Zinc Oxide Nanoparticles: Abiotic Stress Tolerance in Fruit Crops Focusing on Sustainable Production

1  Introduction

Fruits are famous valuable crops because they are rich in minerals and vitamin levels necessary for a healthy life [1]. There is a serious global threat to fruit crop productivity [2]. A drastic decline was observed in crop productivity due to abiotic stresses. Abiotic stresses enhance the generation of free radicals and toxic substances, resulting in hampered photosynthesis, impaired metabolic processes, disturbed carbon assimilation and enhanced membrane permeability in plants [3]. Moreover, several developmental processes necessary for sufficient growth are disturbed due to adversities of abiotic stresses [4]. Mitigation of abiotic stresses is necessary for the sustainable productivity of fruit trees.

Different management strategies, i.e., supplementation of phytohormones, microbes, organic amendments and nanotechnology, are effective for the alleviation of abiotic stress tolerance in fruit crops globally [5,6]. All these management strategies have the potential to improve abiotic stress tolerance in fruit crops. The cultivation of tolerant germplasm can enhance the productivity of crops [7]. Therefore, selection, evaluation and identification of tolerant genetic resources is one of the imperative approaches for the cultivation of tolerant germplasm focusing on higher yield and superior quality.

Innovations in nanotechnology enhanced its utilization in the sustainable farming of fruit trees [8]. Nanoparticles had excellent contributions to orchard management due to their smaller size, greater porosity, better mobility and higher surface area [9–11]. These nanoparticles had the potential to boost plant growth and yield due to their small sizes [12]. Several plant researchers are paying greater consideration to metal oxide nanoparticles (NPs) due to their prodigious performance, less expensive, and climate-friendly nature [13]. The optimum use of metal nanoparticles is effective for sufficient growth and yield, while higher levels than the threshold showed adverse effects on growth and yield due to toxicity [14]. However, the selection of nanoparticles depends on several factors, such as type of crop, stress level, cultural practices, and climatic conditions within the growing regions [15].

Zinc (Zn) is a beneficial micronutrient with a significant contribution to plant homeostasis and regulatory mechanisms that occur in numerous horticulture crops [16]. Zn had an excellent contribution to cell membrane rigidity, chloroplast development, photosynthetic pigment synthesis, and improved endogenous hormones [17]. Zn acts as a cofactor for several enzymes, such as ligases, hydrolases, isomerases, and transferases to regulate cellular metabolism [18,19]. Zinc has a significant involvement in the regulatory functioning of minerals absorption and plant-water relations, which further contributes to alleviating abiotic stress tolerance in plants [20,21]. Hence, from previous literature, it has been explored that Zn is significantly involved in the enhancement of abiotic stress tolerance in crops. However, its application on fruit crops is still limited and needs more investigations to explore the beneficial role of Zn on fruit trees growing under adverse climatic conditions.

Severity in temperatures, heavy rainfalls, speedy winds, degraded soils, floods, and rising CO2 levels are majorly due to climate change. However, the fluctuations in climate change are causing the abiotic stresses in plants [22]. Therefore, fluctuations in climate change are becoming much more challenging for plant researchers and farmers to get higher yields from their fruit crops. The vegetative growth of fruit crops is badly affected by adverse climatic conditions. Fruit crops are perennial and mostly propagated from vegetative parts (asexually) to maintain the productivity and quality of fruit crops [23]. Drought conditions disturbed the emergence of vegetative flushes, reduced water level in leaves, stinted flushes length, and poor number of leaves in mango trees [24]. Flower induction was poor due to water deficit conditions in mango trees. Female flowering was disturbed due to adverse climatic conditions [25]. Earlier research suggested that higher temperature and water stress improve the floral stimulation in trees growing under tropical conditions. However, temperature extremities and water stress for longer periods reduced the productivity of crops [26,27]. Climate change drastically reduced the vegetative and reproductive processes due to the occurrence of abiotic stresses (Table 1). It is imperative to explore the vegetative and reproductive processes that occur in fruit trees subjected to abiotic stress conditions. Moreover, the management of abiotic stresses is necessary for sustainable fruit tree production and superior quality for consumption.

