Plant Nano Biology 4 (2023) 100035

Contents lists available at ScienceDirect

Plant Nano Biology

journal homepage: www.journals.elsevier.com/plant-nano-biology

Nanotechnology in improving photosynthesis under adverse climatic

conditions: Cell to Canopy action☆ ]]
]]]]]]
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Fahima Dilnawaza, , Mohamed Hazem Kalajib,c, Amarendra N. Misrad,

⁎ ⁎⁎

a
Department of Biotechnology, School of Engineering and Technology, Centurion University of Technology and Management, Jatni, Bhubaneswar 752050, India
b
Warsaw University of Life Sciences SGGW, Warsaw, Poland
c
Institute of Technology and Life Sciences - National Research Institute, Falenty, Poland
d
School of Applied Sciences, Centurion University of Technology and Management, Jatni, Bhubaneswar 752050, India

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

Keywords: Climate change has a tremendous influence on plants all over the world, creating severe abiotic stress that alters
Nanoparticles crop growth and development, resulting in major productivity losses. Nanotechnology applications in plants
Nanomaterials have the potential to transform present plant production methods by allowing interactions between nano­
Abiotic stress particles (NPs) and plant responses, such as uptake, localization, and activity. Because of their tiny size, high
Uptake and translocation
surface area, absorption rate, effective catalysis of processes, and adequate reactive sites, nanoparticles (NPs) are
Growth and yield
being employed as an emerging tool to stimulate particular biochemical responses connected to plants and their
eco-physiological output. NPs of various kinds are used as beneficial effectors in plants to mitigate the effects of
abiotic stressors. This review focuses on the positive effects of nanoparticles/nanomaterials in plants, as well as
their mode of action for overcoming a range of abiotic stresses.

1. Introduction
Population expansion has raised the need for productivity per
hectare of agricultural land (Ray et al., 2013; Hinz et al., 2020). This
problem has been compounded by agricultural loss, urbanization, soil
degradation, and climate change, resulting in uncertainties about
enhanced production (Kögel-Knabner et al., 2020). Researchers have
discovered novel and critical strategies to increase agricultural pro­
duction in order to fulfill food demand while mitigating the negative
impacts of climate change on crop yields. Significant effort has been
made to improve agricultural practices and boost crop output in order
to offset these possible difficulties. Despite their effectiveness, these
strategies' efficacy has visibly plateaued since the so-called "green
revolution" (Calicioglu et al., 2019). The global population is pre­
dicted to reach around 11 billion people by 2100, necessitating an
increase in food consumption. There is an urgent need to revolutionize
conventional agriculture in order to reach the UN's second sustainable
development target of "Zero Hunger" by 2030. Furthermore, conven­
tional agriculture must be transformed by utilizing a modernized,
sustainable, and environmentally friendly strategic strategy.
Climate change, which is coupled with abiotic and biotic pressures,
has a significant impact on global food production (Zhang et al., 2022;
Gonzalez, 2021). Abiotic stress is the most significant impediment to
plant growth because it slows physiological responses and obstructs
photosynthetic systems (Chaudhry and Sidhu, 2022; Hussain et al.,
2023). Sustainable Development Goal 12, "Responsible Consumption
and Production," has become the global economy's driving force for
sustainable consumption. This objective promotes sustainable lives by
encouraging the use of fewer resources and achieving more with less.
Goal 15, "Life on Land," may be met by using novel plant stress man­
agement strategies to stop recurrent drought and desertification
(www.un.org/sustainabledevelopment/sustainable-development-
goals/).
Plants continue to be the primary source of food for life on Earth due
to the intrinsic process of photosynthesis, which employs the specific
cellular component chloroplast. Photosynthesis is required for the
creation of oxygen, the regulation of climate, and the powering of
biological processes such as agricultural output. Crop output is a by-
product of the entire system, not only photosynthesis. Development,
leaf and canopy design, source-sink feedback, and other variables all

This article is part of a special issue entitled: “Mechanistic and physiological insights into plant nanotechnologies” published at the journal Plant Nano Biology.
☆

Correspondence to: Department of Biotechnology, School of Engineering and Technology, Centurion University of Technology and Management, Jatni,
⁎

Bhubaneswar 752050, Odisha, India.

