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Seed Priming with Nanoparticles: An Emerging Technique for Improving Plant

Growth, Development, and Abiotic Stress Tolerance

Article in Journal of Soil Science and Plant Nutrition · September 2022

DOI: 10.1007/s42729-022-01007-3

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Journal of Soil Science and Plant Nutrition
https://doi.org/10.1007/s42729-022-01007-3

REVIEW

Seed Priming with Nanoparticles: An Emerging Technique

for Improving Plant Growth, Development, and Abiotic Stress
Tolerance
Mohammad Saidur Rhaman1 · Shaila Shermin Tania1 · Shahin Imran2 · Farjana Rauf1 · Mohammad Golam Kibria3 ·
Wenxiu Ye4 · Mirza Hasanuzzaman5 · Yoshiyuki Murata6

Received: 6 April 2022 / Accepted: 14 September 2022

© The Author(s) under exclusive licence to Sociedad Chilena de la Ciencia del Suelo 2022

Abstract
Crop production is continuously threatened by various environmental stresses. Modern and efficient tactics are required to boost
agricultural output beyond facing stresses. Recently, many studies reported that nanotechnology can enhance plant growth and devel-
opment under stress and non-stress conditions, contributing to agricultural sustainability. Seed priming with nanoparticles (NPs) is
a modern and effective approach that increases germination, plant growth progression, productivity, and postharvest physiology of
plants by regulating morphological and physio-biochemical characteristics. Different plant studies showed that seed priming with
NPs enhances germination, improves nutrient uptake, regulates antioxidant defense, and reduces oxidative damage under stress and
non-stress conditions. Recently, several reviews on the regulatory role of nanotechnology on plant growth progressions have been
published. However, the regulatory roles of nano-seed priming have not been reviewed adequately and systematically. The present
review assembles the current literature on nano-priming and discusses the potentiality of nano-priming in the growth progression
of plants under stress and normal conditions and future directions for sustainable agriculture.

Keywords Abiotic stress · Cross tolerance · Nano-priming · Phytohormones · Seed germination · Seedling vigor · ROS

1 Introduction

Poor seed germination, seedling growth, and improper seed-

ling stand establishment are the major problems to success-
ful crop production (Finch-Savage and Bassel 2016). Last
* Mohammad Saidur Rhaman few years, abiotic stressors such as salt, drought, high heat,
saidursst@bau.edu.bd and heavy metalloids have reduced crop productivity (Rha-
* Mirza Hasanuzzaman man et al. 2022). To overcome the problems related to seed
mhzsauag@yahoo.com germination, growth progresses of seedlings, and the nega-
1 tive consequences of abiotic stresses on crop production,
Department of Seed Science and Technology, Bangladesh
Agricultural University, Mymensingh 2202, Bangladesh plant researchers are continuously focusing on developing
2 diverse approaches to enhance crop production. Among the
Department of Agronomy, Khulna Agricultural University,
Khulna‑9100, Bangladesh different approaches, seed priming is a successful method for
3 improving seed germination, growth progressions of seed-
Department of Soil Science, Bangladesh Agricultural
University, Mymensingh 2202, Bangladesh lings, and crop productivity in normal and abiotic stress set-
4 tings (Farooq et al. 2019; Waqas et al. 2019; Rhaman et al.
Institute of Advanced Agricultural Sciences, Peking
University, Weifang 261000, Shandong, China 2021a).
5 Seed priming refers to an economical and most effec-
Department of Agronomy, Faculty of Agriculture,
Sher-e-Bangla Agricultural University, Dhaka 1207, tive physio-biochemical technique of pre-treating of seeds
Bangladesh for a certain time period which enhances the seeds to ger-
6
Department of Bio‑Functional Chemistry, Okayama minate efficiently (Rhaman et al. 2021b; Tania et al. 2020;
University, Okayama, Japan Farooq et al. 2019; Waqas et al. 2019). Hydro-priming,

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 Journal of Soil Science and Plant Nutrition

