1. Introduction

2. Climacteric fruits and ethylene

• System I of ethylene production is associated with vegetative tissues and fruits in the early stages of development and is characterized by low rates of ethylene production, autocatalytic character (a process by which ethylene induces and controls its own production), and absence of relevant peaks in the production of this phytohormone [30,31].
• On the other hand, system II is present in more advanced development processes, especially in climacteric fruits, and is characterized by high rates of ethylene emission, feedback (higher ethylene concentration in the environment implies higher production of the same), and a high peak of ethylene production at the onset of physiological maturity [30].

4. Regulation of Ethylene Biosynthesis and Signalling

4. Physiological and molecular effects of ethylene on fruit ripening

5. Ethylene managing strategies

5.1. Ethylene Inhibitors

Ethylene inhibitors act at some point in the biosynthesis, signalling, or perception pathways of ethylene as previously developed. By delaying or slowing down these metabolic processes, they hinder the natural progression of product ripening.

5.1.1. 1-Methylcyclopropene (1-MCP)

1-MCP is a chemically synthesized molecule belonging to the family of small-ring hydrocarbons (Figure 4). Under normal environmental conditions (25°C and 1 atm), 1-MCP is gaseous; therefore, in its commercial form, it is encapsulated in γ-cyclodextrins that dissolve with the relative humidity of the environment, releasing this gas [140,141].

This molecule is characterized by having a molecular structure similar to ethylene. This causes it to compete with this phytohormone for the ethylene membrane receptors (ETR1, ERS1, and EIN4) located on the plasma membranes of plant cells, inhibiting the ethylene response [30]. This mechanism can be observed in Figure 5.

Additionally, 1-MCP interacts with ethylene transmembrane receptors in a more intense and rapid manner than ethylene itself, facilitating the binding of 1-MCP to these receptors [142]. Other studies have shown that 1-MCP has other functions besides competing for ethylene reception; however, these mechanisms have not yet been discovered [143]. The effects of applying this molecule have been widely studied in the last 15 years (Table 1).

However, 1-MCP shows lower effectiveness than other ethylene scavengers or deactivators such as palladium compounds or the use of potassium permanganate as ethylene oxidants [30,44]. The reason for this is that 1-MCP is limited to competing with ethylene for cellular receptors, thereby partially preventing its reactivation but not eliminating it. Additionally, 1-MCP does not block all ethylene receptors in plant cells, thus its effectiveness is lower than expected [144].

Table 1. Studies on the application of 1-MCP in different formats and concentrations on the preservation of various fruits.

Table 1. Studies on the application of 1-MCP in different formats and concentrations on the preservation of various fruits.

Format	Concentration	Fruit	Conditions	Significative results	Reference
Gas	0.5 µL L-1	‘Raf’ Tomato	10 °C, 90 % RH, 7 days 20 °C, 90 % RH, 4 days	It reduces both ethylene production and respiration rate and, in turn, delays weight, soluble solids content and total acidity changes.	[145]
Gas	1 µL L-1	Pear cv: ‘Gorham’ ‘Gran Champion’ ‘La France’ ‘Gold La France’	20 °C, 25 days	Reduction of ethylene production, significant delay in the expression of genes related to the change in skin colour of pears and preservation of chlorophyll and fruit firmness.	[146]
Gas Gas	0.1 µL L-1 0.035 µL L-1	‘Unicorn’ Tomato	10 °C, 85 % RH, 15 days	The treatment using a higher concentration of 1-MCP showed a higher conservation of lycopene and weight.	[147]
Gas	0.9 µL L-1	‘Hayward’ Kiwi	20 °C, 95 % RH, 20 days	Inhibition of ethylene production and respiration rate, delay of rot incidence, weight loss, increase of soluble solids content and total bacterial count. Improved preservation of firmness, chlorophyll, total acidity, ascorbic acid and antioxidant capacity.	[148]
Micro-bubbles	100-400 ppb	‘Khai’ Banana	25 °C, 85 % RH, 12 days	Reduced respiration rate and ethylene production. Higher preservation of total chlorophyll content, colour, firmness, total soluble solids, antioxidant capacity and total phenolic compounds.	[149]
Gas	100 nL L-1	‘Gold’ and ‘Rainbow’ Papaya	22 °C, 85 % RH, 25 days 10 °C, 85 % RH, 25 days	Fruits treated with high concentrations of 1-MCP showed increased firmness and delayed colour variation, it menas, delayed ripening. The authors claim that commercial application could lead to a 30 % reduction in papaya ripening.	[150]

5.1.2. Salicylic Acid (SA)

Salicylic acid (SA) (Figure 6) is a phenolic compound found in plants that is currently considered an endogenous plant hormone [151]. This molecule regulates many processes such as stress response, plant development, thermogenesis, photosynthesis, stomatal behaviour, transpiration, ion absorption and transport, seed germination, and glycolysis, among others [152,153].

