modern strategy for introducing artificial
there is much interest among the scientific communities to use nano­ intelligence to agriculture, with a particular focus on stress biology.
particles as nanocarriers for required genetic transformation (Naqvi
et al., 2012) with the advantages of high throughput, low cytotoxicity, 2. Nanoparticles: an overview and applications in agriculture
and wide range of its applicability (Liu et al., 2009). While
Nanoparticle-based delivery methods available now a day are expensive, Nanoparticles are something whose dimension is generally measured
required specialist to handle, and due to high delivery pressure causes in nanoscale (1–100 nm). These can either be organic or inorganic,
damage to the targeted cell/tissues (Husaini et al., 2010). Lew et al. sometimes a fusion of both. Not only photochemical reactions, volcanic
(2018) suggested that the size of nanoparticles and the magnitude of eruptions, forest fires, erosion, etc. but also several plants and micro­
zeta potential are able to determine the distribution of subcellular organisms can produce NPs (Dahoumane et al., 2017). Due to their
nanoparticles via the mechanism of lipid exchange envelope penetra­ eco-friendly nature and synthesizing-simplicity, as compared to others,
tion. Also, for the first time, a universal localization mechanism lipid NPs derived from plants and microorganisms have recently drawn the
exchange envelope and penetration (LEEP) shows kinetic trapping of attention of scientists as a reliable biological source of green synthesis
different nanoparticles in chloroplast which predicts a critical particle (Panpatte et al., 2016). Just like alfalfa (Medicago sativa) and dhaincha
size below which the mechanism fails at all zeta potentials (Wong et al., (Sesbania sp.) derived gold nanoparticles (AuNPs), mustard (Brassica
2016). A species-independent diffusion-based passive biomolecule de­ juncea), alfalfa, and sunflower (Helianthus annus) also own the ability to
livery of genetic material for efficient DNA delivery without transgene manufacture inorganic nanomaterials made of copper (Cu), nickel (Ni),
integration and strong protein expression is an effective tool for cobalt (Co), zinc (Zn), and silver (Ag) (Ghormade et al., 2011; Sharma
target-based nanoparticle delivery in plants (Demirer et al., 2019). et al., 2007; Venkatachalam et al., 2015, 2017). Microorganisms like
Moreover, the aging process of the nanoparticles transforms the diatoms, Pseudomonas stuzeri, Desulfovibrio desulfurican, Clostridium
actual structure of the originally formed one due to environmental ef­ thermoaceticum, etc. are used to generate silicon (Si), gold (Au), zinc
fects (Mitrano et al., 2015; Zhang et al., 2017). Hence a standard sulphide (ZnS), and cadmium sulfide (CdS) nanoparticles. Among all
leaching method is required to estimate the NPs leaching from others, the most effective microorganisms to produce metal and

2
B. Pramanik et al. Plant Physiology and Biochemistry 201 (2023) 107831

metal-sulfide-containing NPs are fungi, explicitly Verticillium sp., On the et al., 2016). A number of metallic oxide nanoparticles (MgO, Al2O3,
other hand, green synthesis or green technology is easy to use, afford­ MnO, SiO2, TiO2, ZnO, and CuO) have been demonstrated to effectively
able, and environmentally beneficial. It has recently attracted a lot of control soil-borne diseases of various crops. Recent research has exam­
attention (Table 1). Several of the green nanotechnology-produced ined the effectiveness of nano-Ag against the phytopathogen Colleto­
nanoparticles have been effectively applied in a variety of fields. In trichum gloeosporioides (Aguilar-Méndez et al., 2016). On the other hand,
general, green nanotechnology may be described as the use of different AgNPs are the predominantly used NMs as antibacterial agents in food
biotechnological methods for biological sources, such as bacteria, fungi, packaging, followed by ZnO NPs. Notably, zinc nanocrystal exhibits
or plants, in order to synthesize nanomaterials (or nanoparticles). The antibacterial and antifungal activity (Duncan, 2011).
resulting nanoparticles are safe for the environment and devoid of Recently, increased crop adaptability necessitates a multifaceted
hazardous substances. Various bacteria, fungi, actinomycetes as well as strategies like hormone modulation, plant enzymatic system activation,
many plants are used to synthesize the NPs as an ecologically beneficial stress gene expression, prevention of water shortage stress, and control
source. A variety of NPs can be generated through green synthesis, of heavy metal translocation (Rajput et al., 2021; Wu and Li, 2021).
which can then be used in the further steps of applications. Many vari­ Improvements in NMs can increase crop output in the current hostile
ables, including pH, temperature, and incubation time, have an impact environment (Myint et al., 2021). In addition to preventing membrane
on the creation of nanoparticles utilizing environmentally friendly damage caused by cold stress, TiO2 NPs treatment significantly reduced
processes; these variables must be taken into account to achieve the best oxidative stress in chickpea (Cicer arietinum) (Mohammadi et al., 2014).
outcome. NPs stimulated crop development in response to high temperature stress
The recent development in nanomaterials-originated slow- or by modifying a number of physiological, biochemical, and molecular
controlled-releasing fertilizers, insecticides, and herbicides ensures processes. CeO and micronutrients NPs were employed to promote
ecological acceptance of those agricultural operations (Panpatte et al., drought-stressed soybean crop growth and development (Karami and
2016; Tripathi et al., 2022b). It aims to release a regulated quantity of Sepehri, 2017). SiNPs outperform traditional fertilizers by depleting
necessary and sufficient agrochemicals over time in order to attain full heavy metal-accumulation in plants (Wang et al., 2016). Also, ZnO NPs
biological competency while minimizing loss along with adverse out­ reportedly inhibited the acquisition of Cd and As in wheat (Triticum
comes (Shojaei et al., 2019). Gradually, the applications of nanotech­ aestivum) and rice (Oryza sativa) plants, respectively.
nology have advanced from laboratory-based experimentation. NPs are
good in delivering pesticides because of their large surface area, ease of 3. Modes of action: biochemical and molecular interventions
adhesion, and fast mass transferring ability (Ghormade et al., 2011;
Prakash et al., 2022). Several processes, e.g. encapsulation in the The effect of the NPs mainly depends on various parameters i.e. the
nano-matrix of active compounds, capsulation, assimilation, weak bond dose, shape, size and composition of the NPs, type of organism of in­
or surface ionic attachments, and encapsulation in the nano-matrix of terest along with duration of exposure. Amongst diversified mechanisms
active ingredients elements, etc. are used to incorporate microscopic or followed by NPs, intracellular and cell surface interaction, ROS gener­
sub-microscopic particles into agrochemicals (Pandey, 2018; Kandhol ation, inhibition of the activities of various proteins like catalase,
et al., 2022b). glutathione reductase, superoxide dismutase (SOD) are reported to be
Since fertilizers are one of the key issues, creating nano-based fer­ the most common. While ROS causes mitotic and chromosomal aber­
tilizers would be a cutting-edge technological advancement in this ration by exhibiting genotoxicity, some other NPs (e.g., Cu/Ce NPs)
sector. The incorporation of NPs into crops increases the content of affect various signalling pathways from fundamental level to create cell
additional elements in addition to the absorbed ones. Nano fertilizers toxicity, which leads to apoptosis, when applied at higher dose (Dimkpa
increase stress tolerance and nutrient utilization efficiency (NUE) by at et al., 2013). On the contrary, mostly moderate or limited dose appli­
least three-folds. These nano fertilizers contain nutrients and growth cation develop antimicrobial, antifungal, antibacterial effect and thus,
stimulants enclosed in nano scale polymers, allowing delayed and tar­ helps in overall plant growth (Wang et al., 2017). Despite the DNA
geted release in order to enhance soil health, control rhizosphere bac­ damage in both nucleus and cytoplasm caused by the over-production of
teria, and finally boost overall crop growth and yield. Moreover, in ROS, these NPs, at a particular dose, can also be efficiently used to limit
comparison to chemical fertilizers, nano fertilizers are less expensive its production, which eventually enhances the generation of antioxi­
and require minimal application, which ultimately reduces pollution. dants. These anti-oxidants helps the plants to adapt effectively in stress
NMs-based agrochemical formulations or nano-sized preparations conditions apart from providing them optimum growth and develop­
can be used to carry biological or synthetic herbicides efficiently for ment (Wei and Wang, 2013). while some NPs camouflage the effect of
effective elimination of weeds along with enriched bioavailability antioxidant and helps in scavenging ROS.
(Shaner, 2014). Nano herbicides provide a greater specific surface area, MgO NPs produce H2O2 on suspension, activate the oxidative stress
which increases the affinity for the target. the nanoscale encapsulation defense genes, which generates ROS like superoxide anion (O2− ) (Huang
also improves spraying efficiency by minimizing splash losses and spray et al., 2005). These easily disrupt the cell membrane of gram-negative
drift. Nano herbicides, blended with soil particles, can harm weeds or bacteria due to its unique crystalline shape with alkaline nature due to
weed seeds. Herbicides like triazine and atrazine could be capsuled for the presence of lipid peroxidation and lack of peptidoglycan layer (Tang
even distribution to crops (Pereira et al., 2014). The development of and Lv, 2014). Moreover, MgO NPs elevate O2− concentration as well as
specialized compounds for NPS-encapsulated herbicides intends to create a strong interaction to the negatively charged cell wall because of
permeate the weed roots in order to stop glycolysis, which eventually their greater surface area (Stoimenov et al., 2002).
cause death of them (Hess, 2018). ZnO NPs possess specific physicochemical properties viz. porosity,
Several particles are used in the nano-formulation of insecticides as particle size, shape etc., due to which their toxicity vary in different
active components, and additional designed nano-structures have ad­ bacteria containing media. Apart from ROS or Zn2+ ions generation,
vantageous insecticidal properties (Haq and Ijaz, 2019). The regulated these also interact with proteins (Dwivedi et al., 2014) in order to
release and persistence of active components inside plants or in rhizo­ destroy the cellular integrity (Kasemets et al. 2009). The surface defect
sphere became possible by the nano-encapsulation and and charge of ZnO NPs has a prime role in the antimicrobial activity as it
nano-formulation of insecticides, even without compromising their ef­ has large potential reactive surfaces. Gram-positive bacteria is more
ficacy. The limitation of poor water-dissolvability can also be overcome susceptible to ZnO treatment than gram-negative bacteria due to the
through these. Enhanced heat endurance, crystallinity, dissolution, difference in cell wall structure, metabolism, and physiology. Notably,
penetration, rigidity, and renewability are displayed in both ZnO NPs at higher concentration destroy gram-negative bacteria too (Hu
nano-encapsulation and nano-formulation of insecticides too (Dwivedi et al., 2012). On the other hand, the fungicidal activity of these NPs

3
 B. Pramanik et al.
Table 1
Impact of singular nanoparticles on biotic stress management in various plant species.
NPs used Conc. (ppm Size Shape Method of Additives Plant Method of Biotic stress Response Mechanism References
or mg/ml* (nm) synthesis application
or mM#)

Ag 2.5 – – High voltage Not used Brassica oleracea In vitro water Fusarium culmorum Resistance against Membrane alteration Kasprowicz et al.,
discharge var. capitata f. suspension fungal spore and disruption of (2010)
method rubra formation transport through cell
membrane
100 4–8 Colloidal – Not used Piper nigrum In vitro Colletotrichum Inhibition of the Damage in mycelial Lamsal et al.,
incubation gloeosporioides growth of the fungus germination caused by (2011a)
in vitro AgNPs
50 – – – Resistance against
anthracnose in vivo
before disease
outbreak
100 – – – Not used Cucumis sativus, In vitro Golovinomyces Prevention of Detrimental effect on Lamsal et al.,
Cucurbita sp. incubation cichoracearum or powdery mildew mycelial growth and (2011b)
– Sphaerotheca fusca disease outbreak conidial germination
caused by NPs
15 mg/μμl – – Green synthesis Not used – – Alternaria alternata, Resistance against Destruction of fungal Krishnaraj et al.,
Sclerotinia fungal pathogens spore through (2012)
sclerotiorum, disruption of
Macrophomina membrane integrity
phaseolina,
Rhizoctonia solani,
Botrytis cinerea,
Curvularia lunata
150 0.25 Chemical Not used Cynodon Soil Meloidogyne spp. Reduction in Change in membrane Cromwell et al.,
4