halo-priming, osmo-priming, hormone-priming, matrix- fresh and dry weight, RWC, proline (Pro) content, ­K+ ion,
priming, bio-priming, magneto-priming, priming with and antioxidant enzymes activities (Khan et al. 2020). In
gamma radiation, UV irradiation-priming, smoke-priming, addition, iron (III) oxide-treated sorghum (Sorghum bicolor)
priming with nanoparticles (NPs), and X-Ray priming are showed higher plant growth by increasing photosynthetic
some of the seed priming agents/procedures that have been attributes such as Chl content, photosynthetic rate, and
developed and are extensively used to overcome the germi- RWC and decreasing lipid peroxidation under saline condi-
nation loss, improving seedling growth and stand establish- tion (Maswada and Djanaguiraman 2018). Likewise, maize
ment, and alleviating abiotic stresses (Paparella et al. 2015; (Zea mays) seeds primed with nano-Cu retained relative
Stanley et al. 2016; Sher et al. 2019; Rhaman et al. 2020a). leaf water status, increased Chl and anthocyanin contents by
Literatures describe that seed priming improves leaf area reducing ROS production under drought stress (Van Nguyen
index, relative water content (RWC), photosynthetic activ- et al. 2022). Silicon priming of marigold (Calendula offici-
ity, net assimilation rate, chlorophyll (Chl) content, phenolic nalis L.) seeds enhanced total flavonoid content and anti-
accumulation, nutrient uptake, and antioxidant enzyme oxidant activities under drought stress (Rahimi et al. 2020).
activities (Bajwa et al. 2018; Iqbal et al. 2018; Tania et al. Abbasi Khalaki et al. (2019) described that drought-stressed
2021). sheep fescue (Festuca ovina) seeds primed with AgNPs had
Among the different priming agents, recently, seed prim- maximum percentage of germination and seedlings growth.
ing with NPs have gained attention in research to enhance According to the available research, NPs have a substan-
growth progress and development of plants (Wang et al. tial influence on how plants react to stress, mostly through
2020; do Espirito Santo Pereira 2021; Kandhol et al. 2022). modulating biochemical and physiological pathways and
The NPs are commonly defined as a small object that is helping to maintain stress tolerance, although NPs have
1–100 nm in diameter, and they can exist as gaseous, liq- been shown to perform a regulatory part in the growth pro-
uid, or solid media state (Buzea et al. 2007; ASTM 2012). gressions of plants in a number of studies. The processes
The NPs have recently become more commonly employed underlying NPs priming-mediated growth progressions of
in commercial products and industrial applications, includ- plants and abiotic stress tolerance are not still elucidated.
ing biosensors, antimicrobial activity computer transistors, Therefore, the present review summarizes the recent reports
electrometers, and wireless electronic logic (Yan and Chen of NPs priming in plant developmental process under nor-
2018; Rico et al. 2014). In the agriculture sector, the NPs are mal condition and abiotic stresses. Besides, in order to gain
commonly used as nano-formulated pesticides, herbicides, in depth-knowledge on seed nano-priming, the purpose of
fungicides, fertilizers, and nano-biosensors for crop produc- this review is to provide an update concerning the potential
tion and protection (Mittal et al. 2020). The NPs application applications of seed nano-priming techniques in agriculture.
in the agricultural field has beneficial and harmful effects This review also discusses the potential mechanisms of NPs
on plants. For instance, NPs can promote phytotoxicity and priming in the alleviation of abiotic stress, as well as future
negatively regulate seed germination and growth. Moreover, directions.
the exclusive properties of NPs can fasten the seed germina-
tion and enhance crop yield (Khot et al. 2012). Therefore,
obtaining appropriate data on NPs properties and application 2 Commonly Used NPs in Seed Priming
procedures for successful crop production are crucial.
The NPs seed priming considered as an effective tactic It is well known that nano-priming is a modern tool of seed
to promote plant growth and development under stress and treatment that uses nano-materials, mainly NPs as priming
non-stress conditions (Jaberzadeh et al. 2013; Mishra et al. agents (Kasote et al. 2019; Shukla et al. 2019). The NPs
2014; Shah et al. 2021). The NPs speed up the activities priming has shown to be an efficient way among the vari-
of antioxidant enzymes, which reduce stress in plants by ous seed priming methods due to its small size and unique
diminishing the negative consequences of reactive oxygen physio-chemical features (Abbasi Khalaki et al. 2021; Acha-
species (ROS), resulting in improved growth and produc- rya et al. 2020; Pérez-de-Luque 2017). It is well known that
tion. Recently, reports showed that nano-priming acceler- the success of priming varies on the species of plants and the
ates morpho-physiological properties of plants under normal characteristics of treating agents. Recently, different NPs like
and stressful situations (Sarraf et al. 2022). For instance, metal-based, carbon-based, and plant extract-based are used
chili (Capsicum annum L.) seeds priming with metal oxide in nano-priming on the basis of both plants and NPs char-
NPs (zinc, titanium, and silver) improved seedling develop- acteristics. Nano-materials such as titanium dioxide ­(TiO2),
ment processes (Dileep Kumar et al. 2020). Under stress silicon dioxide ­(SiO2), Cu/CuO/Cu(OH)2, iron/iron oxides
conditions, seeds of pearl millet pretreated with silver (Ag) (Fe/FeOx), zinc/zinc oxide (Zn/ZnO), silver (Ag), gold (Au),
NPs enhanced salinity tolerance and increased plant height, biochar NPs, lignin NPs, and cerium oxide ­(CeO2) can be