⁎⁎
Correspondence to: School of Applied Sciences, Centurion University of Technology and Management, Jatni, Bhubaneswar 752050, Odisha, India.
E-mail addresses: fahimadilnawaz@gmail.com, fahima.dilnawaz@cutm.ac.in (F. Dilnawaz), misra.amarendra@gmail.com (M.H. Kalaji),
amarendra.misra@cutm.ac.in (A.N. Misra).

https://doi.org/10.1016/j.plana.2023.100035
Received 17 March 2023; Received in revised form 9 June 2023; Accepted 14 June 2023
2773-1111/© 2023 The Author(s). Published by Elsevier B.V. This is an open access article under the CC BY license (http://creativecommons.org/licenses/by/4.0/).
F. Dilnawaz, M.H. Kalaji and A.N. Misra
Fig. 1. Root mediated nanoparticles translocation between the cells through symplastic and apoplastic pathway (a, b).
impact the link between crop yield and photosynthesis (Apostolova,
Misra et al., 2014, Misra et al., 2012a, Misra et al., 2012b). Further­
more, unlike labs or greenhouses, the natural environment has a con­
siderable impact on photosynthesis and development. These interac­
tions and feedbacks have a significant influence on photosynthesis
(Araus et al., 2021). The biological activity under abiotic stresses has a
significant impact on photosynthetic efficiency. Plants under stress
develop reactive oxygen species (ROS), which initiate coordinated
cellular oxidative processes. This causes cellular structural denatura­
tion, protein instability, membrane disintegration, and metabolic dis­
turbance. Excessive ROS generation produces severe oxidative stress in
plants, negatively reducing photosynthesis and yield (Joshi et al., 2013;
Tunc-Ozdemir et al., 2009; Misra, 2018a; Singh et al., 2019; Stefanov
et al., 2019; Moustafa-Farag et al., 2020). Traditional agricultural
practices decrease stress, yet they are insufficient. Managing abiotic
stress is thus the primary objective for increasing crop output when
under abiotic stress. As a result, major attempts to solve this issue with
imminent technology advancements are required. When exposed to
nanoparticles, plant growth hormones are known to participate in a
variety of signaling cascades and act as stress-regulating molecules.
In the context of a changing environment, nanobiotechnology has
gained relevance since it can assist to mitigate the impacts of en­
vironmental stresses and is viewed as a viable approach for enhancing
agricultural productivity (Bora et al., 2022; Singh et al., 2023). Nano­
technology is a multidisciplinary discipline in which scientists and en­
gineers work together to build nanoparticles, nanomaterials, and na­
nocomposites on a nanoscale scale (Dilnawaz et al., 2018; Misra et al.,
2013). Nanotechnology is playing an important role in agriculture. Its
intervention is a new crop production approach (Nadiminti et al., 2013,
Ashraf et al., 2021, Badawy et al., 2021).
Nanoparticles (NPs) have distinct traits such as a high surface-to-
volume ratio, unique physicochemical properties, cargo loading
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Plant Nano Biology 4 (2023) 100035
capacity, biodegradability, and sustained release activity. Furthermore,
due to fast population expansion and environmental deterioration,
traditional agriculture is facing unprecedented problems (Shang et al.,
2019). For many years, fertilizers and insecticides have been critical
components of maintaining agricultural productivity in order to satisfy
food production demands. However, excessive pesticide usage impedes
sustainable agricultural growth (Kumari et al., 2018; Pathak et al.,
2022).
Plant nanobiotechnology, an amalgam of plant biotechnology and
nanotechnology, has the potential to boost agricultural yield. Soil im­
provement mediators, nano-fertilizers, nano-pesticides, growth stimu­
lators, nutrient absorption enhancers, nano-sensors, abiotic stress
management, pathogen detection, plant hormone regulation, environ­
mental pollution reduction, and stress amelioration are all examples of
how nanoparticles and nanomaterials are used to control various agri­
cultural factors on farms (Seleiman et al., 2020, Singh et al., 2022).
Furthermore, plant growth hormones are known to engage in a variety
of signaling cascades when exposed to nanoparticles and to function as
stress-regulating molecules (Manzoor et al., 2022; Sharif et al., 2022;
Tripathi et al., 2022).
Biological activity is particularly sensitive to photosynthetic effec­
tiveness under abiotic stresses. The use of NPs can boost plant pro­
duction by speeding growth, development, and the rate of active pho­
tosynthesis. These applications aid in agricultural production
improvement (Dilnawaz et al., 2018; Manzoor et al., 2022; Misra et al.,
2013; Shang et al., 2019; Wang et al., 2016a). Furthermore, with the
increased use of nanotechnology, agriculture may undergo significant
changes, providing innovative ways to improve sustainable agriculture
and meet food demand through nano-enabled solar energy harvesting
by plants, which could potentially revolutionize agriculture in the fu­
