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Nano-Priming as Seed Priming Technology for Sustainable Agriculture

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 CHAPTER 2
Apple Academic Press

Nano-Priming as Seed Priming

Technology for Sustainable Agriculture
FATIH ÇIĞ,1, ROJIN ÖZEK,2 MURAT ERMAN,3 SIPAN SOYSAL,1
ÖZGE UÇAR,1 ZEKI ERDEN,1 MUSTAFA CERITOĞLU,1

Author Copy
ÇAĞDAŞ CAN TOPRAK,1 SÖNMEZ ÖZBEY,1 MUHAMMAD A. RAZA,1
JAVEED A. LONE,1 SAIFULLAH ABRO,1 MUHAMMAD ARSHAD,1 and
MEHMET EFE1
1Department of Field Crops, Faculty of Agriculture, Siirt University, Siirt,
Türkiye
2KırıkkaleUniversity, Kırıkkale Vocational School, Department of Organic
Agriculture, Türkiye
3Bursa Uludağ University, Faculty of Agriculture, Department of Field
Crops, Türkiye

ABSTRACT

Agriculture is threatened by climate change and by the depletion of

resources and biodiversity. Exploring new farming methods is needed to
increase crop production and ensure food quality and safety in a sustainable
way. Nanotechnology is an emerging trend that contributes to sustaining
agricultural production. Seed nano-priming helps improve seed germination,
seedling emergence, growth, and yield by resisting various plant stresses.
Nano-priming is a more effective method than any other seed preparation
method. Studies have shown several benefits of using seed nano-priming. By
regulating biochemical pathways and the balance between reactive oxygen
species and plant growth hormones, nano-priming helps increase resistance
against stresses and diseases with the limited use of pesticides and fertilizers.

Nanotechnology for Agriculture: A Potential Tool for Abiotic Stress Tolerance.

Ashwani Kumar & Ayman El Sabagh (Eds.)
© 2025 Apple Academic Press, Inc. Co-published with CRC Press (Taylor & Francis)

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 36 Nanotechnology for Agriculture

Moreover, nano-priming prevents the continued damage caused by conven-

tional agriculture, resulting in an environmentally safer system for farmers
and consumers. The present review provides an overview of developments in
the field, showing the challenges and possibilities of using nanotechnology
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in seed nano-priming to contribute to sustainable farming practices.

2.1 INTRODUCTION
In agricultural production, significant losses in yield occur due to biotic
stresses as well as climate change-related events such as salinity, heat,
cold, and drought. It is possible to increase agricultural productivity using

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chemicals such as fertilizers, herbicides, and pesticides. However, excessive
use of agrochemicals can be dangerous for human health and the environ-
ment. Therefore, the development of ‘green’ technologies is an alternative
to sustainable agriculture [1]. Recently, nanotechnology has shown great
potential, which raises hopes for improving agricultural productivity and food
security, considering the negative impacts of chemical inputs on the environ-
ment [2]. Nanotechnology works with nanoparticles (NPs) of less than 100
nm to solve many issues, including agricultural problems [3, 4]. NPs act as
plant growth accelerators that can improve water and nutrient intake [5, 6].
Since NPs trigger metabolic processes in the early germination stage, they
increase the germination rate, stability, plant growth, yield, and quality of
crops [7]. When NPs interact with plants, they cause various morphological
and physiological changes depending on their chemical structure, size, and
dose, and can regulate plants’ interaction with their environment [8, 9].
Utilizing nanotechnology, fertilizer can be delivered at a predictable rate
via stimulus-responsive release and targeted to certain tissues or organisms,
potentially improving nutrient use efficiency (NUE) by gradually delivering
the nutrient for crop uptake [10]. Thus, nano-enabled agriculture is also
anticipated to more effectively target pests with smaller pesticide doses,
minimize extensive effects on soil health and biodiversity, and increase the
soil microbiota to improve soil function and nutrient cycling [11].
Besides, nano-priming is another aspect of nanotechnology that exposes
the seeds to a solution for a certain period of time before planting, allowing
partial hydration but not forming roots [12]. Hence, nano-priming is seed
coating using nanoformulation media to absorb seeds [13]. Nano-priming
can affect seed metabolism, activate some biochemical pathways in the plant,
and affect germination, seedling formation, and the plant’s entire life cycle,
keeping crop uniformity [14]. Nano-priming protects seeds during storage,