13
Journal of Soil Science and Plant Nutrition

used in seed priming (Gottschalk et al. 2009; Pradas del Real activation (Siddaiah et al. 2018; Falsini et al. 2019). Chili
et al. 2016; Kah et al. 2019). seeds treating with metal oxide NPs (zinc, titanium, and
silver) improved germination and seedling development
(Dileep Kumar et al. 2020). In addition, wheat and pigeon
3 NPs Seed Priming in Plant Growth pea seed priming with Fe-NPs enhanced speed of germi-
and Development nation, root/shoot length, seedling dry, and fresh weight
(Fig. 1; Alam et al. 2015). Similarly, F ­ e2O 3 priming of
3.1 Role in Germination and Vigor chickpea seeds increased germination, shoot–root length,
and fresh and dry biomass (Pawar et al. 2019). Likewise,
Germination is the most critical and fundamental stages seeds of sorghum primed with iron oxide increased ger-
in seedling establishment and successful crop production mination and seedling growth (Maswada and Djanaguira-
(Savassa et al. 2018). Better germination of seed is consid- man 2018). In addition, rice seeds priming with zinc NPs
ered as a key prerequisite for proper seedling growth, leaf promoted germination, vigor index of seeds, and growth
expansion, nutrient uptake, and other physiological pro- of seedlings (Itroutwar et al. 2019). According to reports,
cesses including adequate biomass production (Mahakham primed onion seeds with biogenic silver and gold nano-
et al. 2016; Chandrasekaran et al. 2020). Besides, slow particles increased seedling emergence, leaves numbers,
germination is vulnerable to stresses and results in stunt plant biomass, and productivity (Acharya et al. 2019).
and erratic seedling growth which consequently lessens Azimi et al. (2014) described that nano-silica priming of
crop production. Seed priming with NPs is considered as tall wheatgrass enhanced germination rate of seed, height
an efficient technique, which affects not only seed germi- of radicle, and plant dry biomass. Bean seed priming with
nation and vigor but also the whole life cycle of a plant. CuO NPs in low concentration enhanced better germina-
Abbasi Khalaki et al. (2021) reported that NPs function tion and biomass of plants (Singh et al. 2017; Duran et al.
as stimulants and improve seed metabolic rate and seedling 2017). In addition, onion seeds priming with ZnO, Ag,
characteristics. Different NPs such as metallic, biogenic- CuO, and ­TiO2 NPs enhanced the germination, shoot–root
metallic, and polymeric have also been demonstrated length, and seed vigor index (Anandaraj and Natarajan
to have promising in seed germination through enzyme 2017). Moreover, sugar beet, and maize seeds priming

Fig. 1  Role of NPs seed priming in morpho-physiological and post-harvest physiology of plants

13
 Journal of Soil Science and Plant Nutrition

with ­TiO2 improved germination, shoot–root length, bio- and shows that α-amylase induction is dependent on GA
mass, and seedling vigor index (Shah et al. 2021). NPs activity. Although it is unknown what upstream GA sign-
priming may modulate the metabolism of seed, which aling elements are involved in α-amylase-mediated starch
leads to increased water uptake, starch hydrolysis rate, hydrolysis? The investigation of the interactions between
cell wall loosening, endosperm weakening, rapid embryo plant hormones and sugar signaling cues supported by nano-
growth, and rapid root-shoot development. Additionally, priming ought to be fascinating.
seed nano-priming may control many physio-biochemical
processes, such as altering the defense system’s activity 3.3 Role in Antioxidant Enzyme Systems
and upping antioxidant levels and enzyme activities, mak-
ing plants more vigorous as well as resilient to environ- Seed priming with NPs found as an emerging strategy for
mental challenges (Abdel Latef et al. 2017; Kasote et al. antioxidant enzyme regulation in plants (Table 1; Li et al.
2019). 2021). Antioxidant system in plants includes a number of
enzymatic and non-enzymatic antioxidant enzymes such as
3.2 Role in Nutrient Uptake catalase (CAT), peroxidase (POX), ascorbate-peroxidase
(APX), superoxide-dismutase (SOD), phenylalanine ammo-
Plant nutrients are the indispensable elements for plant nia lyase (PAL), glutathione, and glutathione reductase.
growth progressions not only under usual condition but It has been found that priming of maize seeds by sodium
also in stressful conditions. For the successful completion metasilicate elevated the SOD, CAT, and POX activities
of plant life cycle, plant uptakes nutrients through their under salt stressor (Abdel Latef and Tran 2016). Similarly,
roots and leaves. On the other hand, poor nutrient uptake ­TiO2 NPs primed maize seeds elevated the functions of
negatively affects root formation, seedling growth, flower, SOD, CAT, and PAL (Shah et al. 2021). Rice seed prim-
and fruit formation. (Srivastava and Malhotra 2017; López- ing with ZnO nano enhanced the SOD, and POD activi-
Valdez et al. 2018). Furthermore, proper nutrient delivery ties (Li et al. 2021). In addition, seed priming of Egyptian
to a specific crop for successful production is a difficult task Roselle (Hibiscus sabdariffa L.) with ­Al2O3 NPs promoted
by conventional nutrient management systems (Feregrino- the functions of SOD, CAT, POD, and APX (Abdel Latef
Pérez et al. 2018; Kyriacou and Rouphael 2018). Therefore, et al. 2020). The AuNPs and AgNPs priming of Lavender
recently a number of studies suggested that nano-priming (Lavandula angustifolia Mill.) promoted the functions of
technology played important role in proper nutrient manage- APX, POX, and SOD (Jadczak et al. 2020). Wheat seeds
ment. For instance, rice seeds priming with nanoscale zero- priming with Si NPs elevated the functions of SOD, POD,
valent iron (nZVI) promoted the accumulation of macro and and CAT under cadmium stress (Hussain et al. 2019). It is
micronutrients in grains (Guha et al. 2022). The iron pyrite generally known that the antioxidant system regulates ROS
­(FeS2) NPs treatment enhanced calcium, manganese, and production in response to environmental stressors. The anti-
zinc concentrations in the spinach leaves which resulted in oxidant system is crucial for redox management by remov-
high yield of spinach (Srivastava et al. 2014). The ZnO NPs ing ROS and preventing potential cellular molecule damage
reported to increase Zn contents in wheat, which resulted by maintaining ROS homeostasis (Dietz et al. 2016). The
in better growth of wheat (Elhaj Baddar and Unrine 2018). antioxidant enzymes determine how ROS help nano-primed
In addition, the maize, soybean, pigeon pea, and okra seeds seeds germinate in an indirect manner (Anand et al. 2019).
coated with ZnO NPs enhanced indole-3-acetic acid and For instance, it was discovered that increased SOD and CAT
increased the plants Zn requirement, which ensued in bet- activities reduced the activity of ­H2O2 radicals in tomato,
ter growing of plants (Adhikari et al. 2016). Different NPs cucumber, and pea nano-primed seeds (Barba-Espín et al.
priming may alter the metabolism of the seed, break down 2012; Bhardwaj et al. 2012; Anand et al. 2019). However,
the starch that has been stored, and stimulate the release research on the regulation of antioxidant enzymes via NPs
of growth regulators, which ultimately improves nutrient priming is very limited. Therefore, additional investiga-
uptake and fosters the growth and productivity of plants. For tions are required to uncover the effects of NPs priming on
instance, a quick rise in α-amylase activity brought on by antioxidant enzymes especially non-enzymatic antioxidant
nano-priming in germinating seeds may result in improved regulation in different plants under diverse conditions. In
germination rates, seedling vigor, and nutrient uptake (Man addition, ­H2O2 modifies the expression of multiple genes
et al. 2013; Mahakham et al. 2017). Furthermore, gibberellic involved in the germination process by carbonylating pro-
acid (GA) activity is necessary for this increase of α-amylase teins, initiating and changing kinase transduction pathways
synthesis. For instance, Mahakham et al. (2017) revealed (El-Maarouf-Bouteau et al. 2013). In light of this, we sur-
a connection between the lack of GA and the absence of mise that additional research is necessary to clarify the role
α-amylase biosynthesis. This demonstrates signaling cross- of antioxidant scavengers in initiating these transduction
talk between NPs, α-amylase, and GA in nano-primed seeds processes.