2  Influence of ZnO-NPs on Vegetative Growth of Fruit Crops

ZnO-NPs had the potential to mitigate adversities that occur from abiotic stress conditions in fruit crops. These had a better contribution in the enhancement of seed germination, seedling growth, roots emergence, leaf anatomy and vegetative expansion. Its beneficial contribution is based on application level, stress conditions, type of crop, and cultivation practices, which may vary. Amirjani et al. [37] revealed the synergistic interactions between Zn, K, and Mn contributed to the beneficial role of Zn in enhancing the K as well as Mn levels in leaf petioles. However, it has also been indicated that rising Zn levels decreased P and Fe levels, which may have been caused by Zn, Fe, and P differing responses [37]. The supplementation of ZnO-NPs as tiny carriers for the plant auxins indole-3-acetic acid (IAA) and indole-3-butyric acid (IBA) improved the rooting capability of Pyrus species. Karakeçili et al. [38] indicated that the rooting percentage was found to be greater at 400 mg L−1 of IBA-nZnO (50.0%) as well as IAA-nZnO (41.7%). Gupta et al. [39] stated that strawberry growth was improved by application of different nano-fertilizers. In another study by Bayat et al. [40], it has been studied that strawberry farming is more sustainable via the use of nano-fertilizers. However, ZnO-NPs fertilization of 20 ppm enhanced the tree growth-related traits of the Picual cultivar of olive, as reported by Genaidy et al. [41]. Similarly, Carlesso et al. [42] revealed that ZnO-NPs enhanced the physical attributes of strawberry crops growing within the region. Different concentrationss of ZnO-NPs (0 and 2 g L−1) were applied to the pear seedlings. It has been evaluated that vegetative growth comprising seedlings height, diameter, fresh and dry weights, leaf area, and number of leaves was improved by fertilization of ZnO-NPs [43]. Application of ZnO-NPs (180 mg L−1) enhanced the vegetative growth of grape vines cv. Flame Seedless. Another study by Mahdavi et al. [44] indicated that vegetative growth of grape berries was boosted with ZnO-NPs than control growing under calcareous land.

3  Power of ZnO-NPs on Reproductive Growth of Fruit Crops

Fruits are a vital source of nutrients for humans. Moreover, fruit crops are an important source of earnings for farming communities with higher incomes and a source of employment [45]. Abiotic stresses showed adverse effects on the growth phases of fruit crops subjected to adverse climatic conditions. Fruit crops majorly face salinity, drought, temperature, and heavy metal toxicity. Numerous morphological, anatomical, physiological, and biochemical alterations may occur in fruit crops, which further disturb the crop yield and fruit quality [45]. It is imperative to regulate reproductive growth for fruit crops under adverse climatic conditions (Fig. 1).

Figure 1: Abiotic stress conditions reduced the growth, yield and fruit quality of fruit crops by alteration in physiological and biochemical changes

Plant reproductive growth can be improved by the application of ZnO-NPs. The innovations in various management practices can be beneficial for sustainable yields of fruit trees. The bud initiation and formation were delayed due to drought conditions. Moreover, floral bud breakage occurred due to water deficit conditions [27]. The emergence of vegetative flushes was poor in mango plants subjected to drought stress. The decreased number of leaves in a flush, flush duration, and leaf water contents were recorded due to water stress conditions. Drought stress poses a significant threat to the floral induction of mango plants because floral induction is stimulated and produced from mature leaves. Flowering initiation can be enhanced on fruit trees by restricted water applications for a short period. However, prolonged water stress conditions may disturb the tree’s developmental processes [26]. The significant delay in flushing was recorded due to water stress, as depicted by Schaffer et al. [27]. Grapevines have excellent xylem arteries; therefore, these are considered more tolerant to drought stress as compared to some other fruit crops [46].

Banana crop production was affected by the low level of available moisture content in the soil. The carbon absorption was disrupted in plants due to restricted leaf water levels [47]. Moreover, the banana was also found to be sensitive to water deficiency at the flowering stage. Banana cultivar ‘Elakki’ productivity was lower due to water stress at the time of floral differentiation [48]. The maximum yield reduction was recorded in Banana cv. ‘Robusta’ when subjected to drought conditions for five weeks at the time of flowering [49].