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 Nano-Priming as Seed Priming Technology for Sustainable Agriculture 37

ensures germination uniformity, and increases the resistance of seedlings

to stress conditions, improving plant growth [15]. Nano-priming has been
reported to enhance germination and plant growth compared to other seed-
priming methods. The small size of the nanoparticles and high permeability
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to plant cells explain why nano-priming is more effective than other priming
applications [1].
Nanoparticles alter the metabolism of plants and seeds. In addition, they
can strengthen the innate immune system by affecting the production of
hormones [16]. The effects on the innate immune system can increase resis-
tance to pathogens, resulting in limited use of pesticides, ultimately lowering
the concentrations of residual agrochemicals in food. Thus, the consumption

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of these products becomes safer [17]. Nanopriming also increases the produc-
tion of enzymes that facilitate nutrient uptake and improves root growth,
thus reducing the amount of fertilizer used [18]. Therefore, the application
of nano-priming not only improves farmers’ socio-economic status but also
supports agriculture’s sustainability [7].

2.2 NANOPARTICLES USED IN NANO-PRIMING

The application of NPs in seed priming is broadly categorized as metal-
based, carbon-based, and polymeric NPs.
NPs of different metal elements have been used for crop production, yield
increase and disease suppression in sustainable agriculture [6]. The most
common metal Np and metal oxides used in recent years are TiO2, Ag, CuO,
Au, ZnO, and CeO2 [19, 20]. Each primer has different properties and effects
[21]. For example, silver NPs can protect seeds from fungi and bacteria
[22]. CuO NPs act as fertilizers by supporting the growth of plants [23]. In
addition, iron oxide-based nanomaterials are more advantageous than other
nanoparticles because they are biocompatible, easy to prepare, easily usable,
and environmentally friendly [24]. The doses of nanoparticles used are also
very important. Otherwise, it has been found that nanoparticles used in high
doses can disrupt the soil structure and cause a decrease in its biota [25].

2.2.1 METAL-ELEMENT NANOPARTICLES

2.2.1.1 SILVER NANOPARTICLES (AGNPS)
AgNPs are commonly used in many fields due to their antimicrobial prop-
erties that protect seeds from pathogens [26, 27]. By regulating AgNPsCl-
accumulation, Na+ and K+ salinity stress stimulates tolerance. Seed priming

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 38 Nanotechnology for Agriculture

with AgNPs has been found to increase growth under abiotic stresses and
improve the physiology of plants [28]. It reduces the production of reac-
tive oxygen species (ROS) by enhancing the antioxidant defense system,
ultimately resulting in improved plant growth [29, 30]. It has also been
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proven that AgNPs increase the seed viability index under drought stress and
provide speedy seedling growth [31].

2.2.1.2 SILICA NANOPARTICLES (SINPS)

Silica is an important material used in technology and science [22]. Silicon
nanoparticles (Si NPs), among other NPs are crucial for enhancing photo-

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synthesis and plant growth in stressful environments [32]. Si NPs support
plant growth by increasing the photosynthetic capacity in plants. In addi-
tion, it increases plant tolerance and endurance of plant organs under stress
conditions, including salinity stress [33, 34]. Similarly, under drought stress,
amylase increased its activity and seed germination [35]. In addition, some
nano-silica particles can control the ectoparasites of animals and insects [21].