ture (Singh et al., 2021; Djanaguiraman et al., 2018; Tombuloglu et al.,
2019). This review emphasizes the favorable influence of NPs/
F. Dilnawaz, M.H. Kalaji and A.N. Misra
nanomaterials on plants, as well as their mode of operation and strategy
for overcoming a range of abiotic challenges, regarding NPs' impact on
plant photosynthesis.
2. Uptake and translocation of nanoparticles in plants
Nanoparticles can be administered to plants by soil or foliar treat­
ment. Nanoparticle uptake is influenced by their size, shape, surface
charge, and interactions with target cells (Fig. 1) (Pérez-de-Luque,
2017). One of the key hurdles to nanoparticle penetration into plant
tissues is particle size, with size-dependent absorption happening in
plants. Plants have barriers ranging from micrometers (µm) to nan­
ometers (nm). Furthermore, surface functionalization and coating of
nanoparticles can have a considerable impact on plant absorption and
accumulation (Judy et al., 2012). Stomata are two-guard cell pores that
open and close to control gas exchange between the plant and its en­
vironment.
In foliar applications, nanoparticles confront the cuticular resistance
of the leaf epidermis, which has stomata surrounded by guard cells, and
the pore size increases larger during gas exchange (Hu et al., 2020). As
a result, nanoparticles given to plants can enter through stomatal holes.
The characteristics of the cuticle layers and trichomes around the sto­
mata also influence permeation efficiency (Wang et al., 2016a; Wang
et al., 2016b).
In an investigation, lettuce foliar was exposed to freshly-made TiO2-
NPs, and titanium internalization was examined using microscopic and
spectroscopic approaches. TiO2 -NPs were discovered in all sorts of
tissues after being internalized in lettuce leaves (Larue et al., 2014). In
another study, hydrophilic suspensions of carboxy-modified fluorescent
polystyrene nanoparticles of two sizes (43 nm and 1.1 m) were applied
adaxially between the leaf veins of one-week-old Vicia faba plants kept
in a water-saturated atmosphere to investigate size exclusion stomatal
foliar uptake. Larger particle penetration was not detected after
2–9 days, while smaller particles occasionally penetrated and dis­
seminated in the leaf interior through stomatal pores at different
depths, as shown by leaf slices under confocal scanning microscopy
(CLSM). A substantial number of particles were densely deposited in the
areas of cuticular margins and stomatal throats, and some were found
below the aperture of the substomatal cavity (Schönherr, 2006).
The transport of solutes occurs through lipophilic and hydrophilic
pathways in the cuticular pathway (Fig. 2). The lipophilic route is im­
portant for delivering lipophilic chemicals to the cuticle, whereas the
hydrophilic pathway consists of "aqueous polar pores" that transport
polar or ionic solutes (Schönherr, 2006). Foliar spray of 20 nm Fe3O4-
NPs nanoparticles allowed them to penetrate leaf cells and were then
Fig. 2. Entry of nanoparticles through stomata.
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Plant Nano Biology 4 (2023) 100035
distributed throughout the Nicotiana benthamiana plant, as indicated by
enhanced plant dry and fresh weights (Cai et al., 2020).
Another study used two techniques to give Ag-NPs to plants (foliar
and root exposure). Foliar exposure resulted in 2–7 times more Ag
bioaccumulation than root exposure, showing that AgNP transforma­
tion and/or aggregation takes place in plants (Li et al., 2020a, 2020b).
Quantum dots (QDs) less than 10 nm in size and within the range of
plant wall pores may easily penetrate plant tissues (Wu et al., 2017a).
The two basic mechanisms for NP translocation are the apoplast and the
symplast. The symplastic channel transports NPs via the cytoplasm of
adjoining cells (Roberts and Oparka, 2003), whereas the apoplastic
pathway transports NPs through extracellular gaps associated with
neighboring cell walls and xylem arteries (Fig. 1) (Sattelmacher, 2001).
Plasmodesmata transit allows particles to move from cell to cell and
serves as a cytoplasmic bridge between adjacent cells (Dietz and Herth,
2011). The graphic representation displays several modes of nano­
particle application in plants/soil for enhanced agronomic practices
(Fig. 3).
3. Augmented photosynthetic activity by the involvement of NPs
in abiotic stress condition
Plants, as sessile creatures, are exposed to a wide range of stresses
throughout development. The worldwide loss of plant and crop output
is expected to be decreased by 30 % by 2025 (IPCC, 2012; Khalid et al.,
2021; Chaudhry and Sidhu. 2022). Furthermore, the application of
nanoparticles can improve many photosynthetic indices, hence redu­
cing stress (Fig. 4). The parts that follow go through several stressors
and their impact on photosynthesis, from the cellular to the canopy
level. The stress-relieving properties of various nanoparticles and na­
nomaterials are also investigated.
