Advanced Agrochem 3 (2024) 39–42

Contents lists available at ScienceDirect

Advanced Agrochem
journal homepage: www.keaipublishing.com/en/journals/advanced-agrochem

Nano-controlled release of phytohormones will broaden its application on

plant protection
Zixia Liu , Fanglin Wen , Xiaolei Cheng , Zhibing Wu *
National Key Laboratory of Green Pesticide, Key Laboratory of Green Pesticide and Agricultural Bioengineering, Ministry of Education, Center for R&D of Fine Chemicals of
Guizhou University, Guiyang, 550025, China

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

Keywords: Phytohormone is a key regulator of plant growth and development. It has important effects on plant under biotic
Phytohormone and abiotic stresses. However, the dose control of phytohormone is always a difficult problem in the application
Biological/abiotic stress process, which limits the application range of phytohormone. Nanotechnology, because of its characteristics of
Growth and development
controlled release, targeted therapy, non-pollution, high adsorption, lower volatilization of active substances, and
Nanotechnology
Application prospect
low dosage of drug, comes into researchers’ vision. Nanomaterials were directly applicated on crops at the early
stage, and then active substances, such as pesticides, were encapsulated with nanomaterials, also achieved good
results in the field. Currently, more and more attentions have been paid to nano-enabled delivery of phytohor-
mones to plants, and formed a new field in agriculture. In present work, we reviewed the existing literatures,
focused on the important regulatory roles of phytohormones in plant growth and development and their appli-
cation potential, and the development and application prospect of nanomaterials combined with phytohormones
were also have been discussed.

1. Introduction substances exhibit more advantages in reducing pesticide use, controlling

release, reducing volatilization, precise targeting, high adsorption and so
Phytohormones is a class of organic substances with physiological on.
activity that produced by plant metabolism, which play important roles In present work, we focus on the key roles of phytohormones in plant
in plant growth and development regulating1–3 (rooting, germination, growth and development under stress, the application of nanotechnology
flowering and fruiting, cell elongation and division, etc.) by migrating to in agricultural development, and the research status and development
the different tissues and organs of plants.4,5 Auxin (IAA), abscisic acid prospect of the combination of nanotechnology and phytohormones.
(ABA), gibberellin (GA), ethylene (ET), cytokinin (CTK), brassinolide
(BR), salicylic acid (SA), and jasmonic acid (JA) are the common phy- 2. Regulation roles of phytohormones for plant growth and
tohormones, playing crucial roles in plant resistance enhancing, yield development
increasing and the environment adapting, and make them become an
indispensable factor in the response to biotic and abiotic stresses, which Phytohormones can regulate the synthesis of some proteins and
are widely used in agricultural production.6–8 However, during the special enzymes by binding to the corresponding hormone receptor
application, there are much unreasonable using cases, such as abuse, proteins in cells, and show the function of immune regulation. Six main
arbitrary increase of concentration, and no differentiation of application phytohormones16 (IAA CK, GA, ABA, ETH, and BR) played decisive roles
time, which not only affect the normal growth and development of in plant growth and development, and their functions were to regulate
plants, but also reduce the resistance of plants and increase the risk of biosynthesis, metabolism, perception, promote plant growth and devel-
disease and insect susceptibility of plants.9–11 As an emerging technology opment, immunity and signal transduction.17–19 Among them, IAA can
in agricultural development, nanotechnology is affecting the layout of regulate enzyme activity, relax cell wall, promote cell elongation, and
agricultural production. Among them, some nanomaterials can be participate in the growth and development of various organs and parts of
directly used to control pathogens or activate plant resistance,12–15 and the plant20; GA can stimulate the decomposition of starch and other
the nanoparticles that made by metal oxides and pesticide active storage compounds in seeds to promote stem elongation and seed

* Corresponding author.
E-mail address: zbwu@gzu.edu.cn (Z. Wu).