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Journal of Soil Science and Plant Nutrition

Table 1  The effects of nanoparticles seed priming on morpho-physiological responses of plants under normal condition
Plant species Nanoparticles Responses of plants References

Rice (Oryza sativa L) Biogenic silver, biogenic zinc, Promoted germination of seeds, Divya et al. (2019), Itroutwar et al.
iron sulfide, chitosan growth, and biomass. Increased (2019), Mahakham et al. (2017)
functions of antioxidant
enzymes
Sorghum (Sorghum bicolor) Iron oxide Enhanced germination, vigor, Maswada and Djanaguiraman
leaf water content, and plants (2018)
biomass
Water melon (Citrullus lanatus) Silver, iron, manganese, and Increased germination, vigor, Kasote et al. (2019; 2021), Acharya
turmeric nano-emulsion enzymatic and biochemical et al. (2020)
activity
Wheat (Triticum aestivum L.) ZnO, iron oxide, silicon, chitosan Increased biomass and yield, Rizwan et al. (2019), Hussain et al.
enhanced iron absorption and (2019), Sundaria et al. (2019)
deposition, reduced oxidative
stress
Chilli (Capsicum annuum) ZnO, ­TiO2, silver, manganese Increased germination, vigor, Dileep Kumar et al. (2020), Ye
oxide antimicrobial activity, and anti- et al. (2020a, b)
oxidant enzyme activities
Maize (Zea mays L.) Biogenic gold Increased germination, vigor, and Mahakham et al. (2016)
initial seedling growth
Onion (Allium cepa L.) Silver and gold Enhanced germination of seeds, Acharya et al. (2019)
emergence of growth, and incre-
ment of yield
Soybean (Glycine max) Cobalt oxide, molybdenum oxide Boost up germination and growth Chau et al. (2019)
and biomass of plants
Tomato (Solanum lycopersicum) Magnet Enhanced germination capacity, Anand et al. (2019)
increased transcript level of
several genes such as SOD1 and
SOD9
Barley (Hordeum vulgare) Iron/silicon Increased speed of germination, Najafi Disfani et al. (2016)
and lengths of shoot and root
Green gram (Vigna radiata) Zinc oxide Enhanced speed of germination, Lakshmi et al. (2017)
growth, and yield
Switch grass (Panicum virga- Titanium oxide Improved seed germination and Boykov et al. (2018)
taum) enhanced plant developmental
process
Niger (Guizotica abyssinica) TiO2 and ­SiO2 Low concentration of NPs Eskandarinasab et al. (2019)
elevated germination rate and
mean of daily germination

3.4 Role in ROS and Lipid Peroxidation Regulation et al. 2012a; Rui et al. 2015; Zhang et al. 2015). However,
ROS accumulation and signaling triggered the process of
Seeds generally receive NPs as an external agent (Anand seed dormancy breaking and stimulated seed germination
et al. 2019) and after being accumulated in the seed coat (Oracz and Karpinski 2016). This might occur through the
they induce ROS accumulation and activate a number of activation of GA synthesis (Dietz et al. 2016). Moreover,
downstream processes (Guha et al. 2018). ROS is produced increased ROS accumulation in nano-primed seeds strongly
as a by-product in plants during aerobic-metabolism, and interlinked with the abscisic acid (ABA) synthesis, which
is generated in chloroplasts, mitochondria, and peroxi- also triggered seed dormancy breaking and seed germina-
somes (Sandalio and Romero-Puertas 2015; Dietz et al. tion (Bailly and Kranner 2011; Bailly 2019). In addition,
2016; Huang et al. 2016). ROS irreversibly damage DNA nano-priming elevated ROS levels in plant cells that helped
but they are also well known as signaling molecules that let to break the bonding among the polysaccharides in the cell
plants grow normally and mitigate abiotic stresses (Miller wall of seed endosperm and facilitated quick and healthy
et al. 2010; Singh et al. 2019). In the NPs-mediated prim- seed germination (Chandrasekaran et al. 2020). Thus, it
ing, ROS production depended on the NPs characteristics, has been apparent from the literature that ROS acts as a
such as size, shape, concentration, and solubility (Dimkpa positive signaling component for releasing seed dormancy