–
synthesis dactylon treatment nematode permeability due to (2014)
production and gall nematicidal effect of
formation the NPs
2 – – Extracellular Not used Triticum aestivum Spray Bipolaris sorokiniana Resistance against Inhibition of conidia Mishra et al.,
synthesis method spot blotch disease formation as well as (2014)
germination,
accumulation of lignin
in vascular bundles
1# 10 Colloidal Green synthesis Not used Brassica rapa Agar-disc Gloeophyllum Resistance against Formation of Narayanan and
diffusion abietinum, wood-degrading deformed and broken Park (2014)
method G. trabeum, pathogens mycelium with
Chaetomium swollen margins
globosum, caused by the

Plant Physiology and Biochemistry 201 (2023) 107831

Phanerochaete application of AgNPs
sordida
15 20 Spherical – Not used – In vitro Alternaria alternata, Inhibition of fungal NPs cause damage to Ouda (2014)
incubation Botrytis cinerea growth the fungal hyphae,
conidia, and cell wall
components
10 – – Green synthesis Not used Nicotiana Spray Phytophthora Resistance against Inhibition of mycelial Ali et al., (2015)
benthamiana method parasitica, P. capsici plant pathogens growth, germination
and production of
zoospore, germ tube
elongation
5 2–5 Spherical Green synthesis Not used Solanum Foliar spray Alternaria solani Resistance against Higher antimicrobial Kumari et al.,
lycopersicum fungal spore activity of AgNPs due (2017)
formation to the surface coating
of secondary
(continued on next page)
 B. Pramanik et al.
Table 1 (continued )
NPs used Conc. (ppm Size Shape Method of Additives Plant Method of Biotic stress Response Mechanism References
or mg/ml* (nm) synthesis application
or mM#)

metabolites of
Trichoderma viridae,
and their shape and
size
75 – – Green synthesis Not used Oryza sativa Soil Aspergillus flavus Enhanced growth Reduced aflatoxin Ejaz et al., (2018)
application production
100 – – – Not used Cicer arietinum In vitro Fusarium oxysporum Resistance against Antifungal activity of Kaur et al., (2018)
incubation f. sp. ciceri wilt disease the NP through
mycelial growth
inhibition
90 – Colloidal Green synthesis Not used NM In vitro Fusarium Resistance against Higher antimicrobial Gautam et al.,
incubation graminearum, phytopathogenic activities of specific (2020)
F. avenaceum, microbes NPs due to smaller size
F. culmorum, and target-specificity
Pseudomonas syringe
70 – – Erwinia sp.
110 – – Bacillus megaterium
5 2–5 Spherical Bioengineering Not used Arabidopsis Spray Alternaria brassicicola Systemic resistance Increased PAL activity Kumari et al.,
method thaliana method against fungal spore resulting in (2020)
formation production of
antimicrobial
phenolic compounds
50 <100 – Chemical Calothrix Solanum Foliar spray Alternaria alternata Reduction in disease Elicitation of defence Mahawar et al.,
synthesis elenkinii lycopersicum severity and antioxidant (2020)
machineries,
5

increased efficiency of
hydrolytic enzymes by
the combined
antimicrobial effect of
NPs and C. elenkinii
100 – – Chemical Not used Prunus persica In vivo Pseudomonas syringae Resistance against Antibacterial Shahryari et al.,
synthesis treatment pv. syringae bacterial canker activities of NPs due to (2020)
their smaller size
50 – – Green synthesis Not used Trachyspermum Soil Meloidogyne incognita Reduction in number Nematicidal activity Danish et al.,
ammi. application of galls, egg masses, of NPs produced from (2021)
and root knot Senna siamea
50 – – Green synthesis Not used Oryza sativa Foliar spray Aspergillus flavus Control over fungal Decrease in Sultana et al.,
pathogen production level of (2021)

Plant Physiology and Biochemistry 201 (2023) 107831

proliferation enzymatic and non-
enzymatic
compounds,
osmolytes at stress
condition
Ag (ds DNA-directed) 100 18 – Chemical Graphene Solanum Spray Xanthomonas Decreased severity of Enhanced Ocsoy et al.,
synthesis oxide lycopersicum method perforans bacterial spot disease antibacterial property (2013)
of the composite
Au NM – – Green synthesis Not used – Well Puccinia graminis Resistance against Elevated plant Jayaseelan et al.,
diffusion tritici stem rust of wheat immune system and (2013)
method antimicrobial activity
due to the smaller size
Ca3(PO4)2 150 <50 – – Not used Solanum Soaking Rhizoctonia solani Prevention of black Elevated peroxidase El-Shewy, 2019
tuberosum method → scurf disease and catalase activity
soil in potato leaves
(continued on next page)
 B. Pramanik et al.
Table 1 (continued )
NPs used Conc. (ppm Size Shape Method of Additives Plant Method of Biotic stress Response Mechanism References
or mg/ml* (nm) synthesis application
or mM#)

drenching incidence and

after 45 days severity
CeO2 250 8±1 – – Not used Solanum Foliar spray Fusarium oxysporum Reduction in disease Enhancement of Adisha et al.,
lycopersicum f. sp. lycopersici spread antioxidant and (2018)
photosynthetic
efficiency
Ch 1 – – Chemical Not used Eleusine coracana Seed soaking Pyricularia grisea Induced defence Induced production of Sathiyabama and
synthesis (Cke.) Sacc response against ROS and increased Manikandan
blast by reducing level of peroxidase (2016)
sporulation activity, leading to
efficiency delayed symptoms
5* – – – Not used Solanum Seed soaking Fusarium andiyazi Increasing plant Inhibition of radial Chun and
lycopersicum defence against wilt mycelial growth Chandrasekaran
(2019)
NM – – – Not used Triticum aestivum Spray Puccinia triticina Reduced leaf rust Accelerated Elsharkawy et al.,
method transcription level of (2022)
PR1-PR5 and PR-10
genes
CNM (Single walled) 500 – – – Not used NM Suspension Fusarium Induction of Deposition of CNMs Wang et al.,
method graminearum, F. poae antifungal activity on spore surface, (2014)
inhibition of water
uptake, induced
plasmolysis
Cu 20 μg/disc – – Chemical Not used NM Disc diffusion Phoma distructiva, Inhibition of fungal Biocidal effect of Cu Kanhed et al.,
synthesis method Alternaria alternata, growth complex in CuNPs (2014)
6

– – 20 μg/disc Curvularia lunata,

Bavistin Fusarium oxysporum
50 – – – 184 ppm Solanum Foliar spray Clavibacter Stimulation of Increased activity of Cumplido-Nájera
K2SiO3 lycopersicum michiganensis tolerance against the SOD, PAL, GPX, and et al., (2019)
pathogen APX, as well as
reduced glutathione
and phenol content in
leaves
Cu3(PO4)2⋅3H2O NS 10 30 – Chemical Not used Citrullus lanatus Dip method Fusarium oxysporum Repression of fungal Alleviation of stress Borgatta et al.,
synthesis f. sp. niveum infection and spread under diseased (2018)
condition due to the
lack of silicon change
in the cell

Plant Physiology and Biochemistry 201 (2023) 107831

CuO 27.78 g/hl 11–55 – Chemical Not used Solanum Foliar spray Phytophthora Resistance against Antifungal activity of Giannousi et al.,
synthesis lycopersicum infestans late blight and the respective NPs due (2013)
reduced infection on to the bio reactive
leaves nature of Cu
10 30 ± 10 – – Not used Transgenic Dispersion Helicoverpa armigera Insect resistance in Enhancement of Le Van et al.,
Gossypium sp. method Bt cotton expression of (2016)
exogenous gene
encoding Bt toxin
proteins in roots and
leaves
232.75 – – Direct Not used Gossypium sp. Leaf-dip Spodoptera littoralis Resistance against Penetration through Ayoub et al.,
precipitation bioassay cotton leafworm cell membrane and (2018)
method damage of the vital
enzyme
(continued on next page)
 B. Pramanik et al.
Table 1 (continued )
NPs used Conc. (ppm Size Shape Method of Additives Plant Method of Biotic stress Response Mechanism References
or mg/ml* (nm) synthesis application
or mM#)

200 – – Green synthesis Not used Vigna Aqueous Meloidogyne incognita Reduction in Nematicidal effect of Tauseef et al.,
unguiculata suspension nematode CuO due to alteration (2021)
method production, egg of mitochondrial
masses, and number activity, enhancement
and size of root gall of ROS production and
formation triggered apoptotic
pathways
CuS 50 – – Chemical Not used Oryza sativa Seed Gibberella fujikuroi Suppression of Reduction in Shang et al.,
synthesis treatment infection in rice microbial growth due (2020)
to the interaction
between cell and NP/
Cu2+ ions
K2SiO3 NM – – – Not used Coffea arabica Foliar spray Hemileia vastatrix Resistance against Reduced uredia Carré-Missio et al.
coffee leaf rust production on leaves 2014
MgO NM 50 ± 10 – Biosafe method Not used NM Suspension Alternaria alternate, Inhibition of fungal Antimycotic activity Wani and Shah
method Rhizopus stolonifer, growth of NPs due to release (2012)
Fusarium oxysporum, of extracellular
Mucor plumbeus enzymes and
metabolites
0.1% – – – Not used Solanum Soil Ralstonia Induced systemic Production of PR Imada et al.,
lycopersicum application solanacearum resistance to proteins, (2016)
bacterial wilt disease accumulation of β-1,3-
glucanase and tyloses
in the xylem and
apoplast of pith tissues
7

leading to
development of
resistance
O-Ch 2* 297 – – Not used NM Dispersion Nigrospora sphaerica, Resistance against Inhibition of mycelial Xing et al., (2016)
method Botryosphaeria these plant growth due to the
dothidea, Nigrospora pathogenic fungi antifungal activity of
oryzae, Alternaria the NPs
tenuissima
Si NM – – – Not used Rosa rubiginosa In vitro Podosphaera pannosa Reduced powdery Increased Shetty et al.,
incubation mildew severity concentration of (2011)
antimicrobial
phenolic acid and
flavonoids

Plant Physiology and Biochemistry 201 (2023) 107831

3# 249 – Chemical Not used Vicia faba Foliar Botrytis fabae Decreasing incidence Improvement in the Hasan et al.,
synthesis application and severity index of activities of protective (2020)
chocolate spot enzymes, peroxidase,
disease polyphenol oxidase,
proline etc. due to the
treatment of SiNPs
100 – – Green synthesis 0.5 RD Femax Solanum Soil Meloidogyne incognita Reduced nematode Malfunction in El-Ashry et al.,
40% EC + melongena application production, egg cellular mechanism (2022)
Fosthiazate masses, and root gall and reduction in egg
10% WG + formation hatching efficiency
krenkel 75%
EC
Si (Mesoporous) 400 – – One-pot direct Not used Solanum Spray Alternaria solani Resistance against Higher antifungal Derbalah et al.,
template lycopersicum method early blight and efficacy due to high (2018)
approach porosity, smaller
(continued on next page)
 B. Pramanik et al.
Table 1 (continued )
NPs used Conc. (ppm Size Shape Method of Additives Plant Method of Biotic stress Response Mechanism References
or mg/ml* (nm) synthesis application
or mM#)

proper growth of the particle size, and

plant proper shape
500 – – – Not used Citrullus lanatus Foliar spray Fusarium oxysporum Suppression of Strengthening of Buchman et al.,
f. sp. niveum Fusarium wilt natural defence (2019)
disease system of the plant
due to the action of
NPs
SiO2 15 kg/ha 20–40 – – Not used Zea mays Soil Fusarium oxysporum, Resistance against Enhancement of both Suriyaprabha
application Aspergillus niger fungal and bacterial uptake efficiency and et al., (2014)
infection accumulation of silica,
leading to leaf
erectness and defence
response against
pathogens
100 5–15 Spherical – Not used Daucus carota Foliar spray Pectobacterium Resistance against Accumulation and Siddiqui et al.,
carotovorum these microbes along deposition of silica (2020)
Xanthomonas with increasing plant beneath the cell
campestris pv. carotae growth cuticle causes
Rhizoctonia solani formation of a Si-
Fusarium solani double layer, that
Alternaria dauci results in prevention
of pathogen entry.
50 5 – Green synthesis Not used Oryza sativa Foliar spray Fusarium fujikuroi Reduction in Improved silica Elamawi et al.,
sporulation of the content, electrolyte (2020)
fungus, decreasing leakage, and
8

severity index of peroxidase activity

bakanae disease due to the application
of NPs
100 54 ± 7 – – Not used Arabidopsis Suspension Pseudomonas syringae Local and systemic Activation of SA- El-Shetehy et al.,
thaliana method disease resistance dependent defence (2021)
pathway
1500 – Spherical Reverse 1 ml/l non- Citrullus lanatus Foliar spray Fusarium oxysporum Suppression of Effective delivery of Kang et al., (2021)
microemulsion ionic f. sp. niveum Fusarium wilt silicic acid leading to
method surfectant proper growth while
infected with the
fungus
30 – – – Not used Phalaenopsis Aqueous Dickeya dadantii Induced resistance Increased production Nisaq et al., (2021)
pulcherrima spray against soft rot of resistance-related