Three cultivars of banana (Karpuravalli, Robusta, and Rasthali) were subjected to drought stress at the time of flowering. Moreover, the maximum yield reduction was recorded in cvs. Karpuravalli (42.07%), Robusta (25.0%), and Rasthali (18.83%) as illustrated by Ravi et al. [50]. Drought stress in papaya plants resulted in a 50% decrease in leaves, 86% flowering decline, and 58% yield reduction. Moreover, papaya fruit size and yield were also reduced due to drought stress conditions as reported by Masri et al. [51]. Different management practices i.e., growth promoters, use of different mulching materials, modifications managing nutrients, micro-watering, and timely irrigation practices, are important to mitigate adverse conditions that occur from water stress. Selection, characterization, and cultivation of tolerant cultivars need time to mitigate the adverse effects of abiotic stresses in fruit crops. Moreover, rootstock also had a significant contribution to the tolerance mechanism of abiotic stresses. Hence, appropriate management practices can enhance the abiotic stress tolerance mechanism in fruit crops [52].

ZnO-NPs had a positive impact on the improvements of the reproductive phase of fruits subjected to adverse climatic conditions. Different reproductive processes such as floral initiation, pollination and fruit formation of trees were significantly influenced by ZnO-NPs [53]. Similarly, a plant researcher indicated that ZnO-NPs (150 ppm) improved fruit yield and quality of strawberry cv. “Chandler” [54]. Mango yield was enhanced through the supplementation of 100 mg L−1 nZnO by Elsheery et al. [55]. Fruit crops such as pomegranate and mango trees that were sprayed with nano-Zn before flowering showed greater plant resilience to floral malformation disease and improved fruit quality [56,57]. The supplementation of zinc increases fruit retention percentage, fruit set per panicle, and flower biology in mango crops as revealed by the findings of Ahmad et al. [58]. The functionality of many enzymes comprising transphosphorylases, dehydrators, isomers, and aldolases also improved with the supplementation of ZnO-NPs. Furthermore, it has a regulatory contribution in the generation of tryptophan and protein and is occupied in cell division, photosynthetic processes, stomatal regulations, and prevention of membrane integrity [59]. ZnO-NPs boost the endogenous production of auxins. Auxins had a significant contribution in the improvement of reproductive phases of fruit growth and overall tree yield [60,61].

Floral initiation in mango can be influenced by the C/N ratio. Moreover, the accumulation and supply of carbohydrates also depend on the C/N ratio. Several metabolic processes and climatic factors disturbed the C/N ratio in trees, which resulted in the disturbance in the reproductive growth of trees, especially delay and irregular flower initiation observed in mango plants [62]. Modifications in phytohormones, particularly the decrease in gibberellin phases, coincide with a rise in sugars [63]. Hormonal balance is significantly maintained in plants by zinc (Zn) supplementation [64]. In fruit trees, the maximum fruit load on trees in a single year has been observed, while the minimum fruit load in the next year is mostly due to hormonal imbalances within the plants. Therefore, Zn was found to be effective for hormonal regulations in plants. Moreover, management practices can also regulate the fruit load on trees. The thinning of flowering may result in a low number of flowers on a single tree. The lower number of flowers on a tree results in a healthy fruit set and larger fruit size with superior fruit quality. The asymmetrical bearing habit of fruit load in mangoes can be regulated through various adaptive methods that successfully regulate endogenous phytohormones [65]. ZnO-NPs supplementation has improved fruit quality and yield with promising outcomes. Moreover, ZnO-NPs have been shown to improve the reproductive growth of fruit trees. However, its efficacy depends on application level, kind of crop, stress type and numerous other factors.