13
 Journal of Soil Science and Plant Nutrition

by nano-priming. However, it should be noted that knowl- 2017). Moreover, many studies reported that NPs promoted
edge about phytohormones other than ABA/GA interacting germination, vigor, and quality of groundnut, green gram,
with NPs-mediated ROS influx in intercellular trafficking onion, lettuce, and peanut seeds (Zheng et al. 2005; Prasad
is mainly unexplored (Mahakham et al. 2017). Therefore, et al. 2012; Shyla and Natarajan 2016; Lakshmi et al. 2017;
more research needs to be done to understand how ROS Anandaraj and Natarajan 2017).
modify downstream protein targets, as well as how trans- It is well known that harvested crops need to be well
porters involved in intercellular communication interact with packaged during processing and storage to maintain fruit or
ROS and phytohormones in nano-primed seeds. In connec- seed quality. NPs have been widely used for better packaging
tion with this, a thorough genomic and proteomic analysis and storage of fruits and seeds. In this regard, recently, silica
employing ROS signaling mutants is required to ascertain NPs, chitosan NPs, and other inorganics are widely using as
the functions of particular ROS related enzymes taking part a storage materials (Duncan 2011). Likewise, NPs like ZnO
in the nano-priming-mediated crosstalk networks. and MgO enhanced the shelf life as well as the quality of
On the other hand, a number of studies showed NPs stored products. In addition, nowadays, NPs are using as bio-
priming reduces the ROS level under stress conditions and sensors (multiple chips) for product labeling during storage
enhances the crop stress tolerance. For instance, salt-stressed (Yadollahi et al. 2009). Moreover, several studies reported
rapeseed priming with cerium oxide NPs reduced the ROS that the chitosan NPs improved the post-harvest quality of
and MDA levels (Khan et al. 2021). Likewise, maize seed grapes, and banana, by reducing moisture content, total solu-
priming with copper NPs decreased the ROS level under ble solids, pH, and lowered skin discoloration (Lustriane
drought stress (Van Nguyen et al. 2022). In addition, Lathy- et al. 2018; Esyanti et al. 2019). In addition, silver coating
rus odoratus seeds priming with Si NPs reduced the ROS increased physicochemical and nutraceutical properties of
and MDA levels under salt stress (El-Serafy et al. 2021). loquat ( Eriobotrya japonica) in storage ( Ali et al. 2020).
Shah et al. (2021) depicted that ­TiO2 NPs of maize seeds
reduced oxidative damage and increased salt stress toler-
ance by positively regulating antioxidant enzymes. The 4 NPs Seed Priming in Abiotic Stress
lowering of ROS by NPs may result in a large rise in anti- Mitigation
oxidant enzyme levels. Because of the link between ROS
and antioxidant enzymes, nano-primed seeds have a higher The hostile environmental circumstances such as salt, ther-
concentration of antioxidant enzymes due to NPs-mediated mal, drought, floods, and heavy metals extremely affect crop
ROS mitigation. However, further exploration is required to growth development and production (Tahjib-Ul-Arif et al.
uncover the regulatory role of NPs priming in ROS and lipid 2021; Rhaman et al. 2022). The NPs technology brings new
peroxidation remediation in different plants. dimensions in crop production beyond facing the challenges
of abiotic stress (Kumari et al. 2022; Manzoor et al. 2022;
Nile et al. 2022).
3.5 Role in Postharvest Physiology Among the environmental stresses, salinity is a threat-
ening abiotic stress that hampers plant growth and con-
To ensure the high quality of agricultural products with sequently affects yield of crops (Isayenkov and Maathuis
longer shelf-life is an essential demand for crop growers. In 2019). Several studies showed that seeds priming with NPs
this regard, proper management of post-harvest technology increases the salt tolerance level and results in increased
performs a key function in flourishing the food industry to yield (Table 2). Seed priming with Mn NPs promoted root
provide a high-quality product with a longer shelf life. It is elongation, altered available macro-/micro-nutrients that
well known that post-harvest losses occur due to handling, include reallocation of manganese, sodium, and calcium
packaging, storage, and transpiration which reduce the shelf and increased salt tolerance of Capsicum annuum (Ye et al.
life and benefits of the products. Recently, a number of stud- 2020a, b). However, under salinity stress, seeds of maize and
ies described that seed priming with NPs emerged as a novel Paeonia suffruticosa priming with T ­ iO2 enhanced germina-
tactic to improve crop productivity and quality at every stage tion, vigor index, length of shoot and root, biomass of seed-
of plant including, germination, growth and development ling, RWC, total phenolic and Pro contents, and functions
and post-harvest physiology of plants (Shukla et al. 2019; of antioxidant enzymes and lessened concentration of ­Na+,
Acharya et al. 2020). Literature shows NPs priming con- membrane electrolyte leakage (MEL), and malondialdehyde
trols the post-harvest diseases in fruits and improves the (MDA) contents (Shah et al. 2021; Liu et al. 2021). Like-
quality of seeds (Sharma et al. 2019). Shyla and Natarajan wise, seeds of pearl millet pretreated with AgNPs elevated
(2016) reported that NPs like-ZnO, Ag, and T ­ iO2 improved plant height, fresh and dry weight, RWC, Pro content, ­K+
groundnut quality. Likewise, ZnO NPs enhanced param- ion, and antioxidant enzymes activities but at the same time,
eters affecting seed quality in green gram (Lakshmi et al. it decreased ­Na+ ion and N­ a+/K+ ratio and ameliorated salt