Plant Physiology and Biochemistry 201 (2023) 107831

disease proteins and
formation of hard
layer of leaves
1000 – – Stober process Not used Oryza sativa Foliar spray Magnaporthe oryzae Increased resistance Elevated upregulation Du et al., (2022)
against blast of SA responsive genes
TiO2 75# – – – Not used Pelargonium sp. Foliar spray Xanthomonas Controlling Higher rate of Norman and Chen
hortorum pv. Xanthomonas oxidation reaction (2011)
pelargonii bacterial blight leading to reduction in
– – Euphorbia Xanthomonas Controlling lesions in leaves
pulcherrima axonopodis pv. Xanthomonas leaf
poinsettiicola spot
0.5 g/l – – – Not used Beta vulgaris Foliar spray Cercospora beticola Decreasing severity Distortion and Hamza et al.,
of leaf spot disease damage of sporangial (2016)
membrane leading to
death
(continued on next page)
 B. Pramanik et al.
Table 1 (continued )
NPs used Conc. (ppm Size Shape Method of Additives Plant Method of Biotic stress Response Mechanism References
or mg/ml* (nm) synthesis application
or mM#)

40 10–100 – Chemical Not used Triticum aestivum Soil Bipolaris sorokiniana Resistance against Antimicrobial activity Satti et al., 2021
synthesis application spot blotch disease due to smaller size and
other morpho-
physical characters
1
ZnO 10− M 2–28 Spherical Chemical Not used – – Pseudomonas sp. Inhibition of Rupturing of cell Sharma et al.,
– – synthesis Fusarium sp. microbial growth membrane leading to (2010)
the death of the
microbes
>3 mmol/l 70 ± 15 Spherical NM Not used – In vitro Botrytis cinerea Inhibition of fungal Deformation in the He et al., (2011)
incubation growth structure of fungal
hyphae
6 mmol/l – – Penicillium expansum Production of
distorted and
damaged conidia
4# – – Green synthesis Not used – Agar well Aspergillus flavus Resistance against Release of ROS Gunalan et al.,
diffusion fungal growth causing damage to the (2012)
method membrane and
ultimately leading to
death of the fungus
25 – – Green synthesis Not used – Well Aspergillus flavus Inhibition of fungal Antifungal activity Jayaseelan et al.,
diffusion growth due to smaller particle (2012)
technique size
500 – – Microwave Not used – Slide Aspergillus niger Inhibition of fungal Shrivelling of mycelial Patra et al., (2012)
– – assisted route bioassay Fusarium oxysporum growth wall and release of
Zn2+ with fungitoxic
9

effect
NM 30 ± 10 – Biosafe method Not used – Suspension Alternaria alternate, Inhibition of fungal Antimycotic activity Wani and Shah
method Rhizopus stolonifer, growth of NPs due to release (2012)
Fusarium oxysporum, of extracellular
Mucor plumbeus enzymes and
metabolites
500 <100 – – Not used Triticum aestivum In vitro Fusarium Inhibition of fungal Increased antifungal Dimkpa et al.,
incubation graminearum growth activity due to smaller 2013
particle size, charge
and dissolution
efficacy
25 27 ± 5 – Chemical Not used – Modified Aspergillus flavus, A. Resistance against Higher antifungal Rajiv et al., 2013
synthesis Kirby Bauer niger fungal growth activity due to smaller

Plant Physiology and Biochemistry 201 (2023) 107831

disc diffusion size
method
100 20–25 – Precipitation Not used – Modified Aspergillus flavus Resistance against Production of ROS Navale et al.,
method broth fungal growth from the NPs results in (2015)
dilution antifungal activities
Method
16 12–32 – Chemical Not used – Suspension Aspergillus niger Resistance against Disruption of the Jamdagni et al.,
synthesis method this fungus structure of cell (2018)
membrane followed
by the interference
with fundamental
process of the cell
200 ≤100 – – Not used Beta vulgaris Suspension Pectobacterium Management of root Disruption in the Khan and
subsp. vulgaris method betavasculorum, knot-, root rot-, soft development of Siddiqui, 2020
Meloidogyne conidia and
(continued on next page)
B. Pramanik et al. Plant Physiology and Biochemistry 201 (2023) 107831

disturbs the reproductive system and disintegrates the fungal hyphae as

Abbreviations: Ag: Silver; APX: Ascorbate peroxidase, Ca3(PO4)2: Calcium phosphate; CeO2: Cerium dioxide; Ch: Chitosan; CNM: Carbon nanomaterial; Cu: Copper; Cu3(PO4)2⋅3H2O: Copper phosphate trihydrate; CuO:
Copper oxide; CuS: Copper sulphate; GPX: Glutathione peroxidase, hl: Hectolitre; K2SiO3: Potassium silicate; mg: Milligram; mM: Milli molar; ml: Milliliterre; -: Not mentioned; NPs: Nanoparticles; NS: Nanosheet; O-Ch:
Oleoyl chitosan; PAL: Phenylalanine ammonia lyase; ppm: Parts per million; PR: Pathogen resistance; RD: Recommended dose; ROS: Reactive oxygen species; SA: Salicylic acid; Si: Silicon; SiO2: Silica; SOD: Superoxide
well (He et al., 2011).

Sofy et al., 2021

The interaction between AgNPs and cell membrane creates their path

Kairyte et al.,
to enter into the cell cytoplasm, where it creates disturbance in the

References
electron transport chain (ETC) by interacting with the thiol group of

(2013)
proteins. Moreover, they interfere with the free movement of phosphate
and protons and generate ROS (Durán et al., 2010; Thombre et al.,
2016). Besides altering the cellular physiology, these also block the

conidiospores leading
to the death of fungal

Distortion of conidial
heads and shrivelling
Zn-mediated cellular

environmental stress
G2/M phase of cell cycle in fungus. At molecular level, several genes like

of mycelial walls
glutathione reductase and glutathione-S-transferase genes got activated,

integrity and
when exposed to AgNPs, and participates in scavenging ROS (Nair and
Mechanism

resistance
hyphae Chung, 2014).
Exposure to AuNPs can upregulate or downregulate many genes in
bacteria (Cui et al., 2012). Giri et al. (2015) reported that gold nano­
particles targeted the bacterial cell membrane and cell wall to damage
rot-mediated disease

Systemic protection

Inhibition of fungal
complex formation

the entire cell, which ultimately stopped the antibacterial affect. Also it
helped several antibiotics to enter through the peptidoglycan layer of
against ToMV

the Gram negative bacteria’s cell membrane. Notably, the effect of Au on

Response

growth

Gram positive and negative bacteria is different. AuNPs affect the

multidrug resistant ubiquitous Gram positive bacteria via quorum
sensing to reduce the effect of toxins through photodynamical or pho­
tothermal way. (Wang and Wood, 2011).
tobamovirus (ToMV)
Rhizoctonia solani

The dose of Au is also important as it is directly proportional to

Tomato mosaic

Botrytis cinerea

bacterial cell membrane permeability as well as inversely proportional

incognita, and
Biotic stress

to its viability (Adhikari et al., 2013). Different concentration may affect

different organisms via various mechanisms. 4,6-diamino-2-pyrimidine­
thiol (DAPT), although having no antibacterial effect, increases the
permeability of cell membrane in E. coli and Pseudomonas aeruginosa,
when capped with gold nanoparticle resulting in the leakage of nucleic
Foliar spray
application

incubation
Method of

acid from cell. It will also chelate with magnesium (Mg2+) ion and
In vitro

change its concentration either to disturb the lipopolysaccharide layer,

which regulates the outer membrane permeability of cell (Zhao et al.,
2010), or to lyse the cell by forming aggregation on cell membrane
(Hayden et al., 2012). These occurs due to interference in fundamental
lycopersicum

biochemical processes including inhibition of the production of ATP or

Solanum

lack of binding of tRNA to the subunits of ribosomes (Cui et al., 2012).

Plant

Again, increased Ca2+ ion concentrations continuously lead to the

–

fragmentation of DNA (Nagata, 2000; Lee and Lee, 2018). AuNPs’

application increases the ROS and SOS response by decreasing the H2O2
scavenging ability and induces bacterial apoptosis by over expression of
Additives

Not used

Not used

the caspase-like protein (Nagata, 2000) and thus, damages the macro­
molecules causing oxidative stress (Matés et al., 2012). The imbalance
between oxidative, and antioxidant system ultimately reduces the gen­
eration of glutathione (GSH) and induces ROS independent apoptosis
(Lee and Lee, 2018). It has wide bactericidal activity due to its multi­
Method of

valent effect as Au can easily modify itself irrespective of the environ­

Chemical
synthesis

synthesis

ment (Chowdhury et al., 2017; Kalita et al., 2018). In contrast the

antifungal effect of AuNPs, which is ROS independent, they also own
–

dismutase, TiO2: Titanium oxide; Zn: Zinc; ZnO: Zinc oxide.

both cytoplasmic impact, where they affect mitochondrial membrane

potential (MMP) by changing the concentration of Ca2+ ion or
Shape

H+-ATPase-mediated proton pump to disturb the proton gradient ho­

–

meostasis inside mitochondrial cell membrane, and nucleoplasmic ef­

fect, where they interact with P and S inside nucleus to affect protein
function for degrading DNA, and nutrient transportation, which finally
(nm)
Size

400

leads to the death of fungi (Yu et al., 2016; Seong and Lee, 2018).
–

Further to indicate, different antifungal and antibacterial substances like

Conc. (ppm
or mg/ml*

lysozyme, cinnamic acid are coated with Au to enhance its activity

or mM#)

(Gupta et al., 2017; Ahmady et al., 2019). AuNPs also neutralize the
0.5#
100

charge and reduce the number of plasmid when the weightage ratio
between AuNPs and plasmid becomes 15:1. In this circumstance, AuNPs
Table 1 (continued )

ultimately prevent its movement within the cell and thus, causes
(Photoactivated)

apoptosis (Zhao et al., 2010). The extreme light to heat conversion ca­
pacity of Au nanoparticles destroys bacterial cell in 10 min minutes by
NPs used

converting near infrared (IR) light into local heat through protein
denaturation, cell fluid evaporation, and cell wall rupture (Venugopalan
ZnO

et al., 2002; Von-Maltzahn et al., 2009; Huang and El-Sayed, 2010;