4  ZnO-NPs Regulate Morphological and Physiological Changes

ZnO-NPs had the potential to regulate morpho-physiological changes in fruit trees subjected to abiotic stresses. The over-generation of ROS is due to adverse climatic conditions in plants. The excess of ROS increased the oxidative harm in plants. The increased level of lipid peroxidation was found to be effective biomarkers for the determination of stress levels in plants. Different antioxidants (enzymatic and non-enzymatic) were regulated by the generation of various osmolytes and metabolites in plants. The regulation of ROS generation is necessary to protect plants from damaging effects that occur from oxidative harm due to adverse climatic conditions. Abd-Allatif et al. [66] determined the adverse effects of salinity stress in plants by disruption of several enzymes, disruption of cell membranes, protein impairment, and death of certain cells, tissues, and impaired functions. ROS overproduction was regulated by the activation of the plant defense system by modulation of several antioxidants and the formation of various osmolytes. These processes are required to maintain a balance in the formation of ROS, and the plants’ potential to scavenge toxic substances and free radicals depends on the robust immune response. Hence, optimum generation of ROS is important for sustainable fruit cultivation because membrane rupturing may reduce with balanced ROS production within the organelles [67,68]. Transgenic plants are tolerant to adverse conditions that occur from abiotic stresses. Moreover, transgenic plants exhibited higher levels of antioxidants and osmolytes which improved stress tolerance in plants against oxidative harm [69]. Fruit plants can effectively mitigate the negative effects of salinity through suitable fertilization. Fruit trees need to produce several bioactive substances which improve plants tolerance against various environmental challenges. Moreover, bioactive substances were boosted with exogenous supplementation of melatonin as exhibited by Chen et al. [70].

Fruit trees with suitable bioactive compound levels boost defense system, photosynthetic apparatus, stomatal regulation, and respiration rates which further contributes to the improvements of tolerance in plants [71]. Drought is the most critical environmental constraint that disturbs tree productivity [72]. Poor hydraulic conductivity was recorded in the xylem vessels of plants due to water deficit conditions. Moreover, stunted growth and low utilization of carbon levels were observed in plants due to drought conditions. However, poor utilization of carbon can also reduce the tree yield [73]. Root architecture is disturbed due to drought stress conditions in mango trees [74]. Moreover, salinity stress also alter the morpho-physiological activities of mango trees [75]. Drought stress enhanced the salt accumulation in the root biosphere of plants. Therefore, long-term exposure to salts is also toxic for higher plants. However, plants faced different salts such as calcium, magnesium, sodium, and chlorides. However, sodium salt is more dangerous for plants because its concentration may be higher in the root zone [76].

Mango is a valuable fruit crop. Excess of salts is toxic to mango trees because mango are sensitive to salt stress [77]. Salinity stress damages plant growth through osmotic imbalances and ionic distress [78,79]. The maximum leaf droppage was recorded in plants due to salinity stress by inhibition in nutrient absorption, carbon assimilation, and chlorophyll contents [80]. The suitable number of leaves contributes to better productivity of crops. The metabolic disturbances in plants were due to the degradation of proteins, lipids, and nucleic acids. Hence, metabolic disturbances allow the overgeneration of ROS in plants. The irregulates in ROS generation disturbed the morpho-physiological activities in plants [81]. The disturbances in morpho-physiological activities result in stunted tree growth, low yield, and inferior fruit quality, as explored by Ansari et al. [82].

The excessive ROS production is due to free radicals and toxic ions which disrupt homeostasis. The disturbances in osmotic pressure were due to excessive ROS generation. The rupturing of membranes, sugar oxidation, photosynthetic pigments degradation, denatured protein, disruption of antioxidants capability and reduced osmolyte formation are due to due to excessive ROS by altering the morpho-physiological activities. Soni et al. [83] revealed the excessive ROS production, and the oxidation of lipids as biomarkers to determine the severity of stress in plants, which disturbed the plant developmental processes. Plant cells and divisions that are subjected to sodium toxicity can reduce the tree yield. Lipid peroxidation assessment indicates that the damage to the membrane was caused by plants being exposed to more saline than normal conditions. The higher degree of damage to the plant cell membranes was indicated by the elevated presence of lipid peroxidation [83].