It acts as an antagonist of ACC oxidase (ACO), inhibiting its action; this enzyme is crucial in the process of ethylene production. ACO oxidizes ACC to ethylene in its final stage [154]. It has been demonstrated that SA inhibits ethylene production in various plants. Leslie and Romani in 1988 [155] observed that applying a concentration of 80 μM of SA during washing of pears (Pyrus communis L.) reduced ethylene production rate by 80% after 24 hours. Srivastava & Dwivedi in 2000 [156] applied 100 μM of SA to bananas, achieving approximately a 50% reduction in respiration rate. These authors also observed a reduction in the activity of cellulase and polygalacturonase enzymes after SA application. Babalar and colleagues [157] applied SA at a concentration of 2 mM, achieving effective reduction of both fungal rot and respiration rate in 'Selva' variety strawberries. In 'Canino' apricots, Elmenofy et al. in 2021 [158] compared the application of two ethylene inhibitors such as SA (4 mM) and AVG (150 mg L-1), recommending the application of SA (4 mM) which yielded optimal yield, quality, and preservation of this apricot variety when grown under Egyptian environmental conditions, where the research was conducted. Lastly, Mansourbahmani and colleagues [44] compared different ethylene removal methods, including SA. These authors concluded that applying a 1% SA solution to 'Valouro' variety tomatoes reduced ethylene production by 25%, which was lower than other investigated methods such as 1-MCP application or ethylene oxidation by the use of palladium, potassium permanganate, or UV-C radiation.

Recently, applications of SA as an elicitor have been discovered. It refers to a substance or signal that triggers a specific response in a cell, organism, or system. In plants, elicitors can be molecules like phytohormones, which activate specific receptors on the surface of cells, leading to a cascade of intracellular signalling events ultimately resulting in a cellular response. Gong and colleagues [159] reported in their review that SA application during fruit formation (lemon, pomegranate, mango, and strawberry among others) improved phenylpropanoid metabolism and carotenoid biosynthesis, increasing accumulation of flavonoids, ascorbic acid, and carotenoids, thereby enhancing antioxidant activity in the subsequently harvested fruit.

5.1.3. Aminoethoxyvinylglycine (AVG)

Similar to SA, aminoethoxyvinylglycine (AVG) (Figure 7) is a compound that inhibits ethylene production by intervening at some point in the metabolic pathways that leads to its production. Specifically, AVG is an inhibitor of ACC synthase (ACS) as it binds to its coenzyme, pyridoxal phosphate (PLP), preventing the conversion of SAM to ACC and thereby affecting the subsequent production of ethylene [160,161,162].

The postharvest application of AVG has been studied in various fruits: Salveit [163] applied a 10 mM solution of AVG in tomatoes (Lycopersicon esculentum Mill.) of the 'Castlemart' variety, resulting in a 30% reduction in respiration rate and a 70% reduction in ethylene production rate of the fruits. Ozturk and colleagues [164] applied 200 mg L-1 of AVG in plums two weeks before harvest in the 'Black Beauty', 'Black Amber', and 'Fiber' plum varieties. Results showed a 60% reduction in ethylene production and respiration rate. They also observed a delay in the colour change from green to red and prevented firmness loss (70% firmer fruits in treated fruits) and weight loss (15% more weight in treated fruits). The same author in 2019 applied 225 mg L-1 of AVG in kiwifruits (Actinia deliciosa L.) of the 'Hayward' variety, observing similar results, but no differences were observed with the control in the analysis of ascorbic acid and total flavonoids with the application of AVG [165]. Similar effects were observed by Yuan & Carbaugh [166] in apples of the 'Golden Supreme' and 'Golden Delicious' varieties, Muñoz-Robredo and colleagues [167] in apricots of the 'Patterson' variety, Xie and colleagues [168] in pears of the 'Starkrimson' variety, Kim and colleagues [169] in plums of the 'Formosa' variety, and Win and colleagues [170] in persimmons of the 'Sangjudungsi' variety.

However, possible negative effects on fruit aroma development have been reported, resulting in loss of intensity [171,172]. Additionally, its cost is too high for industrial application [161].

5.1.4. Aminooxyacetic Acid (AOA)

Unlike the compounds mentioned earlier (1-MCP, SA, and AVG), AOA (Figure 8) is a nonspecific inhibitor of all enzymes that use PLP as a coenzyme, including ACC synthase, so its application could decrease ethylene synthesis [161,173,174]. Due to its nonspecificity, it may interfere with other enzymatic reactions, negatively affecting other physiological processes [175].