10
B. Pramanik et al. Plant Physiology and Biochemistry 201 (2023) 107831

Gharatape et al., 2016). Near IR light reduces the number of bacteria 4. Stress management and nanoparticles
while local heat helps in decomposition of the same. Although AuNPs
have been proven nontoxic to mammal tissues and cells, and in vitro Throughout the life span, plants undergo various kinds of environ­
experiments, further researches should be executed on the bacteriostatic mental conditions due to fluctuating climatic scenarios, topological
mechanisms and interactions between AuNPs and internal environment. differences, varying soil health and many more. The morphological
Chitosan nanoparticles (ChNPs) act either on the cell membrane or features, growth, and production are adversely affected by these
on the cell wall of the bacteria through an electrostatic interaction be­ external factors. These stimuli are cumulatively known as stress (Bhatt
tween negatively charged cell membrane and positively charged amino et al., 2020). Stress can be caused by numerous biotic as well as abiotic
groups of glucosamine. Consequently, it leads to the generation of os­ agents. The biotic agents include pathogen, insects, nematodes etc.,
motic imbalance and cell death because of increased membrane whereas the latter mentions the non-living entities such as drought,
permeability (Raafat et al., 2008). It causes accumulation of Ch inside salinity, cold, heat etc. All of these stimuli together impart a negative
the cell through influx, which binds with DNA to inhibit the replication impact by interfering the homeostasis maintaining pathways in plants,
and ultimately degrade it. Also, it modifies the ETC of bacterial cell which eventually leads to crop loss. Therefore, as a novel approach, the
(Birsoy et al., 2015; Ivask et al., 2013). ChNPs are able to chelate with application of nanoparticles directs an innovative way to check the
various metal ions in acidic condition such as Fe2+, Mg2+, Ni2+, Co2+, aforesaid problems along with boosting overall growth and develop­
Cu2+, and Zn2+ in order to interrupt the cell membrane integrity (Joshi ment of the plant, and ultimately the productivity.
et al., 2009). There are different types of ChNPs like ChNP3, ChNP4,
which act on the cell membrane with different degrees in specific bac­
terial cells. 4.1. Effect of nanoparticles on biotic stress management
On the other hand, CuNPs are popular antimicrobial agent due to its
greater surface area to volume ratio. It is effective against both Gram Biotic stress on a plant specifically caused by pathogens, insects and
positive and negative bacteria (Theron et al., 2008; Schrand et al., nematodes. Upon attack of any of these agents, plants generate an
2010), besides having anti-cancerous properties (Jose et al., 2011). The immunological response by developing various complicated physiolog­
mode of action of Cu nanoparticles is similar as AgNPs (Ruparelia et al., ical and biochemical processes in order to combat (Almutairi, 2019). In
2008), which penetrate the cell wall, interact with cell membrane due to the same context, nanoparticles play a very crucial role due to their
their affinity towards –NH2 and –COOH groups, and affect DNA double antimicrobial activity, which is widely spread among the plants (Fig. 1,
helical structure. They also form chelate structure through various Tables 1 and 2). Among all, the most commonly used nanoparticle was
biochemical processes by interacting with –SH group (Schrand et al., silver nanoparticles, followed by zinc oxide nanoparticles, silica nano­
2010) in order to degrade those proteins behind cellular toxicity. particles, and copper based one.
Furthermore, due to the small size of Cu, they directly pass through cell
or mitochondrial membrane via membrane transporters or endocytosis, 4.1.1. Silver nanoparticles (AgNPs)
respectively, and produces Cu2+ ions leading to the ROS production. AgNPs were predominantly used for controlling fungal infection in
Although TiNPs have not been explored too much, these NPs cause various families of crops such as Graminae, Crucifereae, Solanaceae and
chromosomal aberrations by interfering in the process of microtubules many more (Table 1). This nanoparticle was mainly known to prevent
formation, which control the segregation of the chromosomes. the formation and growth of mycelia and/or conidia of fungi. Some­
times, development of deformed and broken mycelium (Narayanan and
Park, 2014), fractured cell wall (Ouda, 2014) due to the use of AgNPs
have also been reported. Ejaz et al. (2018) published a report indicating
enhanced growth in Aspergillus flavus infected AgNP-treated rice plant.

Fig. 1. Mechanism and effect of different nanoparticles on biotic stress for plant growth and health improvemnt.

11
 B. Pramanik et al. Plant Physiology and Biochemistry 201 (2023) 107831

In that particular study, they were able to limit the production of afla­

Abbreviations: Ag: Silver; CS: Core-shell; Cu; Copper; FQ: Fixed quat; IrCh: Irradiated chitosan; MV: Mixed variance; NPs: Nanoparticles; OCh: Oleoyl chitosan; PR: Pathogen resistance; ROS: Reactive oxygen species; Se:
Quiterio-Gutiérrez et al.,
toxin from the bacteria with the treatment of AgNPs, which led to the

Nguyen et al., (2017)

Moussa et al., (2013)
biomass enhanced than that of any infected rice plant. Although most of

Manikandan (2018)
Paret et al., (2013)
Chu et al., (2012)

Fan et al., (2021)

Sathiyabama and
Cui et al., (2016)
the studies were conducted employing AgNPs alone, some reports have
been found implementing graphene oxide (GO) and Calothrix elenkinii as
References

additives to increase the efficiency (Ocsoy et al., 2013; Mahawar et al.,

(2019)
2020). The former group of researchers stated that 16 ppm of
Ag@dsDNA@GO composite was capable enough to show antimicrobial
properties in culture, though 100 ppm of the composite showed the same
Production of PR protein for induced defence

Inhibition of cell respiration, cell permeability

increasing efficiency after radiation treatment

mechanism against fungal attack by the plant

change, and damage in cell wall followed by
NPs are able to penetrate the cell membrane

Activation of defence enzymes for improved

in tomato (Solanum lycopersicum) plants. Apart from microbial in­

eliciting the production of defence related

Increasing ROS production leading to cell

Penetration of bacterial cell wall and cell

fections, this nanoparticle can be efficiently used in decreasing number
Combined activity of both NPs through
in order to rupture the cell due to its

of gall and egg formation, and nematode production (Cromwell et al.,

Generation of a cascade of defence

proteins, enzymes, and hormones
2014; Danish et al., 2021). These studies proved the nematicidal activ­
ities of this particular nanoparticle against Meloidogyne sp.
response against pathogen

4.1.2. Zinc oxide nanoparticles (ZnO NPs)

After AgNPs, ZnO NPs was renowned as the most utilized one among
immune response

nanoparticles. It was used for crop improvement for last several years as
Mechanism

membrane
zinc being one of the most important micronutrients for plant develop­
cytoplasm

damage

ment (Amjad and Serajuddin, 2021). Although this nanoparticle has

been tested in vitro for inhibiting microbial growth, its application was
also explored in several crops like wheat, beetroot (Beta vulgaris), tomato
anthracnose and enhanced
Reduction in bacterial spot

etc. (Table 1). Physically, these nanoparticles are smaller in size, and
Development of defence

Reduction in severity of
formation of pathogen

they generate reactive oxygen species (ROS) after pathogen attack,

growth and gray mold
Prevention of fungal

bacterial leaf blight

because of which they possess higher antimicrobial activities. Apart

Inhibition of spore
Resistance against

Resistance against

Resistance against

from interfering in conidia production process, this nanoparticle also

bacterial spot
plant growth

early blight

disrupted cell membrane formation, blocked cellular fundamental pro­

Response

response

cesses of the pathogens to prevent their proliferation inside host plant.

lesions
decay

This can be considered as the one among all the nanoparticles

mentioned in this review, that can provide systemic protection against
tomato mosaic tobamovirus (ToMV), as reported by Sofy et al. (2021).
syringae pv. tomato

Colletotrichum sp.
oryzae pv. oryzae

Pyricularia grisea

Moreover, Khan and Siddiqui (2020) used ZnO NPs to manage a disease
Alternaria solani
Botrytis cinerea

Xanthomonas

Xanthomonas

Xanthomonas
Impact of composite nanoparticles on biotic stress management in various plant species (arranged periodically).

Pseudomonas

complex of root knot, root rot, and soft rot in beetroot caused by Pec­
euvesicatoria
Biotic stress

perforans

tobacterium betavasculorum, Meloidogyne incognita, and Rhizoctonia sol­

ani, respectively.
Pers.

4.1.3. Silica nanoparticles (SiO2 NPs)

Agar macro-dilution

The smaller particles of SiO2 were applied in soil while cultivating

Syringe infiltration
foliar spray + seed
Foliar spray and

maize (Zea mays) for checking the growth of Fusarium oxysporum, and
Spray method

Spray method

Spray method
Disc diffusion

Foliar spray

Aspergillus niger (Suriyaprabha et al., 2014). In that particular study, this

application
Method of

technique
treatment

was applied in soil at the rate of 15 kg/ha. Apart from cereals, SiO2 NPs
method

method

were also employed in resisting disease severity in carrot, Arabidopsis,

watermelon, orchids etc. This nanoparticle mainly targeted enzymatic
pathways to develop resistance against biotic stress (Table 1). It
Capsicum annuum

Eleusine coracana

specially activated salicylic acid (SA) responsive genes to generate SA

lycopersicum

lycopersicum
Piper nigrum
Oryza sativa

dependent defence pathways (El-Shetehy et al., 2021; Du et al., 2022).

(Fragaria ×
Strawberry
Arabidopsis

ananassa)
Solanum

Solanum
thaliana

Kang et al. (2021) also reported of efficient delivery of silicic acid to

Plant

promote sufficient growth of watermelon plant even in Fusarium oxy­

sporum f. sp. niveum-infected condition. Despite enzymatic process, silica
Selenium; Si: Silicon; SiO2: Silica; TiO2: Titanium dioxide.

particles got deposited on the cuticle layer of cell in order to prevent

Spherical

pathogen entry (Siddiqui et al., 2020).

Shape

–
–
–

4.1.4. Copper (Cu) based nanoparticles

Copper based nanoparticles were synthesized in many formats.
<100
(nm)
Size

<8

These include simple copper nanoparticles (CuNPs), complex copper

–

–
–
–

nanoparticles like copper oxide nanoparticles (CuO NPs), copper sul­

Conc. (ppm or

phate nanoparticles (CuS NPs) etc. Borgatta et al. (2018) used a, more
mg/ml* or

complex copper phosphate trihydrate nanosheets (Cu3(PO4)2⋅3H2O NS)

500–800

50 + 50

10 + 50

to alleviate the stress condition under Fusarium oxysporum f. sp. niveum

mM#)

1000
0.1*
125

3.2

attack in watermelon. This nanosheet controlled the concentration of

10

silicon inside cell to maintain this state. CuNPs were implemented along
with a high dose (184 ppm) of potassium silicate to develop resistance
(γ-irradiated)

TiO2–Ag (Light

against Clavibacter michiganensis in tomato plant. The key mechanism

activated)

OCh-SiO2
NPs used

behind this was to elevate the activities of numerous enzymes like su­
SiO2–Ag
Ag-IrCh

MV-Cu
Table 2

FQ-Cu
Cu-Ch

Se–Cu

CS-Cu
Ag–Si

peroxide dismutase, phenylalanine ammonia lyase, glutathione peroxi­

dase, ascorbate peroxidase etc. along with decreased glutathione and

12
B. Pramanik et al. Plant Physiology and Biochemistry 201 (2023) 107831

phenol content in leaves (Cumplido-Nájera et al., 2019). CuO NPs were more singular nanoparticles (Table 2). For successful integration of two
deployed to withstand insect infestation and nematode attack other than nanoparticles, sometimes irradiated nanoparticles have also been used
pathogen infection. A little concentration (100 ppm) of CuO NPs (Chu et al., 2012; Moussa et al., 2013). The latter group of researchers
enhanced the activity of exogenous proteins against lepidopterans in Bt reported that, radiation treatment uplifted the efficiency of
cotton (Le Van et al., 2016), whereas higher concentration (232.75 ppm) silver-chitosan nanoparticles complex due to which these nanoparticles
caused damage to the vital enzymes in cotton leafworm in order to became able to penetrate the cell more effectively in order to rupture it.
inhibit its infestation (Ayoub et al., 2018). According to Tauseef et al. Light-activated TiO2–Ag nanoparticles inhibited cellular physiological
(2021), 200 ppm CuO NPs possessed similar nematicidal activity against activities to prevent Xanthomonas perforans infection in tomato (Paret
M. incognita like AgNPs, as mentioned earlier. Further to mention. The et al., 2013). According to Cui et al. (2016), SiO2–Ag complex nano­
interaction between cellular constituents and Cu2+ ions in CuS NPs was particles increased the production of ROS for damaging cells of Xan­
able to suppress Gibberella fujikuroi infection in rice (Shang et al., 2020). thomonas oryzae pv. oryzae to develop resistance against bacterial leaf
blight in rice. Moreover, a number of composite nanoparticles were
4.1.5. Silicon nanoparticles (SiNPs) utilized to elicit the defence mechanism of various plants to prevent
Silicon nanoparticles were mostly used in vegetables, fruits and floral biotic stresses (Nguyen et al., 2017; Sathiyabama and Manikandan,
crops (Table 1). It actually influenced the biochemical pathways for 2018; Quiterio-Gutiérrez et al., 2019).
improving the resistance mechanism against pathogens. El-Ashry et al.
(2022) reported that 100 ppm SiNPs along with combinations of half of 4.2. Effect of nanoparticles on abiotic stress management
the recommended dose of Femax 40% EC, Fosthiazate 10% WG, and
krenkel 75% EC caused malfunction in intercellular mechanisms, and One of the most important environmental problems facing the world
thus, reduced the egg hatching efficiency of M. incognita. Other than today is abiotic stress, which has prompted academics to focus their
conventional SiNPs, the mesoporous one was also competent enough to efforts on protecting the ecological system for long-term stability. The
develop resistance against various microbes due to its porous nature, major organic stresses affecting agricultural production worldwide are
proper shape and particle size (Derbalah et al., 2018; Buchman et al., drought, salinity, heavy metals, and temperature extremity (high and
2019). low). These abiotic factors interfere the cell osmotic potential which
leads to ionic imbalance and hampers the plant metabolism. Addition­
4.1.6. Chitosan nanoparticles (ChNPs) ally, the whole ecosystem is also at danger due to the deterioration of
Chitosan nanoparticles surfaced in this field later than the others. It agricultural soils. The statistical approximation indicates that these
also possessed antifungal activity due to which radial mycelial growth pressures predominantly affect around 90% of agricultural land areas,
got checked after application. Recently, Elsharkawy et al. (2022) resulting in a 70% decrease in the production of major food crops. To
discovered its activity in the transcription level by accelerating the alleviate the adverse effects of abiotic stress and to encourage plant
transcription rate of several pathogen resistance genes in order to stress adaptation, a number of agricultural and physiological strategies
decrease the severity of leaf rust in wheat. are applied. Nanotechnology has become a fascinating platform in
recent years. The use of nanoparticles improves plant stress resistance by
4.1.7. Titanium dioxide nanoparticles (TiO2 NPs) increasing the radical detoxifying capacity and antioxidant enzyme ac­
This is one of those nanoparticles which was required at very low tivities, which significantly help to regulate various physio-biochemical
concentration, as reported (Table 1). In case of Geranium and Poinsettia, and metabolic processes in plants (Fig. 2). They have demonstrated a
the foliar spray of TiO2 NPs accelerated the rate of oxidation reactions substantial influence on how plants respond to stress, mostly by medi­
inside cell, which led to reduction in lesions on leaves (Norman and ating the physiologically and/or environmentally influenced up-
Chen, 2011). Apart from this, it also disrupted the sporangial membrane regulation of various tolerance gene expression and proteins those al­
of Cercospora beticola, which eventually resulted in the death of the ters the cascade of various biochemical pathways and significantly
fungus (Hamza et al., 2016). contributes in managing stress tolerance.