13
 Table 2  The effects of nanoparticles seed priming on morpho-physiological responses of plants under stress
Plant species Nanoparticles Stress Responses of plants References

Caucasian alder Multi-walled carbon nanotubes (MWCNTs) Drought Increased the germination rate and percentage Rahimi et al. (2016)
(Alnus subcordata) (GP), increased seed vigor index, increased
root and shoot growth
Cape Periwinkle (Catharanthus roseus) Chitosan Drought Boosted plant growth and defense mecha- Ali et al. (2021)
nism, enhanced Pro accumulation, preserved
membrane integrity, mitigated drought-
induced damage
Capsicum Manganese (Mn) Salinity Increased enlargement of roots and promoted Ye et al. (2020a, b)
(Capsicum annuum) salt tolerance
Journal of Soil Science and Plant Nutrition

Cotton Cerium oxide (­ CeO2) Salinity Improved root development, increased root An et al. (2020)
(Gossypium rboretum) biomass, decreased ROS accumulation, K, Ca
and Mg contents
Cucumber Nanoparticles of water treatment residuals Salinity Increased GP, radical length, and total biomass, Mahdy et al. (2020)
(Cucumis sativus) (nWTRs) decreased root radius
Rice Zinc oxide Cadmium Improved amylase activation, antioxidant Li et al. (2021)
(Oryza sativa) enzymes activities, modulated Zn concentra-
tion, and ZnO NPs uptake
Lentil Iron chelate Salinity Improved germination rate and GP, increased Nourafcan and Shahmoradi (2014)
(Lens culinaris) emergence time, plumule length, fresh and dry
biomass
Lupine Zinc oxide Salinity Enhanced photosynthetic pigments, adjusted Latef et al. (2017)
(Lupinus albus) osmoregulation, lowered MDA and Na con-
tents, and increased plant growth
Maize Titanium-di-oxide Salinity Increased seed germination, enhanced anti- Shah et al. (2021)
(Z. mays) oxidant enzymes activities, RWC, Pro, and
total phenolic contents, decreased sodium ion
­(Na+), electrolyte leakage (EL), and MDA,
improved plant growth
Maize Copper Drought Increased anthocyanin, Chl, carotenoid contents, Van Nguyen et al. (2022)
(Z. mays) leaf RWC. Decreased ROS accumulation,
increased plant biomass, total seed number
and grain yield
Marigold Silicon Drought Increased antioxidant activity, quercetin and Rahimi et al. (2020)
(Calendula officinalis L.) total flavonoid content, improved plant physi-
ological and metabolite indices
Milk thistle (Silybum marianum) Chitosan nanoparticle and pyridoxine Salinity Improved germination, Chl, Pro, and the func- Mosavikia et al. (2020)
tions of antioxidant enzymes
Mudan Titanium dioxide Salinity Accelerated seed germination, promoted the Liu et al. (2021)
(Paeonia suffruticosa) functions of SOD, POD, and CAT, elevated
Chl content, lateral root, and seedling dry
weight

13
 Table 2  (continued)
Plant species Nanoparticles Stress Responses of plants References

13
Pearl millet (Pennisetum glaucum) Silver Salinity Increased RWC, Pro content, curtailed oxidative Khan et al. (2020)
injury, enhanced antioxidant enzyme func-
tions, increased K­ +, decreased N
­ a+ and ­Na+/
K+ ratio
Rapeseeds Cerium oxide Salinity Improved the germination rate, water absorb- Khan et al. (2021)
(Brassica napus) ance, SOD and POD activities, α-amylase
activity, and total soluble sugar contents,
lowered ROS accumulation, maintained N ­ a+/
K+ ratio, increased crop biomass
Sheep fescue (Festuca ovina) Silver Drought Increased GP, enhanced seedling growth Abbasi Khalaki et al. (2019)
Sorghum Iron (III) oxide Salinity Increased Chl index, photosynthesis rate, and Maswada and Djanaguiraman (2018)
(Sorghum bicolor) RWC. Decreased peroxidation of lipids,
increased plant growth
Sunflower (Helianthus annuus L.) Greenly synthesized sulfur Manganese Restricted ROS overproduction and oxida- Ragab and Saad-Allah (2021)
tive stress, up-regulated antioxidant enzyme
activities
Wheat Sodium nitroprusside Drought Upregulated antioxidant enzymes activities, Hameed et al. (2021)
(Triticum aestivum) increased glycine betaine content, reduced
total sugars content, enhanced plant biomass,
and yield
Wheat Titanium-di-oxide and Sodium nitropruside Cadmium Increased GR, early seedling growth, and water Faraji and Sepehri (2018)
(T. aestivum) uptake capacity
Journal of Soil Science and Plant Nutrition
Journal of Soil Science and Plant Nutrition