3.1. Salt stress
Salt stress, also known as salinity, is the second most widespread
stress on Earth, after water or osmotic stress (drought stress) on Earth,
influencing global production (Biswal et al., 2002; Hussain et al., 2023;
Misra et al., 1999; Stefanov et al., 2019; Moustafa-Farag as al., 2020;
Noor et al., 2022). Water transport and absorption are critical for plant
growth and development. One of the primary regulatory routes for
protecting plants from abiotic stressors is transmembrane water trans­
port (Lopez-Zaplana et al., 2022).
Salinity stress lowers root water intake and water channel activity,
decreasing plant survival (Singh et al., 2021). The plant utilizes nu­
merous techniques to minimize oxidative stress, including increasing
F. Dilnawaz, M.H. Kalaji and A.N. Misra
Fig. 3. Different modes of application of nanoparticulate formulations in plants/soil.
and maintaining antioxidant activity and increasing the expression of
certain genes in response to salt. The discovery of a variety of plant
aquaporins (AQPs) in roots shows that these molecules may have spe­
cialized roles in diverse cell types and may be activated in response to
varied environmental stimuli (Gautam and Pandey, 2021). As a result,
salt alters AQP expression, demonstrating that these proteins are es­
sential for maintaining water homeostasis and allowing organisms to
tolerate the impacts of salinity (Sohail et al., 2023). AQPs are a diverse
gene family that functions as a multipurpose channel to transport
substances such as carbon dioxide, hydrogen peroxide, glycerol, me­
talloids, and water (Kumawat et al., 2021).
Among all AQPs, plasma membrane intrinsic proteins (PIPs) present
in the leaves and roots of higher plants are the principal water channels
for regulating water homeostasis (Deshmukh et al., 2016). PIPs have a
vital role in plant tolerance to salt and drought. PIPs play an important
role in managing water homeostasis under salt stress. The pumpkin PIP
gene family was recently found and characterized by a genome analysis
in light of the importance of AQPs for salt stress and stress tolerance,
employing the hairy root transformation strategy by producing
CmoPIP1–4 CRISPR mutants (Sohail et al., 2022). The revelation that
CmoPIP1s expression in leaves and roots reacts to salt stress may aid in
the creation of stress-resistant cucurbit crops in the future (Sohail et al.,
2022). The principal targets of the photosynthetic machinery under
Fig. 4. Application of nanoparticles augments the activity of photosynthetic machinery during stress.
4
Plant Nano Biology 4 (2023) 100035
diverse abiotic stress conditions are photosystem II (PS II), Rubisco, and
ATP (Ali et al., 2021; Misra et al., 1990; Misra et al., 2011; Munns and
Tester, 2008). Increased salt concentrations cause salt/ion build-up in
plant cells, reducing the plant's capacity to photosynthesize. Under
stress, these salts concentrate in the apoplast, generating excessive
dehydration and negative effects on chloroplast levels, affecting the
photosynthetic system (Mahmoud et al., 2020; Singh et al., 2021;
Tripathi et al., 2015).
SiO2 NPs can stimulate the photosynthetic system, enhancing chlor­
ophyll synthesis (Tripathi et al., 2015). When TiO2 -NPs were applied to
salt-stressed garlic (Allium sativum) plants, they increased photo­
synthetic pigment content and activity relative to control plants (Bharti
et al., 2018). TiO2 effectively absorbs light and hence transmits energy to
electrons. Furthermore, it boosts RuBisCo enzyme activity (the heart of
the dark cycle or kelvin cycle), which leads to increased carbon ab­
sorption (Poddar et al., 2020; Rizwan et al., 2019; Ze et al., 2011). In
stressed plants, growth and biomass increased, which are markers of net
photosynthetic carbon loss (Bharti et al. 2018). MSNs exhibited no ne­
gative effects on seed germination or when administered to several plant
organs (Lupin, Wheat, Maize, and Arabidopsis). MSNs travel to the roots
via symplastic and apoplastic pathways, and then to the aerial sections of
the plant, including the stems and leaves, via xylem conducting tissues,
demonstrating the ability to enhance photosynthesis by interacting with
F. Dilnawaz, M.H. Kalaji and A.N. Misra
chloroplasts, making them a potential candidate for smart delivery sys­
tems (Sun et al., 2014). Even at very high concentrations (2 g/L), MSNs
do not cause stress and have been proposed as a potential choice for
smart delivery systems (Badawy et al., 2021).
Salt stress has inhibitory effects by forming osmotic barriers, which
impair the capacity of seeds to absorb water and induce toxicity in seed
embryos (Almodares et al., 2007). SiO2-NPs have been proven to im­
prove seed germination and greatly enhance chlorophyll content under
NaCl stress (Siddiqui et al. 2014). In arid and semi-arid regions soil
dryness and salinity are key stress factors affecting agricultural yield
(Misra et al., 2002; Misra et al., 2011; Singh et al., 2019). However, NPs