2.2.1.3 COPPER NANOPARTICLES (CUNPS)

Among the types NPs, copper oxide (CuO) NPs, are widely used because of
low cost, natural abundance, non-toxic with good electrical properties [36].
It is also commonly found in plant tissues being one of the micronutrients
necessary for growth that is effective in many physiological events [37].
CuO NPs increase the chlorophyll content, leaf growth and height of the
plant [23]. At the same time, the use of Cu NPs in drought stress has been a
promising tool for increasing crop production [21].

2.2.1.4 IRON NANOPARTICLES (FENPS)

Iron is one of the essential elements for the plant and plays an important
role in photosynthetic reactions [38]. FeNPs increase the amount of proteins
associated with photosynthesis and improve plant growth by increasing
the number of chloroplasts [39, 40]. At the same time, FeNPs was found to
increase enzymatic activity and reduce oxidative damage [41]. Priming with
Fe2O3 NPs improves germination potential, ensures germination homoge-
neity and increases germination percentage [42]. In addition, FeO NPs can
be used as fertilizers or pesticides [43].

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2.2.1.5 ZINC OXIDE NANOPARTICLES (ZNONPS)

Zinc oxide nanoparticles are the most commonly used NPs globally [44]. Zn
plays an important role in many reactions of plants [45]. ZnO NPs exhibit
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antifungal, antibacterial and antimicrobial activities [46]. Seed priming with

ZnO NPs increases antioxidant enzyme activities. It increases salt stress
tolerance by regulating the amount of Na+ [47]. In addition, it reduces plants’
toxic effects by inhibiting heavy metal intake [22]. ZNPs mobilize the nutri-
ents in the soil and transport them and properly to a plant [48].

2.2.1.6 TITANIUM NANOPARTICLES (NANO‑TI, NANO‑TIO2)

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TiO2, the ninth most abundant element on earth, plays a vital role in plant
growth [49]. Titanium dioxide (TiO2) nanoparticles have biochemical, physi-
ological and morphological effects on plants. It improves oxidative damage
and plant performance [50]. TiO2 increases seed germination and seedling
growth rate of NPS. It also increases the activity of protease and amylase
enzymes [51].

2.2.1.7 GOLD NANOPARTICLES (AUNPS)

Gold NPs (AuNPs) were found to increase germination, leaf area, leaf
number, chlorophyll content, plant height, and yield [1, 52]. Seed coating
with AuNPs increases plant water absorption [4]. It provides fast and uniform
germination of seeds [22].

2.2.1.8 SULFUR NANOPARTICLES (SNPS)

Sulfur is commonly used in agriculture. Sulfur increases tolerance to abiotic
and biotic stresses in plants. It improves plant growth and yield. Priming with
SNPs stimulates SOD and CAT activities. It reduces ROS accumulation and
oxidative stress [53]. It alleviated drought stress by increasing antioxidants
[54]. It also affects the detoxification of heavy metals [55].

2.2.1.9 CERIUM OXIDE NANOPARTICLES (NANOCERIA, CEO2-NPS)

Cerium oxide nanoparticles (nanoceria, CeO2-NPS) are commonly used
because they are a powerful reducer of the harmful effects of ROS accu-
mulation during abiotic stress [56, 57]. Nanoceria improves plant salt stress
tolerance [58].

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 40 Nanotechnology for Agriculture

2.2.2 CARBON-BASED NANOPARTICLES

The use of carbon-based nanoparticles in seed priming includes single-

walled nanotubes, multiwalled carbon nanotubes, graphene, and fullerenes
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that can enhance germination, antioxidant status, and tolerance to a wider

range of extreme environmental events [59]. The carbon nanoparticles are
capable of piercing hard seed testa and altering water imbibition and hydra-
tion during germination [60]. The carbon-based NPs regulate biochemical
reactions such as hydrolysis of carbohydrates, infiltration of oxygen and
water, while down-regulating H2O2 production and electrolyte leakages
in plasma membranes [61]. The multiwalled carbon nanotubes have been

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shown to be reliable in activating antioxidant systems under various extreme
environmental events, improving seed germination in tomato, maize, boreal
forest species, lettuce, and other crop species [62–64].