5  ZnO-NPs Boost Biochemical Activities in Fruit Crops

ZnO-NPs are involved in boosting biochemical activities in fruit trees subjected to abiotic stresses. The antioxidant enzymes and osmoprotectants could regulate and eliminate the excessive ROS within the cell organelles [84,85]. The activation of enzymatic, non-enzymatic, and metabolites contributes to reduce oxidative harm by ROS elimination [86]. Superoxide dismutase (SOD) is one of the most potent cytoplasmic metalloenzymes that contribute to the antioxidant defense network [43]. Moreover, it is comprised of metabolic factors that catalyze the conversion of superoxide anions into dioxygen (O2) and hydrogen peroxide (H2O2). Garg and Manchanda [87] revealed that SOD activity was found to be lower in normal-treated plants as compared to stress-level plants. Therefore, SOD had a significant contribution to the defense system of plants. Catalase (CAT) had the potential to scavenge and eliminate the toxic level of ROS which further reduced the oxidative harm [88]. Moreover, activation of the plant defense system was observed in response to oxidative damage. Hence, elevated SOD, CAT, and POX activities were measured in salt-stressed mango seedlings as compared to non-stressed mango seedlings. Several management approaches have been adopted to enhance the activity of these molecules and purposely over-expressed to control the permeability and ion buildup in mango plants under salt toxicity [89]. Applying NP materials to increase the resilience of plants to abiotic stresses is one of these approaches [90]. Under abiotic stresses, fruit cultivars with a tolerant nature showed greater improvements in antioxidant profiling, whereas sensitive germplasm showed a poor increase in antioxidant profiling [91].

Plants activate their defense system by stimulation of antioxidants within the plant cells and organelles subjected to adverse climatic conditions. Plant metabolism is disturbed due to irregularities in photosynthesis and enzymatic functionality. The regulation of plant metabolic processes is necessary for the activation of antioxidants to scavenge toxic ROS [92]. SOD might eliminate superoxide anions in excess from cells, as proved by the progress of plant defense mechanisms [93]. SOD had a strong chance of balancing O2 from the generation of H2O2. Hence, H2O2 is toxic to plants, as exerted by Dvořák et al. [92]. APX could eliminate the generation of H2O2 [94]. Moreover, APX activation had greater potential to scavenge toxic ROS in plants. Hence, it has been revealed that APX could improve tree tolerance in plants [93]. Fruit trees’ glyoxylic acid-circulating structures and peroxisomes, which effectively convert H2O2 into H2O, are the main locations of CAT. The plants are considered to be more sensitive to H2O2 because CAT significantly lowers H2O2 levels.

The fruit plants had the potential to improve abiotic stresses by the activation of CAT responses across cell divisions. H2O2 generation can be regulated with CAT activity. However, H2O2 is the most reliable indication of abiotic stresses [95]. Moreover, several non-enzymatic activities such as soluble sugars, ascorbic acid, phenols, and tocopherols were reduced under abiotic stresses. ROS regulation is important to reduce oxidative harm (Fig. 2). Fruit tree tolerance is important to improve for higher yield. Ascorbic acid synthesis is effective for improved tolerance levels in fruit crops. The rootstock had a greater contribution to the salinity tolerance of fruit crops by restricting the translocation of excess sodium in plants through roots. Jujube cv. “Gola” was found to be more tolerant due to higher levels of ascorbic acid. Hence, ascorbic acid had a greater contribution to the improvement of salinity tolerance in fruit crops [96].

Figure 2: ZnO-NPs mitigate the adverse effects of abiotic stress on fruit crops focusing on the quality production of fruits

Tocopherol is an integral part of the plant because it can limit lipid radical generation and scavenge harmful ROS. The functioning of antioxidants and non-antioxidants is improved by tocopherol generation. Higher plants are comprised of various tocopherol isomers [97]. However, α-tocopherol had stronger antioxidant properties. Moreover, fruit trees have significant levels of α-tocopherol in their chloroplast envelopes [98]. The oxidation of lipids, carotenoids, and tocopherols had the capability to restore oxidized radicals and limit the extent of the signaling channel [99]. These also contribute to strengthening the defense system of plants subjected to abiotic stresses. Moreover, strong thylakoid membranes had the potential to mitigate the oxidative harms that occur in plants due to adverse climatic conditions.