have shown potential in promoting the growth and production of irri­
gated crops such as rice in these areas (Anjum et al., 2016). Soaking rice
grains in selenium (Se)-NPs and silicon (Si)-NPs resulted in the greatest
relative water content and dry matter, whilst foliar sprays of Se-NPs
during the booting stage resulted in the highest leaf area index when
compared to other treatments (Anjum et al., 2016). The use of Si-NPs
and Se-NPs as nano-nutrients enhanced ion homeostasis in plant cells,
assisting in mitigating the negative effects of salinity and increasing
grain production in rice crops (Badawy et al., 2021).
3.2. Drought stress
Drought is a recurrent abiotic stress that affects crop growth and
development, resulting in lower yield and productivity (Misra et al.,
2006; Misra et al., 2011). Soil dryness is produced by an increase in air
temperature, an increase in ion concentration in the root development
zone, and a dry environment or fast water drainage in arid and semi-
arid locations. (Yadav et al., 2020). Soil water deficiency, often known
as drought stress, is one of the most common abiotic stressors that re­
strict plant photosynthesis and agricultural yield globally through a
variety of biochemical, physiological, and molecular processes (Misra,
2018b; Anjum et al., 2016; Misra et al., 2014; Misra, 1991). To combat
the negative effects of climate change, a variety of strategies have been
implemented, ranging from irrigation management to genetic en­
gineering of crop plants via technological intervention (Liu et al., 2021;
Misra et al., 2006; Misra et al., 2011; Mushtaq et al., 2018; Othmani
et al., 2021; Wu et al. 2017b). Recent advances in nanobiotechnology
hold great potential for enhancing agricultural productivity by im­
proving plant tolerance mechanisms (Liu et al., 2021; Shang et al.,
2019).
SiO2-NPs, as demonstrated in the salt stress section, also show
promise in alleviating drought stress in banana plants (Mahmoud et al.,
2020). By controlling ion balance and ROS metabolism in stressed
plants, SiO2-NPs aid to decrease cellular damage and promote photo­
synthesis (Othmani et al., 2021). Nanoceria, a negatively charged poly
(acrylic acid) nanomaterial, may be sequestered to chloroplasts and
used to improve drought-induced photosynthetic inhibition (Wu et al.,
2017b). Abiotic stress plants treated with nanoceria demonstrated im­
proved quantum yield, net carbon assimilation (Pn), and Rubisco ac­
tivity (Wu et al. 2017b). In water-stressed sesame, studies employing
the magneto-priming approach for rapid incorporation of Si-O2 NPs
into seeds increased seedling establishment, crop establishment, and
crop yield (Janalizadeh et al., 2021). Si-NPs have also been shown to
improve cucumber growth and yield under water deficit and salinity
stress conditions by increasing mineral translocation, regulating the
Na/K ratio, increasing silicon translocation to the leaf, which in turn,
regulates osmotic balance by maintaining ion homeostasis, controlling
stomatal gas exchange and water flow in the soil-plant-atmosphere
system, and improving photosynthesis in stressed plants (Alsaeedi et al.,
2019).
3.3. Heat stress
Heat stress occurs when a plant is subjected to high temperatures for
a brief period of time, ranging from a few minutes to a few hours, and
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Plant Nano Biology 4 (2023) 100035
the air temperature rises over the optimal temperature for plant growth
or development (Geange et al., 2021). ROS, especially hydrogen per­
oxide (H2O2), are expected to play an important signaling function in
the body's response to stress (Mittler, 2017). Controlling ROS levels, on
the other hand, can be critical for maintaining adequate signaling and
minimizing heat stress damage. Heat stress may affect plants at any
stage of development, although it has the greatest impact on early es­
tablishment, flowering, and gametogenesis, as well as floral meristem
growth (Jagadish, 2020).
In a study, Se-NPs were used to treat high-temperature-exposed
sorghum plants, and it was discovered that they were effective at re­
ducing side effects such as membrane damage, reduced pollen germi­
nation, and decreased crop yields by activating the antioxidant defense
mechanism (Djanaguiraman et al., 2018). Another study looked at the
effects of Ag-NPs on plant growth and development under heat stress.
Heat stress lowered plant agronomic properties. AgNPs (50 and 75 mg/
l) application protected plants from heat stress by increasing plant
development parameters compared to the control (Iqbal et al., 2017).
Zinc (Zn)-NPs and iron (Fe)-NPs were utilized in a field investigation to
assess their effectiveness in wheat plants. In the Gimmeza7 cultivar,
NPs exhibited a rising influence on yield amount (the greatest mean
was obtained at 10 ppm Zn-NPs and 0.25 ppm Fe-NPs.). The beneficial
effect of such nanoparticles was followed by improved antioxidant