2.2.3 POLYMERIC NANOPARTICLES

Polymeric nanoparticles are popular in the production of slow-releasing

fertilizers. Polymeric NPs can be made either naturally or synthetically. The
synthetic polysuccinimide polymeric NPs have been successfully adopted
for microbe control in certain plant species and seed priming in maize by
enhancing tolerance to heavy metal phytotoxicity via activating antioxidants
[65]. When the seeds are primed with chitosan NPs, they are effective in
salinity tolerance by increasing photosynthetic capacity and antioxidant
activity [66, 67].

2.3 SEED NANO-PRIMING AND ITS EFFECT ON ABIOTIC STRESS

Although harmful to plants, abiotic stresses can activate various signaling

pathways for short durations and with low intensity. Thus, the plant shows
increased resistance in the event of further stress encounters [14, 68, 69].
Seed priming makes seeds more resistant to future exposure by pre-exposing
them to abiotic stress [12]. The use of seed nano-priming to trigger plant
resistance to abiotic stress is a powerful alternative to reduce production
losses and avoid the side effects of the global climate crisis [17].
Fe2O3 and AgNPs reduce the damage caused by salt stress [70, 71]. An
et al. [58] reported reduced plant salt stress in field conditions in cotton
seeds coated with nanoceria. Nanoceria coated with 0.1 mM polyacrylic

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 Nano-Priming as Seed Priming Technology for Sustainable Agriculture 41

acid increased rapeseed seeds’ germination rate and biomass under salt
stress [72]. Coating with ZnO reduced the effect of salinity stress and
increased germination rate in wheat [73]. Pennisetum glaucum L. coating
of seeds with AgNPs reduced Na+ uptake in the plant and increased salt
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stress tolerance by maintaining the Na+/K+ ratio [28]. In another study,

the adverse effects of salt stress were reduced and germination percentage
increased in wheat seeds coated with AgNP (1 mg L–1) [74]. It was stated
that seed preparation with 60 mg L–1 ZnO could increase salt tolerance
in lupine plants [47]. A study examining the mitigating effects of NaCl-
induced toxicity damage in Brassica napus seed on physiological and
molecular parameters found that ZnO NPs reduced ion leakage [75].

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Wang et al. [76] found that nano-ZnO particles activate the antioxidant
system and improve salt tolerance in wheat by increasing the photosyn-
thetic electron transport efficiency in stressed leaves. Brassica napus L.
seeds were primed with different ZnONP doses (25%, 50%, 100%), it
was found that seed priming with 100% ZnO alleviated Na+ ion toxicity
in colza seedlings, thus reducing salinity stress [10]. Seed priming with
sodium metasilicate NPs increases salt stress tolerance [49]. The process
of priming Origanum majorana L. seeds with TiO2 nanoparticles is an
alternative way to improve seedling growth characteristics in saline condi-
tions [77]. Similarly, seed coating with Mn NPs increased the salt stress
tolerance of Capsicum annuum [78]. Drought stress significantly affects
plant growth. Seed coating with NPs has significantly reduced drought
stress [79]. Nguyen et al. [80], reported that resistance to drought stress
can increase in corn seeds coated with copper nanoparticles. ZnO NPS
was found to reduce the effect of drought in sorghum [81]. SiO2 np was
observed to increase drought stress tolerance and biomass production in
wheat seedlings [36]. Similarly, the effects of four different nano silicone
doses (0, 100, 200, and 500 mg L–1) on Calendula ofcinalis L. seeds, it
was found that Nano silicon applied at low concentrations (100–200 mg/L)
improved the drought plant physiological indices of the plant [82]. AgNPs
increased the percentage of seedling growth and germination in Festuca
ovina under drought stress [83]. Palmqvist et al. [84] reported that γ-Fe2O3
magnetic NPs reduced drought stress and enhanced the growth of Brassica
napus. CSNPs (chitosan nanoparticles) attenuated the effects of drought
stress, which resulted in decreased plant growth, total chlorophyll, relative
water content (RWC), and stomatal conductivity in Catharanthus roseus
[85]. Application of SNP and TiO2 NPs alone or in combination increased
wheat growth and germination under PEG-induced stress [86].