Bulantseva and colleagues [176] applied a solution of AOA at 10 mg L-1 to bananas harvested at three different ripening stages (green, green-yellow, bright yellow). Those treated and harvested in the green and green-yellow stages had ethylene production rates about 30% lower than controls at the same ripening stages. Lima and colleagues [177] immersed the roots of sweet potatoes of the 'BRS Rubissol' variety in 1 mg L-1 of AOA, resulting in better preservation of weight, total sugars, pH, and a reduction in sprouting compared to untreated potatoes, thereby extending the shelf life of treated potatoes.

To date, the application of this compound has been mainly limited to the preservation of harvested flowers [178], while its application in fruits and vegetables has not been sufficiently tested.

5.3. Ethylene removal by catalytic oxidation

This method of ethylene elimination relies on chemical oxidation-reduction reactions in which ethylene irreversibly dissociates into CO₂ and water. The effectiveness of this method depends on the oxidizing capacity of the various compounds that can be used for this purpose.

5.3.1. Potassium permanganate (KMnO₄)

Potassium permanganate (see figure 12) acts on the double bond of the ethylene molecule to oxidize it in the presence of water. Ethylene, when oxidized by potassium permanganate, initially forms acetaldehyde, which is subsequently oxidized to acetic acid and then to carbon dioxide and water.

The redox reaction caused by potassium permanganate results in a colour change in the permanganate itself, shifting from purple (MnO₄ ions) to brown (MnO₂). Studies conducted using potassium permanganate as an ethylene remover are quite diverse. Its effects have been tested in various applications involving apples [192], apricots [106], blueberries [193], tomatoes [44,125,194], pears [100,195], and peaches [45,101], among others. Many studies conducted with this molecule aim to enhance ethylene removal by increasing the contact surface of potassium permanganate with this phytohormone. This is achieved by mixing it with inert materials such as zeolites or activated carbon mentioned earlier, compounds that also aid in the absorption and adsorption of ethylene (see Table 2).

Table 2. Some studies on the application of potassium permanganate, in different formats, in the preservation of fruits.

Table 2. Some studies on the application of potassium permanganate, in different formats, in the preservation of fruits.

Format	Concentration	Fruit	Conditions	Significative results	Reference
LDPE Sachet	Saturated and dried KMnO4 solution (50 mL) together with natural clays and activated carbon	‘Pollock’ Avocado	12 °C, 94 % RH, 21 days	Reduction of CO2 and C2H4 concentrations in the packaging. Maintained good visual quality. Inhibited disease incidence and reduced chilling damage. Delayed weight and firmness losses. Soluble solid content and flesh colour were not significantly affected.	[196]
LDPE Sachet	5 g KMnO4 and vermiculite mixture at doses of 1.5 and 1% w/w	‘Musa’ Banana	18 °C, 70-80 % RH, 16 days	Delayed yellowing of the skin. Slower increase in CSS and decrease in AT. Reduced loss of firmness and weight. Minimal increase in the SSC/TA ratio.	[197]
Sachet	Mixture of natural clays and KMnO4 (9 g per kg of fruit)	‘Hayward’ Kiwi	0 °C, 85-95 % RH, 200 days	Delayed firmness. Reduction of the increase in SSC and decrease in TA. Reduction of total chlorophyll and ascorbic acid degradation.	[198]
LDPE Sachet	20 g KMnO4 together with aluminium oxide	‘Nijisseiki’ Pear	0 °C, 90 % RH, 252 days	Delayed yellowing. Decrease in C2H4 levels and respiration rate. Reduced severity of disorders (browning of flesh and heart).	[199]
Sachet	20 % KMnO4 in bentonite zeolite (2:1 w/w)	‘Valouro’ Tomato	7 °C, 90 % RH, 35 days	Reduction of C2H4. Delayed softening of the fruit. Minimisation of the rate of decrease in phenol content and antioxidant activity. Decreased ascorbic content and increased lycopene content. Delayed water weight loss and reduced severity of rotting.	[44]

This molecule has significant industrial application due to its low production cost, ease of incorporation into storage and transportation systems, and high effectiveness in ethylene removal. The main drawback lies in its rapid consumption and potential toxicity upon contact with food. However, it's worth mentioning that this issue has been addressed by presenting it (in most of its commercial forms) in easily replaceable sachets when they become saturated.

5.3.2. UV Radiation

Short-wave UV radiation between 100 and 280 nm (UV-C) is also capable of acting on the double bond of ethylene, generating oxidizing radicals that ultimately produce CO₂ and water in the reaction process, a phenomenon known as photocatalysis [200,201]. Additionally, this radiation can produce small amounts of ozone at these wavelengths, a molecule that also acts on ethylene, as will be discussed in the following points.