enzyme activity (glutathione S transferase, superoxide dismutase, per­
oxidase, and catalase), the formation of new bands in certain isozymes,
and a decrease in malondialdehyde, a lipid peroxidation result. These
biocompatible nanoparticles were extremely successful in encouraging
wheat plants to survive heat stress (Hassan et al., 2020).
3.4. Cold stress
Cold stress reduces plant water and food absorption, resulting in cell
starvation. Plants produce a variety of proteins in order to endure cold
conditions. They have developed advanced means of protecting them­
selves from cold-related injuries. Plants increase their resistance to cold
stress during this process by synthesizing a number of protective
compounds (e.g., soluble sugars, proline) and proteins such as late
embryogenesis abundant (LEA) for freezing tolerance (Kaplan et al.,
2007; Ruelland et al., 2009). To fight cold stress, nanoparticle (NP)-
mediated techniques have been employed. In one investigation, TiO2
-NPs were administered to sensitive and tolerant chickpea plant geno­
types. Various TiO2 -NP concentrations inhibited membrane damage
indices such as electrolyte leakage index (ELI) and malondialdehyde
(MDA) (Kazemi Shahandashti et al., 2013). Another study looked at the
influence of different TiO2- NP concentrations on membrane damage
variables as ELI and MDA in sensitive (ILC 533) and tolerant (Sel
11439) chickpea plant genotypes. The concentration of NPs in the va­
cuole and chloroplast demonstrated that NPs were taken up by seed­
lings. Under thermal treatments, the sensitive genotype was shown to
be more vulnerable to NPs than the tolerant genotype, and TiO2 levels
were greater under cold stress than at optimal temperatures. NPs
treatments improved physiological markers during thermal treatments.
TiO2 -NP treatments (especially 5 ppm) lowered ELI concentration
during heat treatments, but cold stress treatments at 0 ppm TiO2 en­
hanced ELI content in both genotypes. The study discovered that using
TiO2- NP enhanced genotype redox status after heat treatments
(Mohammadi et al., 2013). These findings imply that NPs in general,
and TiO2 -NPs in particular, might be used to improve cold tolerance in
crops (Mohammadi et al., 2013). In another study, TiO2-NPs were ap­
plied to chickpea plants, which resulted in enhanced expression of
Rubisco and chlorophyll-binding protein genes, decreased H2O2 con­
centration, and increased phosphoenolpyruvate carboxylase activity.
Plants treated with TiO2-NPs adapted to survival or recovery by coping
with the stress of temperature decrease damage and changed metabo­
lism for plant development (Hasanpour et al., 2015).
F. Dilnawaz, M.H. Kalaji and A.N. Misra
3.5. Heavy metal stress
Anthropogenic practices such as mining, irrigation, modern agri­
cultural methods, fertilizer application, sewage disposal, and in­
dustrialization have disrupted the environment by increasing the ac­
cumulation of heavy metals (HM) in soil, water, and air (Jha et al.,
2017; Nazir et al., 2022; Sharma et al., 2018; Singh et al., 2019). Food
crops absorb HMs from contaminated soil, which then accumulate in
edible plant portions and enter the food chain, possibly threatening
human health. As a result, the continuous accumulation of HMs harms
the soil's health and texture. (Jha et al., 2017; Riyazuddin et al., 2021;
Sharma et al., 2018; Singh et al., 2019). Arsenic (As), lead (Pb), cad­
mium (Cd), mercury (Hg), chromium (Cr), aluminium (Al), and ber­
yllium (Be) in polluted soils induce morphological deformities, meta­
bolic diseases, and substantially impair plant and agricultural
productivity (Amari et al., 2017; Sharma et al., 2012). Nanotechnology-
enabled particle formulations offer enormous promise for enhancing
abiotic stress control techniques. Interactions between NPs and metal
stress have an impact on cellular, molecular, and whole-plant physio­
logical processes. Exogenous treatment of CaO-NPs dramatically lowers
the effect of Al toxicity in barley by down-regulating As transporter
genes, allowing for improved calcium absorption, ROS scavenging, and
lowering As uptake and transit from roots to shoots (Nazir et al., 2022).
Biogenic Cu-NPs synthesized utilizing the bacteria Shigella flexneri
SNT22 reduced Cd translocation from soil to plants, enhanced biomass,
nutrient uptake, lowered Cd translocation, and Na+ concentration in
wheat plants (Noman et al., 2020). Foliar application of ZnO-NPs to
wheat plants cultivated in Cd-contaminated soil during a drought en­
hanced leaf chlorophyll content and lowered ROS activity (Adrees
et al., 2021). Silica has an important function in decreasing heavy metal
toxicity (Cui et al., 2017). Silica-NPs were used to assess growth, pho­
tosynthesis, oxidative stress, antioxidant defense system, and mineral
element regulation in Pisum sativum plants growing under Cr (VI)
stress. Application of Si-NPs to plants growing in Cr(VI) contaminated
soils as a nanofertilizer or foliar spray could reduce Cr accumulation in