4.1.8. Others 4.2.1. Drought stress

Excluding the above-mentioned one, carbon (C), gold (Au), calcium One of the most important abiotic factors that lower agricultural
phosphate (Ca3(PO4)2), cerium dioxide (CeO2), magnesium oxide output and quality is probably drought. It happens as a result of dynamic
(MgO), oleyl-chitosan (O-Ch), and potassium silicate (K2SiO3) are some variations in temperature, low rainfall and light intensity that lead to
important metallic and/or non-metallic compounds used to generate moisture stress conditions (Kacholi and Sahu, 2018). Maintaining the
nanoparticles for developing resistance against various biotic stresses water content of the leaf, raising the amounts of chlorophyll, anthocy­
(Table 1). Among them, calcium phosphate nanoparticles elevated anin, and carotenoids, and improving the ROS scavenging capabilities
peroxidase and catalase activity for inhibiting black scurf disease inci­ all contributed to the ability to withstand drought stress. The duration
dence in potato (Solanum tuberosum) (El-Shewy, 2019), whereas Adisha and severity of the stress are crucial considerations during a drought
et al. (2018) reported that cerium oxide nanoparticles, at a higher dose, since they may be directly related to crop production and economic
enhanced the photosynthetic efficiency as well as antioxidant produc­ output losses. Plants counter to drought stress in a variety of ways,
tion in order to reduce the disease spread by Fusarium oxysporum f. sp. altering their growth features, biochemical processes as enzymatic
lycopersici in tomato. On the other hand, deposition of carbon nano­ antioxidant activities, and contents of proteins and metabolites (Selei­
materials (CNMs) on spore surface restricted the pathways of water man et al., 2021). Applying phytogenic NPs to plants is a promising
uptake, which ultimately resulted in plasmolysis of the cell (Wang et al., approach for reducing the negative impacts of moisture stress (Table 3).
2014). Thus, CNMs limited its spread in host cell. Biochemically, MgO When drought stress prevails, various biochemical, molecular,
NPs produced numerous pathogen resistance proteins, and physiologi­ morphological, physiological, and ecological features and cascades are
cally, it helped in accumulation of β-1,3-glucanase and tyloses in the disturbed (Ortiz et al., 2015). Due to a decline in the water potential and
xylem and apoplast of pith tissues. These combined effects led to turgor of plant cells brought on by a lack of water, plants experience
development of resistance against Ralstonia solanacearum in tomato growth inhibition and failure to reproduce as a result of an increase in
(Imada et al., 2016). the levels of the solute in the cell’s cytosol and extracellular matrix.
Abscisic acid along with compatible osmolytes builds up and causes
4.1.9. Composite nanoparticles wilting (Hussain et al., 2019). The elevated levels of ROS and the
Composite nanoparticles refer to the compound formed with two or presence of antioxidants like glutathione and ascorbate make adverse

13
B. Pramanik et al. Plant Physiology and Biochemistry 201 (2023) 107831

Fig. 2. Mechanism of the effect of abiotic stresses on plant (physiological and biochemical aspects) (Source: Kumari et al., 2022).

effects worse (Hussain et al., 2019). There are various drought resistance 4.2.2. Salinity stress
systems in plants. As a result, in reaction to unfavorable circumstances, Osmotic and ionic stressors, salinity stress impacts various funda­
plants can limit resource use and control growth. At the molecular level, mental metabolic processes of plants (Parida and Das, 2005). The ionic
these reactions regarding drought stress are improved by signal trans­ balance, cellular metabolism, and membrane malfunction are all dis­
duction (Zandalinas et al., 2020). The molecular basis of plant response rupted by an excessive buildup of sodium (Na+) and chloride (Cl− ) ions,
involves the modulation of plant stomata through increased ion trans­ which inhibits plant growth and development (Munns and Tester,
portation, gene expression, and Abscisic acid signal transduction. When 2008). Maintaining ionic homeostasis and osmotic potential provide as
the physical preparedness of root systems and foliage is insufficient to the critical parameter for developing salinity tolerance responses in
respond against drought-related signals, plant improvement may be stressed plants in order to counteract these antagonistic reactions.
carried out by incorporating genes that encode regular proteins and According to a recent study, the external application of the salicylic
various signals by crosstalk, which expresses a large number of other acid + nano-Fe2O3 to Ajwain (Trachyspermum ammi L.) significantly
genes in accordance with various regulatory mechanisms (Shahid et al., reduced salinity stress through improving K+ absorption, K+/Na+ ratio,
2020). iron content, several antioxidant enzymes, including sodium oxide dis­
By modifying the C3 cycle’s enzyme activity and controlling the mutase (SOD), peroxidase (POD), catalase (CAT), and phenol peroxidase
photosynthetic pigments necessary for plant growth, nanotechnology (PPO), and the amount of the compatible substances. Together, these
offers the potential to increase plant photosynthesis efficiency. The changes improved the membrane stability index, osmotic potential,
effectiveness of NPs on crop growth and germination varies depending anthocyanin contents, and plant growth. It’s interesting to note that the
on their level and host crop. Nano wax foliar spraying on sorghum NPs are additionally known to safeguard plants against ionic toxicity
moisture stress improved seed output in plants compared to water under salinity stress and have been shown to control mineral homeo­
spraying. Further observed was the possibility that NPs applied as a stasis and increased proline concentration and antioxidant activity of
foliar application could boost soybean photosynthesis, hence increasing enzymes while reducing ROS (H2O2, O2, superoxide radical, and MDA)
yield. When a beneficial bacteria colonizes the roots, these types of re­ generation in response to applied salt stress (Yue et al., 2017). Numerous
actions frequently take place. Since the P5CS gene encodes for proline studies have clarified the role of various NPs in reducing the difficulties
biosynthesis, expression of this gene increases a plant’s resistance to caused by salt stress in plants (Table 4).
many environmental stress situations, which included biotic and abiotic NPs are the successful supplement for triggering stress-responsive
challenges. Conjointly Ca21 and ROS, MAPK2, a MAP kinase gene, is signalling has also been established. It’s interesting to note that the
crucial for controlling phytohormones and antioxidant defence systems NPs are also known for safeguarding plants from ionic toxicity under
in response to various stress situations (Pérez-Labrada et al., 2020). The salinity stress and have been shown to regulate nutritional homeostasis.
transcriptional regulators AREB/ABF are essential for the control of the biosynthesized AuNPs applied as foliar spray had a significant effect on
AREB gene, which encodes ABA, and they play a key role in promoting the ionic contents of the shoot and root, in addition to enhanced nitro­
tolerance to adverse environmental conditions such as salt stress and gen (N) metabolic rate, enzymatic antioxidant production (SOD, GPX,
drought. In response to stress, the ZFHD gene is down-regulated, which APX, and GR), besides non-enzymatic antioxidant encompasses (AsA
is regulated by the abscisic acid production pathway. Conversely, and GSH), with a decrease in the production of ROS and lipid peroxi­
downregulation of the TAS14 gene increases solute aggregation, dation that occurred under salinity stress conditions (Yue et al., 2017).
including K1 and sugars, and lowers osmotic pressure, making the plants MiR164 expression is down regulated during NP-mediated root
more resilient to moisture and salinity stress (Pérez-Labrada et al., development, which is associated with auxin hormone signalling.
2020). Lateral roots can form and blooming can be sped up as a result of
elevated miR169 transcription and reduced miR167 expression

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B. Pramanik et al. Plant Physiology and Biochemistry 201 (2023) 107831

Table 3
Impact of various nanoparticles on drought stress management in various plant species.
NPs Conc. (ppm Size Shape Plant Method of Response Mechanism References
used or mg/l* or (nm) application
mg/kg#)

Ag 10 μg/ml Lens culinaris Seed Enhanced germination Increased activity of antioxidant defense Hojjat.
treatment percentage, shoot and root system and accumulation of osmolytes (2016)
length
Cu 4.444# 30–40 Zea mays Soil Enhanced plant growth and Stabilized leaf water potential, and Nguyen et al.
application biomass enhanced chlorophyll and carotenoid (2020)
content
Fe 100# 50–100 Triticum Soil Enhanced plant height, spike Eradication of oxidative stress by Adrees et al.
aestivum application length, dry weight, and activating antioxidative defence (2020)
chlorophyll A content. mechanism.
Si 100# ≤50 Triticum Soil Increased shoot and root Ameliorated oxidative stress alongside Khan et al.
aestivum application length along with biomass decreased MDA and H2O2 content, and (2020b)
electrolyte leakage.
SiO2 100* 10–30 Amorphous Crataegus sp. Soil Increased plant biomass and Significant improvement in Ashkavand et
sp. application xylem water potential at photosynthetic rate, stomatal al. (2015)
seedling stage conductance, and electrolyte leakage
3200 Gossypium sp. Seed Enhanced plant vigour, boll Elevated phenolic content, total free Shallan et al.
treatment formation and ultimately amino acids, and proline content, (2016)
yield increase increased antioxidant activity of various
enzyme like CAT, POD, and SOD
TiO2 100* 10–25 Linum Foliar Increased number of capsules Decreased H2O2 and MDA content Aghdam et al.
usitatissimum application per plant increased in plants, (2016)
enhanced seed weight
50 Gossypium sp. Seed Enhanced plant vigour, boll Elevated phenolic content, total free Shallan et al.
treatment formation and ultimately amino acids, and proline content, (2016)
yield increase increased antioxidant activity of various
enzymes like CAT, POD, and SOD
ZnO 1.0% 18 Globular Triticum Urea coated Reduction in panicle Enhanced Zn uptake Dimkpa et al.
aestivum application initiation time and (2020)
significantly increased grain
yield
100* Mangifera Foliar Enhanced leaf area, total Enhanced leaf NPK content, total Elsheery et al.
indica application yield, phenolics and several carbohydrates, TSS, and proline content, (2020)
physiochemical and SOD, POX, and CAT activities
characteristics
100* 20–30 Triticum Foliar Improved biomass, total Enhanced activities of SOD, POX, etc. Adrees et al.
aestivum application growth and crop yield. (2021)
100 Solanum Foliar Improvement in plant Improved macro- and micronutrients’ Semida et al.
melongena Application growth and productivity uptake capacity and relative water (2021)
content

Abbreviations: μg: Microgram, Ag: Silver, CAT: Catalase, Conc.: Concentration, Cu: Copper, Fe: Iron, H2O2: Hydrogen peroxide, K: Potassium, kg: Kilogram, l: Literre,
MDA: Malondialdehyde, mg: Milligram, N: Nitrogen, -: Not mentioned, nm: nanometre NPs: Nanoparticles, P: Phosphorus, POD: Peroxidase, ppm: Parts per million, Si:
Silicon, SiO2: Silicon dioxide, SOD: Superoxide dismutase, TiO2: Titanium dioxide, ZnO: Zinc oxide.