https://doi.org/10.1016/j.aac.2023.11.004
Received 30 September 2023; Received in revised form 2 November 2023; Accepted 15 November 2023
Available online 17 November 2023
2773-2371/© 2023 The Authors. Publishing services by Elsevier B.V. on behalf of KeAi Communications Co. Ltd. This is an open access article under the CC BY-NC-ND
license (http://creativecommons.org/licenses/by-nc-nd/4.0/).
Z. Liu et al. Advanced Agrochem 3 (2024) 39–42

germination21; CK can participate in cell division, proliferation and dif- 3. Application of nanotechnology in plant growth and disease
ferentiation, and also delay cell senescence22; ETH can promote fruit control
ripening and cell senescence23; ABA can catalyze the shedding of leaves,
fruits, etc.24; and BR can play a crucial role in various physiological Nanotechnology means that the research and development of mate-
processes, including cell elongation, differentiation, and stress rials and structures on the basis of nanoscale.34,35 The "small size"
response.25 (<100 nm) of nanomaterials enabled them easily cross the biological
Exogenous phytohormones were usually applied on plants that under barriers and enter the vascular system of plants after sprayed in the root
stress to ameliorate the adverse effects caused by abiotic/biotic stress or leaf surface, and kept a metastable state in the plant body,36 then the
(Fig. 1). Such as, Wang et al. found that exogenous zeatin (ZT) and GA3 effective substances were slowly released, playing a regulating role in the
could effectively alleviate cell damage in leaves under drought condi- growth and development of plants.
tions26; Tang et al. verified that exogenous SA (100–200 mg/L) could The application of materials that made with nanotechnology in
significantly increase the plant height and biomass of lettuce under heavy agriculture mainly has two forms:
metal cadmium (Cd) stress, and down-regulate the expressions of Nramp
5, HMA 4 and SAMT, reduce the absorption of Cd by lettuce roots and (a) Nano-materials directly acted on plants as agricultural production
block the upward transport of Cd27; Wang et al. found that exogenous supplies. Nanobionics provides an opportunity to embed nano-
melatonin (MT) had a positive effect on wheat germination under salt structures into plant systems, which can improve light capture and
stress by the analysis from physiological, biochemical indexes, and energy conversion ability of plants, enhance plant photosynthesis,
transcriptome level, which could not only reduce oxidation load and and promote plant growth and development.37 As the most com-
germination energy metabolism of wheat to maintain normal cell mon nanomaterials, carbon nanotubes and metal oxide nano-
morphology, and oxidoreductase activity and plant hormone-dependent particles can directly act on plants.38 Ma et al. sprayed nanoscale
signal transduction pathways also could be altered28; Li et al. found that hydroxyapatite (nHA) on tomato leaves three times to resist the
SA regulated the phosphorylation of FoSNF1 through the FoSNF1-FotorC1 infection of tomato wilt, and found that the above-ground salicylic