roots and shoots and increase ROS scavenging mechanisms by up-reg­
ulating the antioxidant defense system, thereby improving plant
growth, metabolism, stress-regulating genes, and photosynthesis, ulti­
mately increasing plant biomass (Tripathi et al., 2017). Similarly, in a
Pb-contaminated environment, foliar treatment of Si-NPs reduced Pb
accumulation and reduced ROS activity in coriander plants (Fatemi
et al., 2021). Mercury (Hg) contamination is also a worldwide issue.
Mercury is absorbed by plants through their leaf surfaces as well as by
their roots from water and soil (Bishop et al., 1998). To minimize the
growth restriction caused by Hg in soybean plants, nano-SiO2 was
added to mercury-contaminated soil (Li et al., 2020a, 2020b). Alumi­
nium (Al) toxicity is the principal constraint on crop yield in acidic soil.
The majority of Al is bound in non-phytotoxic forms (Kochian et al.,
2015). Si-NPs have been found to be highly effective in reducing Al
toxicity (Hong et al., 2005a). Si-NPs have been shown to alleviate Al
toxicity in maize plants grown in acidic soil. These enhancements in­
clude increased accumulation of photosynthetic pigments such as
chlorophylls, increased activity of the Rubisco enzyme, decreased levels
of photorespiratory enzymes and NADPH oxidase, and activation of
antioxidant defense mechanisms (de Sousa et al., 2019). These findings
imply that Si-NPs can help plants deal with Al toxicity and increase
their overall development and health in harsh soil conditions.
3.6. Submergence stress
Submergence is a type of abiotic stress that causes hypoxic condi­
tions owing to waterlogging. Plant roots are the first and deadliest
victims of floods. The accumulation of ethylene in plants as a result of
hypoxia impedes root development and permeability. In one in­
vestigation, Ag-NPs were utilized to regulate submergence. Specific
amounts of Ag-NPs (0, 40, 80, or 120 ppm) were applied to saffron
6
Plant Nano Biology 4 (2023) 100035
corms before to planting, either under flooding stress or in non-flooding
circumstances.
After 10 days of flooding stress, root number, root length, root fresh
and dry weight, and leaf fresh and dry weight all decreased. Saffron corm
soaked with Ag-NPs (40 or 80 ppm) showed enhanced root number and
dry weight (Rezvani et al., 2012). Submergence induces hypoxia, which
results in morphological alterations and cellular acclimation responses.
Silver ion-mediated reactions appear to alter the ethylene, polyamines,
and calcium-mediated pathways, which are important for regulating
morphogenesis, particularly root induction (Licausi et al., 2011; Mishra
et al., 2008; Taylor and Whitelaw, 2001). Silver ions are utilized to replace
copper ions in receptor proteins, and they inhibit ethylene activity by
preventing it from attaching to its receptors in plant cells. During stressful
settings, both internal and external cues closely restrict ethylene bio­
synthesis. Except for wetland plants like rice, ethylene build-up as a result
of waterlogging may impede root growth and survival. Waterlogging-in­
duced ethylene build-up in plants may impair root development and
survival (Visser and Pierik, 2007, Wang et al., 2002). As a result, by
controlling ethylene production or activity, growth, and development
under stressful conditions such as floods may be regulated.
A proteomic method was utilized to investigate putative interactions
between various sizes of Al2O3-NPs and soybean under flooding stress
(Mustafa and Komatsu, 2016). The ascorbate/glutathione system con­
trolled scavenging activity in soybean plants exposed to Al2O3-NPs with
diameters ranging from 30 to 60 nm, which expanded the length of the
roots, including the hypocotyl, and suppressed the production of gly­
colysis-related proteins (Hashimoto et al., 2020).
3.7. UV-B stress
There is rising global concern about an upsurge in ultraviolet radiation,
particularly UV-B, reaching the Earth's surface as a result of the ozone hole
caused by human activities in the polar areas (Hong et al., 2005a; Lei
et al., 2008). UV-B exposure can cause an increase in ROS, changes in
antioxidant enzymes, cell membrane, and DNA damage, a decrease in
photosynthetic rate, and eventually a decrease in crop output (Gohari
et al., 2020; Hong et al., 2005a; Lei et al., 2008; Sicard et al., 2011). Plants
may benefit from nanoparticles such as TiO2, Ag, and Si by improving
photosynthesis and antioxidant enzyme activity (Kataria et al., 2014;
Mousavi et al., 2022). Because of the redox reactivity of Ce 3+/Ce 4+ in the
NPs, Cerium oxide (CeO2)-NPs can also offer UV-B defense to photo­
synthetic organisms by efficiently absorbing damaging UV radiation and
lowering oxidative stress (Gohari et al., 2020; Sicard et al., 2011;
Wojcieszek et al., 2019). A foliar spray of CeO2 on plants has been shown
to elevate membrane lipid peroxidation and leakage, cell wall and chlor­
oplast structure, chlorophyll production, and antioxidant enzyme activity