(Tolaymat et al., 2017). Under salinity stress, rapeseed plants (Brassica deal with stress, the effect of heavy metals on the plant’s system both
napus L.) were treated with foliar Zn NPs, which decreased the levels of chemically and mechanically varies depending on the kind of plant. In
expression of some genes, including MYC, SKRD2, and MPK4, and plant species like poaceae (rice), solanaceae (potato, pepper, tomato
increased the expression of others, including ARP and MPK, which are etc.), fabaceae (soybean, lentil, etc.), and brassicaceae (Arabidopsis,
involved in hormonal and physiological reactions and mustard, etc.), heavy metals are transported by the naturally occurring
transcription-related factors, as well as MYC and SKRD2, which are resistance-associated macrophage protein (NRAMP) gene family (Tian_e­
essential for plant cells to tolerate abiotic stress (Hezaveh et al., 2019). t_al_2021). Si NPs inhibit the expression of the NRAMP5 and low-affinity
Proteomics investigation of tomato plants under salinity stress revealed cation transporter 1 (LCT1) genes, which are involved in facilitating the
that Si impacted the genes for cytochrome bb6f (Cytb6f), movement and intake of Cd from the root to the shoot. Both the silicon
ATP-synthesizing complexes, and photosynthesis. This component, as absorption gene low silicon rice 1 (LSI1) and the heavy metal ATPase 3
demonstrated by Siddiqui et al. (2020), was also implicated in upregu­ (HMA3) gene, are upregulated which transports Cd into the vacuoles.
lating the expression of the OsNAC protein, a stress-response protein. Ahmed et al. (2021a) discovered Cd transporter genes in the phloem and
xylem, including OsHMA2 and OsHMA3, which are in charge of heavy
4.2.3. Heavy metal stress metal transportation, and OsLCT1, which is in charge of Cd trans­
Rapid globalisation severely pollutes the environment by releasing location. All three genes expressed much less in rice after using hydrogel
more harmful metals into the atmosphere. Heavy metals harm soil dy­ and FeO nanoparticles. Phytogenic NPs, however, have the capacity to
namics after they are deposited in the soil as well as microbiological induce tolerance mechanisms by modulating the overall growth of the
structure (Emamverdian et al., 2015), resulting in decreased in crop plant and physiological functions in exposure to heavy metal stress, as
productivity and soil health. Crop failure is frequently caused by heavy shown by a number of studies (Table 5).
metal stress, which has a negative impact on the water potential,
photosynthesis, and various physiological process (Nazir et al., 2020). 4.2.4. Flood stress
In plants exposed to heavy metals, NPs retain membrane stability, Fields may flood if there is improper drainage after a period of heavy
increase chlorophyll pigments, and accelerate photosynthesis (Ahmed rain. Flooding is also brought on by river overflow and a rising
et al., 2021b), each plant species has a unique defence mechanism to groundwater table, both of which have the potential to wipe out the

15
B. Pramanik et al. Plant Physiology and Biochemistry 201 (2023) 107831

Table 4
Impact of several nanoparticles on salinity stress management in various plant species.
NPs Conc. Size Shape Plant Mode of Response Mechanism References
used (mg/l or (nm) application
mg/kg*
or ppm#)

Ag 1.5 – – Solanum Seed priming Increased germination Upregulation of genes like Almutairi et al. (2016)
lycopersicum percentage, germination AREB, MAPK2, P5CS and
rate, root and shoot CRK1, along with
length, and weight of the downregulation of TAS14,
seedling DDF2 and ZFHD1 genes
30 20 Spherical Trigonella Seed priming Improved seed Optimum osmotic Hojjat and Kamyab
foenum-graecum germination adjustment and ion (2017)
homeostasis
2 mM 15–29 Spherical Triticum aestivum seed priming Improved biomass and Enhanced total soluble Mohamed et al. (2017)
yield due to increase in sugars and proline contents,
water potential, and POD activity, and decreased
osmolytes accumulation CAT activity
1 – – Triticum aestivum seed priming Improved various growth Triggered IBA, NAA, BAP, Abou-Zeid and Ismail
parameters, and and various pigment (2018)
photosynthetic efficiency contents, and reduced ABA
as well as phytohormonal content
balance
20 mM 50–100 – Pennisetum seed priming Reduced oxidative Enhanced relative water Khan et al. (2020a)
glaucum damage by improving and proline content,
various antioxidant decreased Na+/K+ ratio,
enzymatic activities total phenolic and flavonoid
content
300# – – Triticum aestivum Foliar spray Enhanced proline Enhanced ion homeostasis, Wahid et al. (2020)
metabolism and N non-enzymatic antioxidant
assimilation, and content such as AsA and
reduction in peroxidation GSH, and activities of SOD,
of lipids APX, GR, and GPX
Au 300# – – Triticum aestivum Foliage Improved N assimilation Maintenance of ion Wahid et al. (2022)
application and antioxidant enzymatic homeostasis, and various
activities non-enzymatic compounds
like AsA and GSH
C 20 – – Brassica napus Seedling Enhanced plant salinity Reestablishment of redox Zhao et al. (2019)
treatment tolerance balance and ion
homeostasis
50 – – Ocimum Mixed with Enhanced various Induced phenolic Gohari et al., 2020
basilicum Hoagland physiological parameters compounds and
solution such as chlorophyll and antioxidative enzymes like
carotenoid content, and APX, CAT and GP activity
essential oil content
Ce 1000* 20–110 – Brassica napus Sand and clay Higher plant biomass and Enhanced efficiency of the Rossi et al., 2016
mixture chlorophyll content photosynthetic apparatus
500* 52.6 Spherical Brassica napus Mixed with dry Several anatomical Reduced apoplastic barriers Rossi et al., 2017a
sand changes in plant roots led to allow higher Na+
to better physiological transportation to shoots
performance
Cu 10 – – Solanum Mixed with Increased stem diameter Proper regulation of Hernández-Hernández
lycopersicum nutrient solution and plant height oxidative stress ionic et al., 2018
balance by upregulating
gene expression of JA and
SOD
250 20–50 Spherical Solanum Foliar spray Enhanced plant height, Increased antioxidant Pérez-Labrada et al.,
lycopersicum fresh and dry weight activity, phenolics, vit-C, 2019
glutathione, and Na+/K+
ratio
Fe 2000 – – Helianthus annus Foliar spray Improved shoot dry Increased antioxidants and Torabian et al., 2018
weight and seed yield soluble carbohydrates
content, and decreased
MDA content and ROS
generation and
maintenance of osmotic
balance
0.8# – – Strawberry Mixed with Improved growth and Significant increase in the Mozafari et al., 2018
(Fragaria × culture media pigment content, RWC, total protein content, and
ananassa) MSI, and Fe and K content activities of various
of the mature plants antioxidative enzymes like
POD, CAT, and SOD
3 mM – – Trachyspermum Foliar spray Improved membrane Improved K+ uptake, Fe Abdoli et al., 2020
ammi stability index, leaf water content, SA, activities of
content, leaf pigments, antioxidative enzymes like
root and shoot growth, SOD, CAT, peroxidase, and
and seed yield
(continued on next page)

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B. Pramanik et al. Plant Physiology and Biochemistry 201 (2023) 107831

Table 4 (continued )
NPs Conc. Size Shape Plant Mode of Response Mechanism References
used (mg/l or (nm) application
mg/kg*
or ppm#)

polyphenol oxidase and

several osmolytes
K 235 – – Medicago sativa Mixing with Enhanced shoot dry Reduced electrolyte El-Sharkawy et al.,
Hoagland’s weight, relative yield, root leakage, Increased activities 2017
solution length and root dry weight of CAT, proline, and several
antioxidant enzymes
Mn 1 20–30 Spherical Capsicum Seed priming Improved root growth as Controlled oxidative stress, Ye et al., 2020
annuum well as yield and upregulation of Mn-
SOD expression to scavenge
ROS
Mn3O4 1 – – Cucumis sativus Foliar Increased plant biomass, Increased accumulation of Lu et al., 2020
application and chlorophyll content photosynthates, and
endogenous antioxidants
PHF 120 mM – – Triticum aestivum Seed pre- Higher chlorophyll, free Decreased MDA and H2O2 Shafiq et al., 2019
treatment amino acids, vit-C, and content, and enhanced
soluble sugars content, antioxidative enzymatic
higher K+content in root activities of CAT, POD, and
and P contents in both root APX
and shoot
Si 1 mM 20–30 – Glycine max Seed treatment Improved plant dry Increased antioxidant Farhangi-Abriz and
weight, and enhanced activities, and non- Torabian (2018)
accumulation of proteins, enzymatic compounds, but
chlorophyll and phenolic decreased Na+ conc., lipid
compounds peroxidation, and ROS
200* 10 – Cucumis sativus Soil application Improved growth, Enhanced N and K uptake, Alsaeedi et al., 2019
productivity, and NUE regulated ion homeostasis
and osmotic balance
50 10–20 – Strawberry Foliar Enhanced salt tolerance Accumulation of less Avestan et al., 2019
(Fragaria × application along with better proline, and properly
ananassa) chlorophyll, and developed epicuticular wax
carotenoid content structure on leaves
100 50–100 Spherical Triticum aestivum Hoagland’s Improved salt stress Well-developed cell wall Mushtaq et al., 2019
solutions tolerance, germination, extensibility in the apical
and chlorophyll contents regions, and decreased Na
uptake
100 – – Musa acuminata Added with Increased chlorophyll Lowered electrolyte leakage Mahmoud et al., 2020
culture media content, and improvement and MDA content, and
in shoot number and proper adjustment of
length various phenolic
compounds
500 – – Solanum Added with Increased fruit weight as Maintained chlorophyll Pinedo-Guerrero et al.,
lycopersicum culture media well as yield conc., activities of GSH, 2020
PAL, and vit-C
TiO2 60# 10–25 Amorphous Zea mays Seed priming Improved germination Enhanced K+ conc., relative Shah et al., 2021
. percentage, germination water content, total phenol
energy, and seedling vigor and proline contents, SOD,
index, and root and shoot CAT, and PAL activities,
length, fresh and dry and lowered Na+ ion
weight, and reduced mean concentration, MDA, and
emergence time electrolyte leakage
ZnO 15 – – Solanum Hoagland Improved antioxidant Higher transcription of SOD Alharby et al., 2016
lycopersicum solution→ Soil defense system leading to and GPX genes
transplantation increased biomass.
2000 – – Helianthus annus Foliar spray Elevated biomass, and High proline content and Torabian et al., 2016
various antioxidant SOD activity
enzymes
60 – – Lupin ustermis. Seed priming Increased conc. of Enhanced activities of SOD, Abdel et al. 2017
chlorophyll, phenolics, CAT, and POD and reduced
vit-C, and various organic MDA and Na+ content
solutes
10 30 – Brassica napus Foliar spray Improved water retention Upregulated oxidative Farouk and Al-Amri
and photosynthetic rate defense system by osmotic (2019)
adjustment, and ionic
regulation
10 16–35 Spherical Abelmoschus Foliar spray Increased chlorophyll Enhanced activities of SOD Alabdallah and
esculentus content, and increased and CAT, and decreased Alzahrani (2020)
yield. proline accumulation and
total soluble sugars
40 – – Solanum Soil application Increased plant growth Alteration in cytosine Haliloglu et al., 2020
lycopersicum and development methylation to induce
positive anti-genotoxic
effect and facilitate DNA
(continued on next page)

17
B. Pramanik et al. Plant Physiology and Biochemistry 201 (2023) 107831

Table 4 (continued )
NPs Conc. Size Shape Plant Mode of Response Mechanism References
used (mg/l or (nm) application
mg/kg*
or ppm#)

repair pathway for reducing

DNA damage
1000* 10–30 – Trigonella Foliar Improved antioxidant Increased proline and Noohpisheh et al., 2020
foenum-graecum application defence mechanism and trigonelline content,
photosynthesis, and antioxidative enzymes, and
increased biomass decreased MDA, aldehydes,
sugars, H2O2
50 16–35 Spherical Solanum Foliar spray Enhanced photosynthetic Increased chlorophyll Faizan et al., 2021
lycopersicum machinery, root and shoot content and other
length, biomass, and leaf photosynthetic attributes,
area protein content, and
activities of various
antioxidative enzymes like
POX, SOD, and CAT
17 – – Carthamus Foliar spray Enhanced yield and yield Upregulation of ion Yasmin et al., 2021
tinctorius attributes like number of homeostasis, and
capitulum per plant antioxidant defence
mechanism to minimize the
ROS production