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 42 Nanotechnology for Agriculture

Shang et al. [87] found that nanoparticles improve the effects of pollut-
ants such as heavy metals. NPs alleviate metal stress in plants with their
large specific surface area, small size and high reactivity [65]. 2000 mg L–1
Fe3O4 NPs reduced the growth inhibition caused by heavy metals in wheat
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seedlings [88]. Cd concentrations and Cd toxicity in grains, shoots, and roots

of wheat seedlings whose seeds were coated with zinc oxide (ZnO) and
iron nanoparticles (Fe NPs) were significantly reduced [89]. The cadmium
stress of rice seeds was reduced coated with ZnO NPS, increasing amylase
activation and seedling growth [90]. It has been reported that SiO₂ NPs can
protect the plasma membrane of bamboo plant and protect the integrity of
cells against Pb-induced reactive oxidative (ROS) stress [91]. Ragab and

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Saad Allah [53] reported that sulfur NPs increased antioxidant defense and
decreased Mn toxicity in Helianthus annuus (L.) seedlings.
Triticum seeds primed with TiO2 and Phyllostachys edulis seeds primed
with SiO2 reduced the negative effects of Cd stress on germination and seed-
ling growth [92, 93]. ZnO NPs can be used as a Co stress relieving agent in
maize. Similarly, in wheat, seed priming with silicon nanoparticles has been
shown to reduce Cd content by 12–75% in grains, 10–52% in shoots, and
11–60% in roots [94].

2.4 NANO-PRIMING AND GERMINATION AND SEEDLING GROWTH

NPs improve nutrient uptake and plant growth by altering the metabolism
of the seed. These also stimulate the release of growth regulators [95].
Chandrasekaran et al. [4] reported that NPs increase the amount of
gibberellin and the mobilization of storage proteins. Improved seedling
vigor and seed germination were observed in Vigna radiata L. seeds coated
with 100 mg L–1 MgO NP [96]. Itroutwar et al. [97] reported an increase
in seedling viability index, seedling growth and germination in rice seeds
primed with ZnO nanoparticles. As a result of coating pepper seeds with
ZnO, Ag and TiO2 NPs, germination and seedling growth increased [98]. In
sorghum seeds coated with five different Fe2O3 doses, soaking with n- Fe2O3
at 10 mg/L was the best treatment to improve germination percentage and
rate. It was stated that seed preparation with n-Fe2O3 at 50 and 100 mg L–1
was the most effective application in improving seedlings [70]. Similarly,
germination, dry and fresh biomass, and root-shoot length increased in
chickpea seeds primed with Fe2O3 [99]. Rice seeds primed with 20 mg
L–1 FeO NPs showed a 50% increase in biophysical properties such as dry
weight and root length [43]. Onion seeds primed with ZnO, CuO, TiO2,

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 Nano-Priming as Seed Priming Technology for Sustainable Agriculture 43

and Ag NPs increased seed viability index, germination and root-shoot

length [100]. As a result of coating the corn seeds with Au nanoparticles
resulted in higher seed emergence, viability index and chlorophyll content
compared to the control [101]. Onion seeds prepared with AuNPs showed a
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63.2% increase in emergence percentage compared to the unprimed control

(37.4%) [13]. Similarly, when primed with ZnO NPs, wheat seeds’ increased
crop productivity and grain nutritional qualities were observed [102]. ZnO
NPs administered at low doses increased the rate of root growth and root
elongation in Brassica napus [103].
Similarly, priming Paeoniasu fruticosa seeds with TiO2 increased germi-
nation, viability index, seedling biomass, shoot and root length [104]. Ye et al.