signaling pathway to inhibit the activity of FoTORC1, resulting in growth acid content of tomato plants increased by 10% ~ 45% and the
stagnation, cell viability, and pathogenic reduction of Fusarium oxy- activity of phenylalanine ammoniase and total phenol content
sporum, and prevented its invasion and reproduction29; Liu et al. found increased by 30–80% and 40–68%, respectively. All the results
that compared with the untreated group, the expression of flavonoid indicated that there was a potential relationship between antiox-
synthesis related genes in watermelon plants of SA treated group was idants and plant hormone pathways in NHA-promoted anti-fungal
up-regulated, the total flavonoid content was increased, and the proba- infection defense, and nHA could enhance plant disease resistance
bility of infection with Cucumber green mottled virus (CGMMV) was lower by improving hormone levels in plants.39
than that in the untreated group, which indicated that exogenous SA (b) Nanomaterials were used as the carrier, the active substances
could prevent CGMMV from infecting watermelon plants30; Huang et al. (nutrients or pesticides et al.) were embedded or covered on its
found that exogenous SA could effectively prevent the invasion of rice by surface, then the nanomaterials were used to achieve the purpose
Rice black stripe dwarf virus (RBSDV).31 In general, the application of of slow release and prolonged efficacy by their characteristics.
exogenous phytohormones could play an immunomodulatory role on There were two main types of nanomaterials combined with plant
plants. hormones, encapsulation and loading. Liposomes, because of their
Although the using of exogenous phytohormones could alleviate the lipophilic/hydrophilic and compartmentation properties, are
adverse effects caused by biological/abiotic stress and promote the usually used for encapsulation40; Layered carriers (such as mag-
normal growth and development of plants. However, natural phytohor- nesium aluminum layered double hydroxides), which can trans-
mones were prone to degradation because of the influence of light and port drugs to the target, and can also adsorb heavy metals and
enzymes during the application. Meanwhile, phytohormones are dose- other contaminants41,42; Chitosan nanoparticles are widely used
dependent, could not be applied at once and need to be added and sup- because of their biocompatible, biodegradable and non-toxic
plemented to achieve the effect of slow release.32,33 Therefore, the using characteristics.43 For example, Cao et al. successfully coated the
concentration and frequency of phytohormones were very important, water-soluble chitosan derivative (N-(2-hydroxyl)
which further revealed that accurate dosage and slow-release control was propyl-3-trimethyl chitosan ammonium chloride (HTCC) on the
the primary challenge facing the current practical application of plant surface of MSN loaded with pyrrolidine. HTCC modified MSN
hormones. achieved the same therapeutic effect despite reducing the using
amount of pyridoxylate by half.44 Wang et al. combined graphene
oxide (GO) with three pesticides (Pyridaben (Pyr), Chlorpyrifos