Abbreviations: ABA: Abscisic acid, Ag: Silver, APX: Ascorbate peroxidase, AREB: abscisic acid response element-binding protein, AsA: Total ascorbate content, Au:
Gold, BAP: 6-benzylaminopurine, C: Carbon, CAT: Catalase, Ce: Cerium, CRK1: Cysteine-rich receptor-like protein kinase 42-like, Cu: Catalase, DDF2: Dwarf and
delayed flowering 2, Fe: Iron, GP: Guaiacol peroxidase, GPX: Glutathione peroxides, GR: Glutathione reductase, GSH: Glutathione, H2O2: Hydrogen peroxide, IBA:
Indole-3-butyric acid, JA: Jasmonic acid, K: Potassium, kg: Kilogram, MAPK2: Mitogen-activated protein kinase 2, MDA: malondialdehyde, , mg: Milligram, ml:
Milliliter, mM: Milli molar, Mn: Manganese, Mn3O4: Tri-manganese tetra-oxide, MSI: membrane stability index, N: Nitrogen, Na: Sodium, NAA: 1-Naphthalene acetic
acid, -: Not mentioned, nm: nanometre, NPs: Nanoparticles, NUE: Nutrient use efficiency, P:Phosphorus, P5CS: Delta-1-pyrroline-5-carboxylate synthetase, PAL:
Phenylalanine ammonia lyase, PHF: Polyhydroxy fullerene, POD: Peroxidase, ppm: Parts per million, ROS: Reactive oxygen species, RWC: Relative water content, SOD:
Superoxide dismutase, TAS14: Abscisic acid and environmental stress inducible protein, ZFHD1: Zinc finger homeodomain transcription factor.

standing crops. Additionally, excessive irrigation might cause temporary downregulation of NmrALike is substantially activated by Al2O3NPs
waterlogging or flooding. All of these circumstances could result in under flooding, according to a transcriptional study of genes that are
either a partial anaerobiosis (hypoxia) or a total anaerobic state susceptible to Al2O3NPs. Conversely, under flooding, Al2O3NPs suppress
(anoxia), which would exhaust the available soil air (Vartapetian et al., the Flavodoxin-Like Quinone Reductase (FQR1) gene (Mustafa et al.,
2014). Flooding not only results in a lack of oxygen, whose diffusion 2015).
velocity is 104 times slower in water than in air, but also in the pro­ FQR1 and NmrA-Like both are under NmrA-like family genes, which
duction of poisonous chemicals that stunt plant growth and frequently possess significant antioxidant properties and reactive oxygen species
leads to in plant death (Armstrong and Drew, 2002). In Hypoxia and (ROS) scavenging capacity (Kim et al., 2010). Transcriptome study in­
anoxia condition results in energy-deficit conditions, due to inhibition of dicates that in comparison with flooded plants, AgNPs suppress the
respiration, and upregulation of ethylene and abscisic acid expression of genes for pyruvate decarboxylase 2 (PDC) and alcohol
synthesis-related genes as strategies for adaptation to waterlogging dehydrogenase 1 (ADH). Pyruvate undergoes catalytic action by the PDC
(Komatsu et al., 2009). Other coping mechanisms include the produc­ enzyme in the very initial step of the alcohol fermentation pathway,
tion of nodal roots during the air-water transition and the growth of which results in the production of CO2 and acetaldehyde. The ADH
aerenchymatous cells in the root cortex to aid oxygen diffusion enzyme then reduces acetaldehyde to ethanol.
(Yamauchi et al., 2013). Therefore, it is crucial to understand if NPs Beta Ketoacyl Reductase 1 (BKR1) is the most interacting protein
increase plants’ ability to withstand floods or reduce its negative con­ when exposed to AgNPs at a thickness of 15 nm; nevertheless, despite
sequences (Bailey-Serres and Colmer, 2014). this, its enzyme activity is reduced. BKR1 has this pattern of expression
Following exposure to NPs under flooding stress, the transcriptomics when under flooding stress. The expression of this enzyme, however,
and proteomics study of the soybean (Glycine max L.) plant were declines while its enzyme activity steadily rises over the post-flooding
analyzed. The results of each study showed that AgNPs and Al2O3NPs recovery phase (Khan et al., 2015). The production of fatty acids and
can mitigate the negative impacts of flooding stress. In plants that were wax is the primary function of the BKR1 enzyme. According to the
not treated, plants that were subjected to flooding, and plants that were findings of Mustafa et al. (2016), AgNPs appear to play a protective role
exposed to Al2O3NPs while they were under flooding, the expression of in flooding by controlling the production of wax and amino acids.
172 common proteins drastically changed, according to a proteome Together, the three studies (Mustafa et al., 2015a, 2015b, 2016) suggest
study. These proteins are involved in metabolism, which is affected less that the NPs’ defense against flooding stress may involve controlling the
by Al2O3NPs than by flooding stress. Al2O3NPs caused the induction of production of protein and the purification of harmful substances, as well
genes involved in protein synthesis, posttranslational alterations, as limiting root cell death and lowering O2 starvation stress by switching
glycolysis, and lipid metabolism. Glycolysis and the fermentation pro­ from aerobic to anaerobic metabolism of energy.
cess can take over aerobic energy metabolism in the absence of oxygen Furthermore, some other physiological and biochemical strategies
(Sousa and Sodek, 2002). Interestingly, when flood stress is prevalent, such as switching from glycolysis to fermentation in the metabolism of
Poly(A)-Binding Protein 2 (PABP2) is the most reactive protein in soy­ carbohydrates. (Banti et al., 2013), in tolerant species, and down­
bean root, those plants treated with Al2O3NP. In order to promote the regulation of the alcohol dehydrogenase enzyme (Mustafa et al., 2015b).
initiation of translation, this protein attaches to the poly(A) tail of an Further thoughtful research is necessary in light of the apparent dearth
mRNA (Mustafa et al., 2015). The PABP2 gene belongs to a group of of evidence on the relationship between NPs and plants. However, based
cytoplasmic particles called stress granules that protect RNAs from on the facts at hand, it may be hypothesised that NMs are essential for
harmful circumstances inside the cell when a cell is under stress. The reducing the negative consequences of flooding stress by altering

18
B. Pramanik et al. Plant Physiology and Biochemistry 201 (2023) 107831

Table 5
Impact of different nanoparticles on heavy metal stress management in various plant species.
NPs Conc. (mg/ Size (nm) Shape Plants Mode of Response Mechanism References
used kg or mg/l*) application

Ce 500 41.7 ± 5.2 – Glycine max Sand mixing Significant increase in the Improved Fv/Fm ratio and Ce Rossi et al., 2017b
accumulation of Ce in roots transportation through roots
and older leaves.
200 9–22 Spherical Oryza sativa Hydroponics Improved crop growth, Reduced activities of SOD, POD Wang et al., 2019
photosynthesis and the level of and MDA
8-OHdG
Cu 100 <100 – Triticum Soil Rise in plant height and Reduced Cd transport, Noman et al.,
aestivum application biomass, increase in N and P maintained ion homeostasis, 2020
uptake and increased SOD and POD
content
50 – – Enhanced in growth and Reduced Cr bio-availability,
biomass elevated uptake of essential
nutrients and antioxidant
content
Fe 20* 50–100 – Triticum Seed priming Enhanced plant growth, Decreased loss of electrolytes Rizwan et al.,
aestivum biomass, and grain number and activity of SOD and 2019a
peroxidase enzymes
Fe3O4 2000* 6.8 – Triticum Seedling Minimization of the inhibitory Improved the level of SOD and Konate et al.,
aestivum exposure action of various heavy metals peroxidase, and ion 2017
homeostasis.
100 – – Oryza sativa Soil Retardation in the Augmented detoxifying Ahmed et al.,
application mobilization of Cd, improved enzymes, reduced ROS activity, 2021a
biomass, chlorophyll content lowered gene expression
supporting Cd transport
100 – – Triticum Soil Increase in plant length, and Blocked Cd transport, increased Manzoor et al.,
aestivum application biomass NPK uptake, increased 2021
antioxidants and pigment
content
Si 10 μM – – Pisum sativum Seed priming Improved growth in presence Enhanced synthesis of defense Tripathi et al.,
of Cr enzymes, maintained ion 2015
homeostasis and augmented
nutrient uptake
1.0 mM 19 – Oryza sativa Foliar Reduced metal uptake Proper maintenance of cellular Cui et al., 2017
application integrity, and altered
expression of genes responsible
for metal acquisition
2000* 30 – Glycine max Hydroponics Increased seedling vigour and, Repression in the growth Li et al., 2020
photosynthates by enhanced inhibitory action of Hg, and
chlorophyll content. reduced its content in root and
shoot
Ti 100* 21 Amorphous Oryza sativa Hydroponics Significant increase in net Enhanced activities of SOD and Ji et al., 2017
photosynthetic rate, and POD; reduced MDA and Cd
chlorophyll content accumulation in roots and
leaves
0.1–0.25% – – Vigna radiata Seed priming Augmented radical length and Minimized ROS content and Katiyar et al.,
biomass lipid peroxidation, 2020
upregulation of gene expression
of antioxidative enzymes
100-250* 6.5 ± 0.76 – Zea mays Foliar Significant lowering in the Enhanced the activities of SOD Lian et al., 2020
application accumulation of Cd in shoots and GST, and upregulation in
the metabolic pathways to
alleviate Cd stress
100* – – Zea mays Foliar Increased shoot and root dry Elevated activities of Zhou et al., 2020
application weight antioxidative enzymes
ZnO 25* – – Leucaena foliar Diminished heavy metal Improved photosynthetic Venkatachalam
leucocephala application induced toxicity, enhanced pigments and soluble proteins, et al., 2017
seedling growth, antioxidant defense enzymes,
photosynthetic activity and decreased peroxidation
100* – – Zea mays Foliar Spray Cd content lowered in root and Enhanced gas exchange Rizwan et al.,
shoot along with enhanced characteristics, and boosted 2019b
plant growth, biomass, and activity of antioxidant enzymes
chlorophyll content
– – Triticum Seed priming Improved plant growth, Decreasing loss of electrolyte,
aestivum biomass, and grain number, and activity of SOD and
and reduced Cd accumulation peroxidase
100* 20–30 Spherical Oryza sativa NA Increased the growth of Elevated phytochelatin level Yan et al., 2021
seedlings, and diminished
accumulation of As in root and
shoot

Abbreviations: μM: Micromole, 8-OHdG: 8-hydroxydesoxiguanosine, As: Arsenic, CAT: Catalase, Cd: Cadmium, Ce: Cerium, Cr: Chromium, Cu: Copper, Fe: Iron,
Fe3O4: Magnetite, GSH: Glutathione, K: Potassium, kg: Kilogram, l: Liter, MDA: Malondialdehyde, mg: Milligram, mM: Millimolar, N: nitrogen, NA: Not applicable, -:
Not mentioned, nm: nanometre, NPs: Nanoparticles, P: Phosphorus, POD: Peroxidase, ROS: Reactive oxygen species, Se: Selenium, Si: Silicon, SOD: Superoxide
dismutase, Ti: Titanium, Zn: Zinc, ZnO: Zinc oxide.