stress (Khan et al. 2020). Sorghum seeds treated with iron nanozymes aid plants during stress by scavenging ROS
(III) oxide NPs under salt stress showed higher plant growth and enhancing enzymatic abilities (Palmqvist et al. 2017).
by increasing Chl content, photosynthesis rate, and RWC Additionally, bio-priming of seeds with rhizomicrobes that
and decreasing peroxidation of lipids (Maswada and Djana- promote plant growth, similar to nano-priming, is a crucial
guiraman 2018). Moreover, priming of rapeseed with cerium strategy for reducing abiotic stress as well as for sustain-
oxide increased germination, water absorbance, SOD, POD, able agriculture (Mahmood et al. 2016). Bio-nano-priming
α-amylase activities, total soluble sugar contents, and ­Na+/ is a convergent area of beneficial technology that combines
K+ ratio while lowered the ROS accumulation and enhanced the advantages of applying nano and bio-agents for enhanc-
the salinity tolerance (Khan et al. 2021). In addition, lentil ing crop growth (Pudake et al. 2019). For instance, bacterial
seeds with iron nano chelate (Nourafcan and Shahmoradi priming of wheat increased photosynthesis rate and plant
2014), cucumber seeds with nanoparticles of water treat- biomass under drought stress (Timmusk et al. 2014). Metal
ment residuals (Mahdy et al. 2020) and milk thistle with or metal oxide NPs may alter the profiles of the secondary
chitosan nanoparticle (Mosavikia et al. 2020) alleviated salt metabolite and protein synthesis carried out by plant growth-
stress by improving physiological characteristics of plants. promoting rhizomicrobes in a concentration-dependent way
Latef et al. (2017) described that ZnO NPs priming of lupine (Haris and Ahmad 2017) and improved the production of
seed promoted photosynthetic dyes, organic solutes, total plant hormones (IAA) (Dimkpa et al. 2012b) that may be
phenols, Zn, and APX, POD, SOD, and CAT activities and able to fine-tune the plant growth-promoting benefits in
decreased the MDA content and N ­ a+ under salinity stress treated crops (Timmusk et al. 2018).
condition. Similar to this, priming with nano-ZnO improved Metal toxicity is one of the detrimental abiotic stresses
salt tolerance in wheat by activating the antioxidant system that hampers plant growth progressions. Recently, several
to reduce the oxidative burst and increasing the effectiveness studies showed that NPs priming decreases the toxic met-
of photosynthetic electron transport and sucrose production als accumulations and mitigates the negative consequences
in stressed leaves (Wang et al. 2020). According to Baz in different crop production (Table 2). For instance, Mn-
et al. (2020) lettuce seed priming with water soluble carbon stressed sunflower (Helianthus annuus L.) seeds priming
nanoparticles (CNPs) increased seed germination and Chl with greenly synthesized sulfur stimulated antioxidant
contents under salt stress. enzyme activities and decreased the ROS and lipid peroxi-
Drought stress severely stunts plant growth and reduces dation (Ragab and Saad-Allah 2021). Under cadmium (Cd)
plant productivity (Osakabe et al. 2014). The NPs-mediated stress, ZnO NPs priming increased amylase activation, POD
priming exhibited significant effects on different plant growth and SOD activities, and seedling growth of fragrant rice (Li
performance to mitigate the drought stress (Table 2). Rahimi et al. 2021). In addition, wheat seeds priming with T ­ iO2
et al. (2016) described that seeds priming with multi-walled increased germination percentage, seedling growth, and
carbon nanotubes increased the rate of germination, vigor water uptake capacity under Cd stress (Faraji and Sepehri
index and root-shoot growth of alnus subcordata (Cau- 2018). Similarly, under Cd stress, S ­ iO2 NPs enhanced ger-
casian alder) under drought stress condition. In addition, mination rate and seedling growth of Moso bamboo (Phyl-
drought-stressed cape periwinkle (Catharanthus roseus L.) lostachys edulis) (Emamverdian et al. 2021). In maize, under
seeds primed with chitosan NPs showed enhanced Pro accu- cobalt stress seed priming with ZnO NPs reduced the ROS
mulation, membrane integrity, and plant growth (Ali et al. and MDA level and improved plant growth, biomass, and
2021). Moreover, maize seeds priming with copper NPs photosynthetic machinery (Salam et al. 2022). In general,
increased RWC, Chl, carotenoid and anthocyanin contents, it has been discovered that NPs regulate plant physiological
and decreased ROS accumulation during drought stress and biochemical parameters to reduce the detrimental effects
(Van Nguyen et al. 2022). Silicon NPs priming of mari- of HMs (Rajput et al. 2021). However, NPs priming can
gold (Calendula officinalis L.) seeds enhanced quercetin, be a promising technique for heavy metal stress mitigation.
total flavonoid content, and antioxidant activities under Therefore, this area needs to clarify further by investigating
drought stress condition (Rahimi et al. 2020). Likewise, the effects of NPs priming in various heavy metal-stressed
Abbasi Khalaki et al. (2019) found increased germination plants. Also, additional research is required to settle the ben-
percentage and seedling growth of sheep fescue (Festuca eficial effects of NPs priming under UV, nutrient deficiency,
ovina) plants under drought stress condition while seeds temperature, water-logging stresses.
were primed with AgNPs. Nanozymes, magnetic NPs of The above discussions clarify that seed priming with NPs
magnetite ­Fe3O4 and magnetite γ-Fe2O3 have been reported positively regulates ionic homeostasis, secondary metabo-
to be effective in plant growth progressions under drought lites, photosynthetic attributes, and antioxidant enzyme
stress. For example, γ-Fe2O3 NPs reduced H ­ 2O2 and lipid activities, and decreases oxidative stress to alleviate abiotic
peroxidation and enhanced growth of Brassica napus under stress in plants (Fig. 2).
drought stress (Palmqvist et al. 2017). According to report,