and ROS scavenging activities (Cao et al., 2018; Jahani et al., 2019;
Jurkow et al., 2020). In addition, Ag-NPs have been shown to dramatically
prevent the negative effects of UV-B on garden thyme by positively in­
creasing root development, photosynthetic pigment content, and essential
oil output (Azadi et al., 2021).
4. Use of nanotechnology in smart agriculture
In smart agricultural systems, remote sensing methods are being
utilized to monitor plant function, stress impacts, and photosynthesis by
monitoring CO2 assimilation using chlorophyll fluorescence or gas
analyzers (Misra et al., 2012a, Pérez-de-Luque, 2017). Biosensors based
on near-infrared (nIR) fluorescent single-walled carbon nanotubes
(SWCNTs) have been created to detect the essential signaling chemical
H2O2, which is related with the initiation of plant stress caused by UV-
B, photoinhibition/high light, and pathogen-induced biotic stress
(Amari et al., 2017; Li et al., 2018a, 2018b; Nazir et al., 2022;
Riyazuddin et al., 2021; Sharma et al. 2012). The optical nano-sensor
can detect early signals of stress, helping researchers to gain a deeper
understanding of plant stress communication (Wu et al., 2020). Salt
F. Dilnawaz, M.H. Kalaji and A.N. Misra
stress or salinity is the second most prevalent stress on Earth, influen­
cing global production, after water stress (Biswal et al., 2002; Misra
et al., 1999; Stefanov et al., 2019). Salinity also produces ROS and re­
duces photosynthesis in plants (Misra et al., 1990; Stefanov et al.,
2019). Nanoceria applied to A. thaliana leaves increased resistance to
ROS-induced damage and controlled ion balance in salt-stressed plants,
enhancing photosynthetic efficiency (Wu et al., 2018; Wu et al., 2017b).
Currently, fluorescent carbon dots (CDs) are potential replacements for
regular QDs. CDs are low in phytotoxicity and have little influence on
physiological metabolism. They make it possible for plants to absorb more
light energy. The photosynthesis of chloroplasts can be increased by uti­
lizing CDs with light-harvesting properties. CDs absorb UV light quite well,
whereas chloroplasts do not. The dispersion of CDs promotes the absorp­
tion of blue and red light by chloroplasts, hence boosting photosynthetic
activity in leaves by improving light capture and electron transfer effi­
ciencies in photosystem II (PS II) (Li et al. 2018a, 2018b). Using QDs,
optical glucose signaling in wild-type plants is also conceivable in vivo.
QDs were used to conduct optical glucose sensing. The probe was created
by combining thioglycolic acid-capped QDs that act as an internal glucose
reference with boronic acid (BA)-conjugated QDs that extinguish fluores­
cence in response to glucose. QD fluorescence response is active in the
visible window when the photosynthetic tissue background is minimal.
Confocal microscopy was utilized to study a standoff Raspberry Pi camera
system that can report glucose from a single alga (Chara zeylanica) and
plant tissues (A. thaliana). This technique allows for in vivo glucose
monitoring in photosynthetic organisms (in the presence of plant cell wall
porosity) as well as selective monitoring in situ (Li et al., 2018a, 2018b).
5. Conclusion
Plants integrate cellular, physiological, and molecular responses to
tolerate single or multiple stress factors that occur in nature and affect
plant/crop yield. These stress factors disrupt cellular processes such as
osmoregulation, ion homeostasis, ROS, hormone metabolism, and stress
signaling. This, in turn, affects plant organs and the entire plant and
canopy as a whole.
The application of various NPs, either supplemented through seed
priming, soil, or foliar application, has been shown to ameliorate stress and
alleviate stress-induced adverse effects at the chloroplast, leaf, and various
stages of crop production. As a result, NPs hold promise and potential to
increase plant/crop productivity in the future. To supply the demand for
food, fuel, and fodder in a world where marginal and unfavourable soil and
climate conditions exist, NPs could play a significant role.
The recent use of nanotechnology in smart agriculture is also a be­
ginning for the use of nano-sensors. These sensors can help monitor
plant health and stress levels, allowing farmers to make informed de­
cisions about when and how to intervene to optimize crop yields. This
innovative approach to agriculture, combining nanotechnology and
advanced sensor systems, is expected to contribute to more sustainable
and efficient farming practices in the future.
CRediT authorship contribution statement
FD; Conceptualization, original draft preparation, MHK; con­
ceptualization, review, and editing, ANM; conceptualization, review,
and editing.
Data availability
No data was used for the research described in the article.
Declaration of Competing Interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influ­
ence the work reported in this paper.
7
Plant Nano Biology 4 (2023) 100035
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