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[78] reported that an average of 33% longer root occurs in Capsicum annuum
seeds primed with Mn NPs than in control. Copper nanoparticles increased
grain weight, number of grains per ear and ear length in wheat seeds [39].
CuO NPs at low concentrations (≤20 μg mL–1) have been shown to increase
plant growth in Lactuca sativa L. seeds [105]. Lining watermelon seeds with
iron nanoparticles increased germination and growth of shoots and roots
[106]. In wheat lined with multi-walled carbon nanotubes (MWCNTs), an
increase in stomatal density, grain number, biomass and water uptake were
observed [107]. An almost 4-fold increase in fruit production was observed
in tomato seeds primed with chitosan nanoparticles [108]. Improvements in
seedling strength and germination were observed in Solanum lycopersicum
seeds coated with TcTiONP [109]. Germination and seedling strength
increased in rice primed with 5 and 10 ppm AgNO3 compared to control
[110]. Germination and growth were increased in watermelon seeds [7].
nZVI priming increased seedling viability, promoted nutrient uptake and
ensured homogeneous germination of seedlings [111]. Shinde et al. [112],
Mg (OH) nanoparticles showed positive results in germination and growth in
Zea mays seedlings. The priming of Astragalus squarrosus seeds with SiO2
supported the germination percentage [113]. Silicon dioxide nanoparticles
have been found to affect yield enhancement and phytoremediation [114]. In
cotton seeds priming with poly (acrylic acid) coated cerium oxide nanopar-
ticles (PNC), seedling root length (56%), dry weight (38%), fresh weight
(41%) and root viability (114%) increased compared to the control [58].

2.5 EFFECT OF NANO-PRIMING AND ENZYME ACTIVITY

Nano-priming is used for antioxidant enzyme regulation in plants. Jadczak
et al. [115] reported that seed coating with silver and gold nanoparticles

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had a positive impact on antioxidant enzyme catalase (CAT), ascorbate

peroxidase (APX), guaiacol peroxidase (POX), and superoxide dismutase
(SOD) activity. Hibiscus sabdariffa L. seeds primed with 0.01% Al2O3 NPs
showed improvements in chlorophyll α, chlorophyll β, soluble sugars, carot-
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enoid content, protein, proline, amino acids, and defense enzymes such as
CAT, SOD, and peroxidase [47]. Afzal et al. [43] reported that seed priming
with FeO NP stimulated sugar, amylase, and sugar content in rice seeds
and increased antioxidant enzyme activity. The activities of CAT, SOD,
phenylalanine ammonia-lyase (PAL), proline, total phenolic contents, and
RWC were increased in maize seeds primed with TiO2 NPs [49]. Similarly,
applying ZnO NPs to rice seeds increased their biochemical and physi-

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ological activities, such as SOD activity at the shoot and root and peroxidase
(POD) activity at the root. Priming wheat seeds with Si NPs increased
chlorophyll content and antioxidant enzyme activity compared to the control
[94]. In addition, enzymes such as SOD, CAT, ascorbate peroxidase (APX),
and glutathione reductase (GR) showed a significant increase in activity with
ZnO NPs [6]. Polyacrylic acid-coated nanospheres provided high SOD, POD,
and α-amylase activity [72]. TcAg NPs increased the activity of antioxidant
enzymes such as superoxide dismutase, peroxidase, and catalase [109]. SiO2
NPs increased the capacity of the antioxidant enzymes CAT, SOD, PAL, and
GR [91].

2.6 EFFECT OF NANO-PRIMING AND ROS

ROS production is the immediate response of plants under abiotic stresses

such as salinity, drought, heat and heavy metals, which cause significant
damage to the cell structure. Nanoparticles reduce cell damage by lowering
ROS levels [18]. CeO2-NPs priming with poly (acrylic acid) reduced leaf
ROS levels in Arabidopsis thaliana [116]. Ragab and Saad-Allah [117] found
that seed coating with sulfur nanoparticles reduced total ROS overproduc-
tion and oxidative stress. Seed preparation with cerium oxide nanoparticles
(PNC) resulted in a 46% reduction in ROS accumulation in seedling roots
[58]. AgNPs developed increased ROS removal ability in primed rapeseed
[28]. El-Serafy et al. [118], reported a reduction in ROS levels when priming
Lathyrus odoratus seeds with Si NPs α-Fe2O3 nanoparticles reduced ROS
production of maize and rice seeds [24]. ZnO nano-priming provided
osmotic protection and reduction of ROS accumulation [10]. Priming maize
seeds with ZnO NPs and copper NPs reduced the level of ROS [119, 120].