Fig. 1. The regulatory effects of plant hormones on plants.

40
Z. Liu et al. Advanced Agrochem 3 (2024) 39–42

(CHL), and Cyfluthrin (Cyf)) to obtain nano-materials, and used for GA3, and ALGCS-GA3 nanoparticles were found to be the most effective in
the prevention and control of phyllobiformis and cyclo- increasing leaf area and chlorophyll and Carotenoid levels, and also
phyllobiformis, which found that GO exhibited obvious synergistic could promot plant growth and seed germination55; Korpayev et al.
effects for three pesticides.45 Therefore, after combining the found that Ag and CS nanoparticles loaded with IAA and indole butyric
nanomaterials with active substances, the application efficiency acid (IBA) could improve the growth ability of roots, and the nano-
became higher, the energy was more saving, the toxicity of sub- particles that combined with hormones and Ag or CS exhibited better
stances were reduced, and the safety was improved. effects on plant growth than free IAA or nanomaterials alone56; Clemente
et al. extracted lipids from olive residues to prepare nanocarriers and
Nanomaterials have better control properties than traditional agro- combined with IAA, found that it could improve root growth ability.57 It
chemicals, making them be a more sustainable and environmentally could be seen that although some nanomaterials may directly act on
friendly option. However, nanomaterials still have some problems in plants as agricultural products, but the effect was greatly improved after
their applications. First, most of them are prepared by physical and phytohormones combining with nanomaterials. Therefore, nanoparticles
chemical methods (such as grinding, laser ablation, thermal decompo- prepared with phytohormones and nanomaterials have great potential
sition or hydrothermal), and these processes are very energy-consuming, for plant control and growth.
the auxiliary reagents are expensive, which may also result in toxic
compounds that harmful to the environment during the procedures.46 5. Conclusion
Second, when the accumulation of nanomaterials reaches over a certain
value in soil or water, it may exhibit negative effects on plants, ecology In recent years, with the gradual deepening of research on phyto-
and even human health. In order to solve the problems, more and more hormones, it was found that their performances in improving plant
researchers have changed their attention on the biosynthesis processes. resistance and promoting plant growth and development were surprising.
Biosynthetic nanomaterials, using active molecules for capping and sta- As a new agricultural product, phytohormones can improve plant toler-
bilizing, do not require external reducing agents and stabilizers, which ance. However, the acquisition of exogenous plant hormones is still faced
will reduce cost and save time. Meanwhile, toxic chemical reagents are with no small difficulties, and exogenous plant hormones are mainly
not used in the preparing procedure, friend to the environmental and less prepared by synthetic methods, whose production requires significant
threat or effect on the environment and human beings.47 material costs and may produce additional chemical pollution. Therefore,
the development of methods for obtaining physiologically active sub-
4. Nano delivery of phytohormones stances for agriculture from natural biological raw materials containing
plant hormones has become an important research direction in the
Phytohormones can promote plant growth and improve the effects future. In addition, after the combination of nanotechnology and phy-
caused by abiotic or biotic stress. However, when phytohormones were tohormones, it has effectively improved the unstable characteristics of
combined with nanotechnology to form nanoparticles with nano- hormones in the use environment, better stimulated the performances of
materials as carriers and phytohormones as active substances, which hormones, and greatly reduced the negative impacts of improper dosage
exhibited multiple advantages, such as targeted therapy, precise control, on plant growth and development when it was used alone. Based on the
reduced dosage, slow release, and no pollution after being transported to existing literatures, it was demonstrated that the interactions between
plants (Fig. 2). There are two different nano delivery ways of plant nanotechnology and phytohormones have only been preliminarily stud-
hormones. One is by xylem (root absorption), which transports water and ied. In addition, the regulatory relationships between exogenous hor-
minerals from roots to stems and leaves, and further be transported with mones and hormones produced by plant-self also need to be deep studied
exocytosis and symplastic transport.48–50 The other is by the phloem (leaf in order to improve plant tolerance better. And the selection of different
spray), which refers to the spraying of adsorbed nanomaterials through spray methods according to different plants and diseases also brings new
the leaf surface and then transports to the phloem system through cuticle requirements and challenges to the preparation process of nanocarriers.
penetration or directly through pores (e.g. stomata and drainage pores), Most nanoparticles, prepared by the combination of plant hormones and
distributes to non-photosynthetic tissues and organs via sieve elements in nanomaterials, are currently in the laboratory stage, and have a long way
the phloem.51,52 For example, Polyakov et al. studied the control-release to be applied in the field. Before its large-scale application, it is crucial to
system of SA nanoparticles using various organic and inorganic com- anticipate both short-term and long-term impacts on the environment
pounds as the carriers, which mean that SA was wrapped in nano- and its ecological equilibrium and assess potential risks to food safety and
materials to achieve the purpose of control-release53; Hanif et al. human health. Certainly, the research and development of new nano-
synthesized CuO nanomaterials served as nano-carriers of IAA, and materials with better loadability, eco-friendly and lower cost will be the
enhanced the physiological parameters (root length, shoot length and focus of future phytohormones nano-delivery research.
biomass) of chickpea plants54; Pereira et al. used alginate/chitosan Nanoparticles, prepared by nanomaterials and phytohormones, not
(ALG/CS) and chitosan/Tripolyphosphate (CS/TPP) as nano-carriers of only can reduce their using amount, but also accurately control its slow-

Fig. 2. Preparation and application of nanoparticles contain hormones.