The aerial plant components, i.e., leaf, pollen, flower, and fruit portions are high in flavonoids. Fruit plants commonly store them as glycosides in their vacuoles. In higher-level plants, they are also effective ROS explorers as revealed by Gadi [100]. Moreover, it has been linked to decreased lipid peroxidation and lipoxygenases. A plant researcher explored that flavonoid (1 mM) was very effective at limiting lipoxygenase in saline circumstances as explored by Potapovich and Kostyuk [101].

6  ZnO-NPs Enhance Shelf Life and Fruit Quality

Fruit quality can be enhanced with supplementation of ZnO-NPs. The practical implications are necessary to explore the fruit productivity to feed a huge population in the future. The customers are interested in fresh-cut fruits because of their novel and natural substances. Fruits’ deceased and living tissues are vulnerable to bacterial and fungal invasions, which results in spontaneous decomposition when the fruit is handled and stored. Fruit browning is attributed to the destruction of phenolic molecules that are o-quinone counterparts, which subsequently form polymers into intricate dark-colored pigments [102]. Several kinds of chemical and physiological remedies are used to retard deterioration and extend the freshness of fruit products. Immobilization of the enzyme is a crucial technique to increase their function and longevity. There are two techniques used to immobilize enzymes: covalent adhesion and physical adsorption. There are disadvantages to physical adsorption, including diminished enzyme activity, encapsulation of the active areas, and unwanted polarity of the molecules. However, the enzyme’s covalent immobilization with the particle surface releases the bound active compounds and also leads to an increase in performance [103].

Its thermal resilience was raised by the covalent adherence of GOx on altered iron oxide nanoparticles. The results of the investigation revealed that the greatest activity of both free and immobilized enzymes was observed at 30°C. However, compared to free enzymes, immobilized enzymes remained more resilient when the temperature was enhanced [104]. Because of their fixed authorization, the enzymes affixed to the surface of the nanoparticle demonstrated greater resilience and durability than their free equivalents (Table 2).

The application of Zn-ONPs for the immobilization of enzymes is an intriguing strategy because it offers several advantages. Akbar et al. [114] revealed ZnO-NPs are used in food processing. Their opinion is that ZnO-NPs have a significant antibacterial impact and are a valuable source of Zn for food supplements, making their usage in the food business both acceptable and desired [114]. In light of the commercial significance of GOx hybrids, researchers conceptualized a novel antibacterial strategy to extend fruit shelf life. The GOx/ZnO-NPs bioconjugate was dissolved in a buffer to develop a spraying solution that was used on peach fruit [115]. The results of the treatment were tracked for 15 days at a temperature of 25°C ± 1. Fruit quality after storage was assessed using common quality indicators such as weight loss, fruit hardness, total soluble solids (TSS), and antioxidant activity in removing free radicals [115].

ZnO-NPs management of abiotic stresses in fruit crops seems to be a practical and long-term strategy that could improve production and quality. However, to fully realize this technology’s promise in real-world farming environments, more study and a thorough grasp of the agronomic as well as environmental variables are required.

7  Conclusion

ZnO-NPs had excellent potential to improve tree yield and fruit quality to combat abiotic stresses by improving the defense system. ZnO-NPs have high reactivity and small size; therefore, fruit crops must strengthen the membranes of plants under adverse climatic conditions. Fruit tree tolerance can be enhanced by supplementation of ZnO-NPs. Moreover, ZnO-NPs supplementation is effective in improving growth, yield, fruit quality, defense system, activation of antioxidants, osmolytes generation, and production of bioactive compounds to improve tree resilient mechanism. However, optimal concentrations of ZnO-NPs can be addressed in the future depending upon the severity of stress, the kind of crops, and their cultivars.

Acknowledgement: None.

Funding Statement: The authors received no specific funding for this study.

Author Contributions: Meryam Manzoor and Konstantin Korolev designed and wrote the paper, Maryam and Riaz Ahmad reviewed and edited the manuscript. All authors reviewed the results and approved the final version of the manuscript.

Availability of Data and Materials: The availability of data is not applicable and all the required data is included within the manuscript.

Ethics Approval: Not applicable.

Conflicts of Interest: The authors declare no conflicts of interest to report regarding the present study.

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