19
B. Pramanik et al. Plant Physiology and Biochemistry 201 (2023) 107831

metabolism and gene expression (Table 6). OsCu/ZnSOD2, OsCu/ZnSOD, OsPRX89, OsPRX65, OsCATA, sPRX11,
and OsCATB) and some transcription factors those are chilling respon­
4.2.5. Temperature stress sive (OsbZIP52, OsMYB30, OsNAC5, OsWRKY76, OsMYB4, and
The atmospheric CO2 concentration has been rising rapidly since the OsWRKY94) (Song et al., 2021). Upon application of ZnO NPs, in soy­
industrial revolution, contributing to global warming. It has been ex­ bean seedlings, up regulated the HSF-34, WRKY1, EREB, HDA3, CAT,
pected that increasing temperatures will negatively affect crop produc­ R2R3MYB, and MAPK1 genes these raised proline content, antioxidant
tivity (Asseng et al., 2015; Kumar 2016), and around the world, efforts enzyme activity, photosynthetic pigments, and ultimately plant yields
are being done to stop the temperature rise and develop sustainable (Mirakhorli et al., 2021).
agriculture. When temperatures reach a certain point, both temperature
extremities i.e. high temperatures (heat stress) and low temperatures 4.2.5.1. Heat stress. Heat stress, also known as high temperature stress,
(cold stress or chilling/freezing stress) start to harm plants. is a surge in temperature that lasts long enough to permanently harm a
NPs are primarily carried by lateral roots from the soil to the plant. plant’s ability to plant growth, development, and yield (Wahid, 2007).
The NPs penetrate the entire plant body through the xylem tissues (Jalil Heat stress induces oxidative stress, which leads to membrane lipid
and Ansari, 2019). The application of NP aids in maintaining ion ho­ breakdown, electrolyte leakage, and biomolecule denaturation. Heat
meostasis in plants, which in turn promotes membrane stability and stress also increases the overproduction of ROS (Savicka and Skute,
osmotic control. Plant productivity is increased when it can continue to 2010; Karuppanapandian et al., 2011; Aref et al., 2016). and decreases
transmit water and nutrients under pressure. Under stressful circum­ the chlorophyll content and photosynthesis rate (Prasad et al., 2011). In
stances, development is also ensured by preventing enzymatic oxidative reaction to heat stress, the molecular chaperons known as heat shock
damage. Under circumstances involving heat and cold stress, the syn­ proteins (HSPs) are produced (Schulze et al., 2005). Other proteins are
thesis of enzymatic anti-oxidizers, such as SOD, POD, CAT, and APX, are helped to maintain their stability under heat stress by HSPs and various
promoted by the use of NPs. Toxicities induced by NPs can be extremely other related solutes (Wahid, 2007) and contribute significantly to
dangerous at greater levels, hence the concentration of NP application is thermotolerance. Nanotechnology’s current advances have modernized
crucial (Wang et al., 2016). the agricultural system and show promise for improving crop growth
In rice ZnO NPs used as foliar spray activates the chilling-induced and development in stressful environments. According to several
gene expression of various anti-oxidative system (OsCu/ZnSOD3, research listed in Table 6, NPs may be used to increase crop plants’

Table 6
Impact of nanoparticles on other abiotic stresses management in various plant species.
NPs Conc. Stresses Size Shape Plant Mode of Response Mechanism References
used (mg/l or (nm) application
mg/kg*
or ppm#)

Ag 80# Flood – – Crocus Corm Increased root Blocked ethylene signalling Rezvani et al., 2012
sativus treatment biomass, and leaf and increased stored
biomass carbohydrates
2# 15 – Glycine max Seed Increased abundance Suppression of cytotoxic by- Mustafa et al., 2015
treatment of stress-related products of glycolysis,
proteins, enhanced regulated ascorbate/
seedling growth glutathione pathway
5 Cold – – Arabidopsis – Improved cold stress Upregulation of MeCu/ZnSOD Kohan-Baghkheirati
thaliana tolerance and MeAPX2 antioxidant and Geisler-Lee (2015)
genes
75 Heat – – Triticum Foliar Protection against Improved osmotic potential by Iqbal et al., 2017
aestivum application heat stress and upregulation of various
significantly organic solutes and osmolytes
improved plant accumulation
growth
CeO2 400* Heat 10 ± 1 – Zea mays Soil Enhanced tolerance Enhanced cross talk among Zhao et al., 2012
treatment response under stress H2O2, heat shock protein and
condition lipid peroxidation
Se 10 Heat 10–40 – Sorghum Applied with Improved chlorophyll Activated H2O2 quenchers Djanaguiraman et al.,
bicolor Hoagland content, biomass, (GPX, CAT, POX, and APX) (2018)
solution pollen germination, and decreased MDA
and seed set production
TiO2 5 Cold 7–40 Amorphous Cicer Foliar Improved antioxidant Increase various antioxidants Mohammadi et al.,
arietinum application defence system, and (like SOD, peroxides), 2013
tolerance response decreased H2O2 content and
electrolyte leakage
100 Heat – – Solanum Foliar Increased Regulation of energy Qi et al., 2013
lycopersicum application accumulation of dissipation and various
photosynthates metabolic pathways causing
cooling of plant canopy
through induced stomatal
opening
5 Cold – – Cicer Seed Tolerance against Overexpression of RuBisCO- Hasanpour et al., 2015
arietinum treatment cold stress and chlorophyll-binding
protein genes, enhanced
activity of PEP-carboxylase,
and decreased H2O2 content

Abbreviations: APX: Ascorbate peroxidase, CAT: Catalase, CeO2: Cerium dioxide, Conc.: Concentration, GPX: Glutathione peroxidase, H2O2: Hydrogen peroxide, kg:
Kilogram, l: Liter, mg: Milligram, -: Not mentioned, nm: nanometre, NPs: Nanoparticles, POD: Peroxidase, ppm: Parts per million, RuBisCO: Ribulose bisphosphate
carboxylase/oxygenase, Se: Selenium, SOD: Superoxide dismutase, TiO2: Titanium dioxide.

20
B. Pramanik et al. Plant Physiology and Biochemistry 201 (2023) 107831

ability to withstand heat stress. 5. Artificial intelligence integration in plant stress nanoscience

4.2.5.2. Cold stress. Farmers are especially concerned about the Application of nano-sciences in the form of nano-sensors involves a
frequent and uncommon occurrence of low temperatures. While wide variety of detection for plant hormones, pesticide detection, dis­
freezing stress (below zero degrees) results in the development of ice ease monitoring, investigation of the soil acclimatization (Yata et al.,
crystals in plant tissues and causes frost-killing of crops, cold stress 2017). Amongst which nano-sensors play a vital role in monitoring of
(0–15 ◦ C) causes harm without doing so (Hasanuzzaman et al., 2013). the development and the growth of the plant due to their excellent
The two main impacts of cold stress are loss of membrane fluidity and electrical, optical and the mechanical properties when compared to its
electrolyte leakage. Reduced crop production, poor germination, and loose features. Most of the commonly studied nano-sensor techniques for
delayed growth are all symptoms of plants under cold stress (Welti et al., the detection of the stress induced on the plants are surface enhanced
2002; Suzuki et al., 2008). However, susceptibility to cold stress varies Raman Scattering, fluorescence resonance energy transfer, corona-phase
among species and cultivars. Greater levels of tolerance in plants result molecular recognition (Fig. 3) (Zhang et al., 2013).
in less membrane damage compared to sensitive species. (Heidarvand Nano-sensors work in the living plants for its overall growth, stress
et al., 2011). detection and for the induction of plant signals. Firstly, the numerous
TiO2 NPs may affect the breakdown of energy through various signalling molecules present in the plants, such as gas producing signals,
mechanisms, such as malate, as evidenced by the greater expression of electrical, chemical and phytohormones could be detected with the help
the PEP carboxylase (PEPC) gene in plants that are resistant compared to of the nano-sensors (Tsuchiya et al., 2015). Oxygen and the nitric oxide
susceptible ones (Hatch, 1987). Malate dismutation cascade is a typical signals contribute in response to the biotic and abiotic stress (Giraldo
defence in prolonged oxygen deprivation. Phosphoenolpyruvate pro­ et al., 2015). Nitric oxide detection has been made possible using a
duced by glycolysis is changed by the PEPC enzyme into oxaloacetate, fluorescent ratio metric sensor based on the carbon nanotubes which has
which is then reduced to malate, as part of the anaerobic energy meta­ a potential to respond to the complexity prevailing in the environment
bolism. This malate enters the mitochondrion and there it degrades via (Mittal et al., 2020). Additionally, the calcium signalling molecules are
malate dismutation (Edwards et al., 2013). The PEPC gene is upregu­ connected with the abiotic and biotic stresses. Fluorescent indictors such
lated under cold stress, indicating a change from aerobic to anaerobic as GCaMP and YC3.6 and genetically coded ones are used for ocular
energy metabolism. Photosynthesis, a very crucial physiological process, imaging on the basis of Fluorescence resonance energy transfer tech­
is vulnerable to cold stress. Cold stress causes plants to produce less nique; which provides speedy, noticeable and high likeness behaviour
chlorophyll, assimilate less CO2, and degrade the activity of RuBisCO for the detection of transitory calcium molecules (Wu et al., 2014).
enzyme (Yordanova and Popova, 2007; Liu et al., 2012). Beneficial ef­ Besides, a sensor based on the needle is used for detecting calcium sig­
fects of NMs on photosynthesis promote RuBisCO carboxylation (Gao nalling molecules in the living cells but at the same time doesn’t function
et al., 2006), chloroplast light absorption capacity (Ze et al., 2011), in the plants (Son et al., 2011). Scientists have also discovered sensors
electron transport rate, and inhibition of ROS generation in chloroplast for the rapid detection of the auxin plant hormones like 1-napthalene
(Giraldo et al., 2014) (Table 6). To make a concrete determination acetic acid (NAA) and 2,4 dichlorophenoxyacetic acid (2,4-D) (Tripa­
regarding the function of NPs in improving cold tolerance of several thi et al., 2022a). Volatile compounds are amongst those chemical sig­
significant crops, however, much more research is undoubtedly nals produced by the plants which affect the quality of food. Sensors
necessary. used for sensing the chemical signals are essential in maintaining the
shelf life of the food product also aiding towards stress tolerance. Sci­
entists have also shown the nano-probe based on the surface enhanced
Raman scattering for detecting the several stresses induced impulsive

Fig. 3. Application of nano-sensors in plants against environmental stress (created in Biorender: https://biorender.com).

21
B. Pramanik et al. Plant Physiology and Biochemistry 201 (2023) 107831

molecules in the plants. This nano-probe works in the region of infrared becoming a hindrance regarding this blooming technology. The
region at 785 nm by getting placed in the inner space in order to avoid knowledge of structure, size, surface area, thermal stability etc. must be
any disruption from the autofluorescence produced in the plants. In the considered as the mark of importance for their commercial application.
presence of biotic or abiotic stress, optical sensors can effectively mea­ Hence, in order to harness the maximum potential advantages of NPs,
sure extracellular adenosine triphosphate (ATP), glutathione and the more research works need to be implemented, in order to gain more
level of salicylic acid and phytolexin in Hordeum vulgare, Triticum aes­ knowledge regarding the mechanisms of action of different NPs,
tivum L, Nasturtium officinale which are responsible for the inception of plant–NPs interaction in definite ecosystem at molecular level, optimi­
the plant diseases. Such nano-sensors based on the surface plasmon ef­ zation of proper concentration and size, their effect in source-sink
fect can contribute towards the before time diagnosis of the stress and regulation and most importantly, their negative effects towards the
can lead in the efficient supervision of any plant disease (Son et al., plant body as well as the environment. In a nutshell, this present review
2022). offers a comprehensive concept regarding the multifaceted utilization of
Several studies have revealed the ability in monitoring plant growth a diverse range of NPs as stress management strategies as well as draws
by the process of fabrication using nano-sensor based on the strain by the attention towards the future pathways of research on this newly
the synthesis of graphite powder in the nanoscale range and the chitosan emerged arena.
solution, on direct exposure to the Cucumis sativus (Tang and Lv, 2014).
Upon the introduction of carbon nanotube ink, the results indicated 1%– Authors’ contribution
8% increase in the plant growth within the strain with a gauge factor
value of 64 with 60% stretchability. A higher sensitivity and the reso­ BP conceived the idea; BP, PS, RKG, RB, SP collected data and pre­
lution demonstrated a recurrent pattern in the plant growth (Zhang pared the basic manuscript; BP, PS, DM, SS edited the manuscript; BP
et al., 2022). Environmental factors such as change in the humidity and did some correction and addition to the manuscript; PS, SS incorporated
the temperature conditions are moreover responsible for hampering the the pictorial diagrams; BP, PS, DM, SC revised and finalized the manu­
growth of the plants. Research findings have demonstrated the use of script. All the authors read and approved the manuscript.
nano-sensors based on graphene which are basically placed on the back
surface of the leaves of the plants particularly maize which can easily Funding
detect the alterations in the pattern of relative humidity on the leaf
surface. On irrigation the relative humidity of the leaves changes which The manuscript has not received any funding from any source.
is articulated as the change in the resistance of the nano-sensor. Simi­
larly, the fabrication of nano-sensor based on the relative humidity was Declaration of competing interest
performed by the deposition of gold and titanium on the film made up of
polyimide for forming an electrode of an interdigitated form. Monitoring The authors declare that they have no known competing financial
of the transpiration rate of the stoma was evaluated by the nano-sensor interests or personal relationships that could have appeared to influence
which helps in the conversion of the variation produced in the plants the work reported in this paper.
into the electrical signals which was further sent to the devices present
in the wireless form particularly mobile devices in the real time. The Data availability
invention could be more meaningful when used in combination with the
devices which are wireless in form and could help in analysing the data Data will be made available on request.
but still the stability is questionable and needs in-depth research.
Several other reports have reported the detection of soil pollutants
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