41
Z. Liu et al. Advanced Agrochem 3 (2024) 39–42

release, which will make this kind of nanoparticles become a new di- 23. Bleecker AB, Kende H. ETHYLENE: a gaseous signal molecule in plants. Annu Rev Cell
Dev Biol. 2000;16:1–18.
rection for future agricultural research and development, and have a
24. Nguyen CH, Yan D, Nambara E. Persistence of abscisic acid analogs in plants:
significant impact on future agricultural development. chemical control of plant growth and physiology. Genes. 2023;14(5).
25. Zhang Z, Chen Z, Song H, Cheng S. From plant survival to thriving: exploring the
CRediT authorship contribution statement miracle of brassinosteroids for boosting abiotic stress resilience in horticultural
crops. Front Plant Sci. 2023:14.
26. Wang C, Yang A, Yin H, Zhang J. Influence of water stress on endogenous hormone
Zixia Liu: Writing – original draft, Investigation. Fanglin Wen: contents and cell damage of maize seedlings. J Integr Plant Biol. 2008;50(4):427–434.
Investigation. Xiaolei Cheng: Investigation. Zhibing Wu: Conceptuali- 27. Tang W, Liang L, Xie Y, et al. Foliar application of salicylic acid inhibits the cadmium
uptake and accumulation in lettuce (Lactuca sativa L.). Front Plant Sci. 2023:14.
zation, Writing – review & editing. 28. Wang J, Lv P, Yan Di, et al. Exogenous melatonin improves seed germination of
wheat (Triticum aestivum L.) under salt stress. Int J Mol Sci. 2022;23(15).
Declaration of competing interest 29. Li L, Zhu T, Song Y, et al. Salicylic acid fights against Fusarium wilt by inhibiting
target of rapamycin signaling pathway in Fusarium oxysporum. J Adv Res. 2022;39:
1–13.
The authors declare that they have no known competing financial 30. Liu M, Kang B, Wu H, et al. Transcriptomic and metabolic profiling of watermelon
interests or personal relationships that could have appeared to influence uncovers the role of salicylic acid and flavonoids in the resistance to cucumber green
mottle mosaic virus. J Exp Bot. 2023;74(17):5218–5235.
the work reported in this paper. 31. Huang R, Li Y, Tang G, et al. Dynamic phytohormone profiling of rice upon rice
black-streaked dwarf virus invasion. J Plant Physiol. 2018;228:92–100.
Acknowledgement 32. Butova VV, Bauer TV, Polyakov VA, Minkina TM. Advances in nanoparticle and
organic formulations for prolonged controlled release of auxins. Plant Physiol Biochem
(Issy les Moulineaux, Fr). 2023:201.
This work was supported by the National Natural Science Foundation 33. Sampedro-Guerrero J, Vives-Peris V, Gomez-Cadenas A, Clausell-Terol C. Efficient
of China (No. 32160655), and Breeding Program of Guizhou University strategies for controlled release of nanoencapsulated phytohormones to improve
(No. 201931). plant stress tolerance. Plant Methods. 2023;19(1).
34. Saleem H, Zaidi SJ. Recent developments in the application of nanomaterials in
agroecosystems. Nanomaterials. 2020;10(12).
References 35. Sarraf M, Vishwakarma K, Kumar V, et al. Metal/metalloid-Based nanomaterials for
plant abiotic stress tolerance: an overview of the mechanisms. Plants. 2022;11(3).
1. Berens ML, Berry HM, Mine A, et al. Evolution of hormone signaling networks in 36. Lowry GV, Avellan A, Gilbertson LM. Opportunities and challenges for
plant defense. Annu Rev Phytopathol. 2017;55(1):401–425. nanotechnology in the agri-tech revolution. Nat Nanotechnol. 2019;14(6):517–522.
2. Bl