13
 Journal of Soil Science and Plant Nutrition

Fig. 2  Nano-priming-mediated stress tolerance mechanisms in plants. matal aperture but the role of nano-priming not been investigated.
Seed priming with NPs maintained ionic homeostasis, enhanced anti- Also, the role of NPs priming in the regulation of stress related gene
oxidant enzyme activities, reduced oxidative damage and enhanced expressions not fully elucidated
stress tolerance. The NPs inhibits stress-induced reduction of sto-

5 Constraints of Using NPs Seed Priming showed positive results, proper care must be taken during
application of NPs. The proper and safe use of NPs in seed
Seeds priming with NPs not only have beneficial effects priming is essential not only for agriculture but also for the
but also have some negative consequences on plant growth environment and many other industrial sectors (Scott et al.
progression and ecosystem. Plant species, NPs properties, 2018).Taking into consideration, the possible ecotoxicity of
and culture media potentially caused the divergent results NPs, logical, legal, and conscience agendas are essential for
(Yang et al. 2017). Inappropriate NPs size, high concentra- manufacturing of NPs, disposal of industrialized waste, and
tion, and high duration of priming can cause germination in agricultural appliances.
inhibition and reduction of plant development by altering
plant metabolism and cell structure (Dileep Kumar et al.
2020; Gross et al. 2020). The toxic effects of NPs priming 6 Future Research on NPs Priming
showed seed emergence deterioration, seedling numbers
reduction, root and stem growth prohibition, and delay- Seed priming with NPs has great advantages than foliar and
ing of flowering and ultimately reduced the yield (Hayes soil supplementation as treatments in seeds minimize the
et al. 2020; Pelegrino et al. 2020). Additionally, nanopar- exposition of NPs. Low concentrations of NPs have been
ticles like carbon nanotubes, metals and metal oxides, and found in studies to have a superficial effect on plant growth
fullerene showed diverse levels of toxicity on the ecosys- and development. To select an appropriate concentration
tem (Klaine et al. 2008; Palimi et al. 2015; Pachapur et al. of NPs for both effective and negligible toxic effects, it is
2016). Several reports described the negative consequences critical to clarify how the physio-chemical properties of NPs
of NPs on plant physiology of Arabidopsis (Wang et al. disturb seed priming. Also, the interaction effects of NPs and
2016), black mustard (Amooaghaie et al. 2015), sorghum plants need to elucidate how different NPs interact with the
(Lee et al. 2012), and maize (Zhang et al. 2015). Copper NPs plant development. Also, the residual effects of NPs in the
inhibited germination of mungbean and wheat seeds and environment need to clarify.
decreased the elongation of shoot–root length by entering Plant evolves different sophisticated mechanisms to coun-
into the cell membranes (Lee et al. 2008). The AgNPs toxic- teract the effects of stresses. Stomatal regulation is an indis-
ity have been found as a severe issue for the environment and pensable mechanism of plants under stress conditions to reg-
plant growth progressions. Although NPs in seed priming ulate gas exchanges and transpiration (Rhaman et al. 2020b;

13
Journal of Soil Science and Plant Nutrition

Islam et al. 2020; Ye et al. 2020a, b). Recently, NPs are using
in photosynthetic attribute, stomatal movement regulation.
Faizan et al. (2021) reported that Cu-induced stomatal aper- Author Contribution MSR and MH conceived the idea and prepared
the outline. SST, SI, FR, and MGK prepared the original draft; MSR,
ture reduction was improved by ZnO NPs application. Iron WY, MH, and YM reviewed and edited the manuscript. All authors
NPs-induced stomatal opening in Arabidopsis by activating have read and agreed to the published.
plasma membrane ­H+-ATPase and consequently enhanced
­CO2 uptake (Kim et al. 2015). Therefore, NPs application Declarations
under stress and non-stress condition improve crop growth
by regulating stomatal movements. However, still, there is Conflict of Interest The authors declare no competing interests.
no report available of NPs seed priming whether regulates
stomatal movement in plants.
Seed priming with NPs has been shown to begin or modify
a number of genes expressions (Chandrasekaran et al. 2020).
References
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Timmusk S, Abd El-Daim IA, Copolovici L, Tanilas T, Kännaste A, Publisher’s Note Springer Nature remains neutral with regard to
Behers L, Nevo E, Seisenbaeva G, Stenström E, Niinemets Ü jurisdictional claims in published maps and institutional affiliations.
(2014) Drought-tolerance of wheat improved by rhizosphere bac-
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