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2.7 EFFECT OF NANO-PRIMING ON PATHOGENS

Nanoparticles act directly against pathogens. They can change the metabo-
lism of plants and hormone production, and seeds strengthen the immune
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system. In addition, plants increase their resistance to abiotic stress or

diseases [16, 17]. Dhabaan et al. [121] reported in their study that nano
chitosan has activity against pathogens in seedlings. Kumaraswamy et al.
[122] reported that chitosan NPs have antimicrobial and pathogen-removing
properties. Chitosan nanoparticles showed increased resistance to pathogens
and changes in the immune system in Panicum seeds [123]. Similarly,
Choudhary et al. [124] stated that zinc-containing chitosan nanoparticles

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make corn plants more resistant to pathogens. In addition to all these, AgNPs
were found to prevent the formation of bacterial cancer disease in tomato
seeds [125]. Rahman et al. [126] reported that platinum, silver, and gold
nanoparticles decreased in the root microbiota region of pea seeds, rhizobial
colonization, and mycorrhizal fungi. Chitosan nanoparticles showed anti-
fungal activity in rice seedlings [127]. TiO2, ZnO, and Ag NPs suppressed
pathogens such as Colletotrichum capsici, Aspergillus flavus, Aspergillus
fumigatus, and Aspergillus niger in pepper seeds [98].

2.8 CONCERNS

Nanomaterials have properties that can directly affect agricultural activities

[128]. Therefore, it is necessary to understand the mechanism of action of
nanoparticles very well. In addition, it is very important to develop nanopar-
ticles that can be used safely in laboratory and field conditions. Priming
should be done considering the nanoparticle concentration, exposure time,
and size, because these factors may show side effects such as reducing plant
growth, inhibiting germination, and adversely affecting root-microbiota
interactions [89, 126, 129]. Khalaki et al. [22] reported that the toxic effects
caused by NPs depend on the concentration, type, seed size, plant species,
and the NPs. Their application in high concentration may adversely affect
plant growth and germination by altering inappropriate preparation time, cell
structure, and plant metabolism [98, 129]. In addition, some nanoparticles
can be toxic to plants or seeds by re-oxidation. For example, silver nanopar-
ticles can be re-oxidized, inactivating enzymes. This may cause harmful
effects on the plant and seed [130]. Nanoparticles in agriculture have been
seen as having great potential in many studies. However, some nanoparticles

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 46 Nanotechnology for Agriculture

can be toxic to living things in the soil and can also alter the microbiome
selected by the roots [17]. NPs used in agricultural areas should be non-
toxic, environmentally friendly, biocompatible, and cost-effective [43, 131].
All these are reasons for concern in the use of NPs in agriculture.
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2.9 CONCLUSION
Seed nano-priming has the potential to improve crop production sustainably.
It increases the resistance of plants against stress conditions and protects
seeds by suppressing pathogens and diseases. In addition, some reports have
demonstrated that seed coating with NPs can alleviate heavy metal stress.

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Nano-priming can help optimize the amount of fertilizers and pesticides. It
can also reduce the amount of work on agricultural machinery by preventing
soil compaction. However, the interactions of NPs with plants need to be
clarified. The NP type and doses used in seed priming are very important. In
addition, the residues left by NPs in the environment and their effects need
to be clarified. This review has been prepared to contribute to the literature
by providing information about the nano-priming application.

KEYWORDS

• abiotic stress
• enzymes
• germination
• nanoparticles
• priming
• reactive oxygen species
• seedling growth
• sustainable agriculture

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