azquez MA, Nelson DC, Weijers D. Evolution of plant hormone response pathways. 37. Mony C, Kaur P, Rookes JE. Nanomaterials for enhancing photosynthesis: interaction
Annu Rev Plant Biol. 2020;71(1):327–353. with plant photosystems and scope of nanobionics in agriculture. Environ Sci Nano.
3. Wang L, Hart BE, Khan GA, et al. Associations between phytohormones and cellulose 2022;9:3659.
biosynthesis in land plants. Ann Bot. 2020;126(5):807–824. 38. Tarrahi R, Mahjouri S, Khataee A. A review on in vivo and in vitro nanotoxicological
4. Saini S, Kaur N, Pati PK. Phytohormones: key players in the modulation of heavy studies in plants: a headlight for future targets. Ecotoxicol Environ Saf. 2021:208.
metal stress tolerance in plants. Ecotoxicol Environ Saf. 2021:223. 39. Ma C, Li Q, Jia W, et al. Role of nanoscale hydroxyapatite in disease suppression of
5. Verma V, Ravindran P, Kumar PP. Plant hormone-mediated regulation of stress fusarium-infected tomato. Environ Sci Technol. 2021;55(20):13465–13476.
responses. BMC Plant Biol. 2016;16(1). 40. Laouini A, Jaafar-Maalej C, Limayem-Blouza I, et al. Preparation, characterization
6. Han X, Kahmann R. Manipulation of phytohormone pathways by effectors of and applications of liposomes: state of the art. J Colloid Sci Biotechnol. 2012;1:
filamentous plant pathogens. Front Plant Sci. 2019;10. 147–168.
7. Li M, Zhu Y, Li S, et al. Regulation of phytohormones on the growth and development 41. Verma KK, Song XP, Joshi A, et al. Recent trends in nano-fertilizers for sustainable
of plant root hair. Front Plant Sci. 2022;13. agriculture under climate change for global food security. Nanomaterials. 2022;12.
8. Zhao B, Liu Q, Wang B, Yuan F. Roles of phytohormones and their signaling 42. He T, Li Q, Lin T, et al. Recent progress on highly efficient removal of heavy metals
pathways in leaf development and stress responses. J Agric Food Chem. 2021;69(12): by layered double hydroxides. Chem Eng J. 2023:462.
3566–3584. 43. Seon Sim H, Su Jo J, Jeong Woo UJ, et al. Abscisic acid, carbohydrate, and
9. Kavino M, Harish S, Kumar N, et al. Induction of systemic resistance in banana (Musa Glucosinolate metabolite profiles in Kimchi cabbage treated with extremely high
spp.) against Banana bunchy top virus (BBTV) by combining chitin with root- temperatures and chitosan foliar application. Sci Hortic. 2022:304.
colonizing Pseudomonas fluorescens strain CHA0. Eur J Plant Pathol. 2007;120(4): 44. Cao L, Zhang H, Cao C, et al. Quaternized chitosan-capped mesoporous silica
353–362. nanoparticles as nanocarriers for controlled pesticide release. Nanomaterials. 2016;
10. Niedz RP, Garmendia A, Beltr
an R, et al. Gibberellic acid in Citrus spp. flowering and 6(7).
fruiting: a systematic review. PLoS One. 2019;14(9). 45. Wang X, Xie H, Wang Z, He K. Graphene oxide as a pesticide delivery vector for
11. Zhang J, Vrieling K, Klinkhamer GL, et al. Exogenous application of plant defense enhancing acaricidal activity against spider mites. Colloids Surf, B. 2019;173:
hormones alters the effects of live soils on plant performance. Basic Appl Ecol. 2021; 632–638.
56:144–155. 46. Omran Basma A, Baek KH. Control of phytopathogens using sustainable biogenic
12. Cao X, Wang Z. Application of nano-agricultural technology for biotic stress nanomaterials: recent perspectives, ecological safety, and challenging gaps. J Clean
management: mechanisms, optimization, and future perspectives. Environ Sci Nano. Prod. 2022;372:133729.
2022;9(12):4336–4353. 47. Yang Y, Waterhouse GIN, Chen Y, et al. Microbial-enabled green biosynthesis of
13. Kannan M, Bojan N, Swaminathan J, et al. Nanopesticides in agricultural pest nanomaterials: current status and future prospects. Biotechnol Adv. 2022;55:107914.
management and their environmental risks: a review. Int J Environ Sci Te. 2023; 48. Schwab F, Zhai G, Kern M, et al. Barriers, pathways and processes for uptake,
20(9):10507–10532. translocation and accumulation of nanomaterials in plants-critical review.
14. V
azquez-Nú~ nez E. Uso de nanomateriales en la agricultura y sus implicaciones Nanotoxicology. 2016;10:257–278.
ecol
ogicas y ambientales. Mundo Nano. Revista Interdisciplinaria en Nanociencias y 49. Wang Z, Xie X, Zhao J, et al. Xylem- and phloem-based transport of CuO
Nanotecnología. 2022;16(30):1e–25e. nanoparticles in maize (Zea mays L.). Eviron Sci Technol. 2012;46:4434–4441.
15. Wang P, Lombi E, Zhao F, Kopittke PM. Nanotechnology: a new opportunity in plant 50. Larue C, Veronesi G, Flank AM, et al. Comparative uptake and impact of TiO2
sciences. Trends Plant Sci. 2016;21(8):699–712. nanoparticles in Wheat and rapeseed. J Toxicol Environ Health A. 2012;75:722–734.
16. Wang Y, Irving HR. Developing a model of plant hormone interactions. Plant Signal 51. Eichert T, Kurtz A, Steiner U, et al. Size exclusion limits and lateral heterogeneity of
Behav. 2011;6(4):494–500. the stomatal foliar uptake pathway for aqueous solutes and water-suspended
17. Anfang M, Shani E. Transport mechanisms of plant hormones. Curr Opin Plant Biol. nanoparticles. Physiol Plantarum. 2008;134:151–160.
2021;63. 52. P
erez-de-Luque A. Interaction of nanomaterials with plants: what do we need for real
18. Bittner A, Cie
sla A, Gruden K, et al. Organelles and phytohormones: a network of applications in agriculture? Front Environ Sci. 2017;5.
interactions in plant stress responses. J Exp Bot. 2022;73(21):7165–7181. 53. Polyakov V, Bauer T, Butova V, et al. Nanoparticles-Based delivery systems for
19. Nascimento FX, Glick BR, Rossi MJ. Isolation and characterization of novel soil- and salicylic acid as plant growth stimulator and stress alleviation. Plants. 2023;12(8).
plant-associated bacteria with multiple phytohormone-degrading activities using a 54. Hanif S, Javed R, Khan A, et al. IAA-decorated CuO nanocarriers significantly
targeted methodology. Access Microbiology. 2019;1(7). improve Chickpea growth by increasing antioxidative activities. 3 Biotech. 2023;
20. Matsuo S, Kikuchi K, Nagasuga K, et al. Transcriptional regulation of auxin 13(3).
metabolic-enzyme genes during tomato fruit development. Sci Hortic. 2018;241: 55. Pereira AES, Silva PM, Oliveira JL, et al. Chitosan nanoparticles as carrier systems for
329–338. the plant growth hormone gibberellic acid. Colloids Surf, B. 2017;150:141–152.
21. Kim YC, Nakajima M, Nakayama A, Yamaguchi I. Contribution of gibberellins to the 56. Korpayev S, Karakeçili A, Dumano glu H, et al. Chitosan and silver nanoparticles are
formation of arabidopsis seed coat through starch degradation. Plant Cell Physiol. attractive auxin carriers: a comparative study on the adventitious rooting of
2005;46(8):1317–1325. microcuttings in apple rootstocks. Biotechnol J. 2021;16(8).
22. Andreeva AA, Vankova R, Bychkov IA, et al. Cytokinin-regulated expression of 57. Clemente I, Menicucci F, Colzi I, et al. Unconventional and sustainable nanovectors
Arabidopsis thaliana PAP genes and its implication for the expression of chloroplast- for phytohormone delivery: insights on olea europaea. ACS Sustainable Chem Eng.
encoded genes. Biomolecules. 2020;10(12). 2018;6(11):15022–15031.

42