Bu and colleagues [202] applied UV-C radiation at 254 nm to tomatoes of the 'Zhenzhu' variety. Despite the direct impact of UV radiation on the fruit potentially causing adverse effects, these researchers observed a significant 20% reduction in the ethylene production rate. Mabusela and colleagues [203] in their review article studied ultraviolet radiation at 254 nm and 185 nm in a vacuum environment. At both wavelengths, ethylene elimination was observed through two pathways: firstly, the direct action of UV-C radiation on the double bond of ethylene, and secondly by producing ROS by reacting with water and oxygen from the environment (considering that the relative humidity of fruit storage environments shows values around 90% and 21% for oxygen). These ROS then react with ethylene, producing CO2 and water (Figure 11).

The effectiveness of the treatments studied by Mabusela and colleagues [203] depends on numerous factors that greatly influence the ethylene removal rate. One example is relative humidity. Water is the main source of ROS when attacked by radiation; therefore, low relative humidity significantly affects ethylene removal using this method. Currently, there is no consensus on the performance of this ethylene oxidation technique.

Additionally, UV-C radiation has been extensively studied for its ability to eliminate spores of microorganisms present in fruit storage environments [204]. This is an effect that must be considered, as fungal and bacterial diseases, among many other effects, increase the ethylene production rate in infected fruits [203].

This method has numerous advantages, such as ease of implementation, low energy consumption, effective ethylene removal under optimal conditions, and the elimination of spores, minimizing fungal and bacterial infections. However, it has several disadvantages that, if not considered, would reduce its effectiveness:

• Relative humidity: as mentioned earlier, environmental water is the main source of certain ROS, crucial for the efficacy of this method. The lower the relative humidity, the lower the ethylene removal achievable with UV-C radiation [205].
• Initial ethylene concentration: increasing the initial concentration of ethylene, while keeping the photon energy constant, reduces the energy received by a greater number of ethylene molecules, thus reducing ethylene removal [137,206].
• Oxygen concentration: oxygen is the precursor of certain ROS and ozone that facilitate this process. A low concentration of this gas will hinder ethylene removal [206].
• Direct incidence on fruits: when UV-C radiation is directly targeted at fruits, it causes structural changes that negatively affect their quality [203].

5.3.3. Titanium Dioxide (TiO₂)

Photocatalysis is another approach to eliminate ethylene [17,200,207]. Titanium dioxide (TiO2) is a semiconductor material and one of the most frequently employed photocatalysts for ethylene degradation, a capacity primarily attributed to its unique photochemical reactivity and physical properties, which include high brightness (due to high refractive index) and resistance to discoloration [30].

The complete mechanism of organic compound oxidation by UV light radiation on titanium dioxide is described by Pathak and colleagues [206]. In summary, the reaction is based on the irradiation with UV wavelengths (hν) (around 240-380 nm) which causes titanium dioxide to generate electrons (e-) and other effects (reaction 1). These, when acting on the water in the environment, produce highly reactive hydroxyl radicals (·OH) (reaction 2). These radicals then react with organic compounds, resulting in CO₂ and water (reaction 3) [200,201].

-

Reaction 1: TiO₂ + hν (UV energy/radiation) → TiO₂ + e- + h+

-

Reaction 2: H₂O + h+ → ·OH + H+

-

Reaction 3: ·OH + Organic compound (ethylene) → CO₂ + H₂O

Although the mechanism of photocatalytic oxidation has been described [206], the exact mechanism of reaction with ethylene remains a topic of debate due to the presence of various reaction intermediates that have not been fully elucidated [137].

Numerous studies have been conducted on the application of this ethylene elimination technology. Alves and colleagues [208] applied it to cherry tomatoes stored at 18°C with a relative humidity of 85%. The treatment involved passing a constant airflow through the titanium dioxide and UV light system. During the storage period, cherry tomatoes treated with titanium dioxide irradiated with UV light showed a lower concentration of ethylene, lower respiration rate, lower total soluble solids content, and higher concentrations of lycopene and titratable acidity compared to the control, demonstrating that fruits treated with photocatalysis did not reach full ripeness. Li and colleagues [209] obtained similar results using the same tomato variety for their study.

In recent years, many studies on the effect of titanium dioxide-UV on ethylene have focused on its application as a component of films or fruit packaging. Among them, Fonseca and colleagues [210] evaluated the efficacy of polyethylene foam nets coated with a photocatalytic nanocomposite composed of gelatine and titanium dioxide for degrading the ethylene produced by papayas (Carica papaya L.) of the 'Golden' variety. The fruits treated with gelatine-titanium dioxide and irradiated with UV light showed a 60% lower ethylene accumulation than the fruit in the control group after four days of storage, as well as higher firmness and lower ripening index (soluble solids/titratable acidity ratio). Ghosh and colleagues [211] applied photoactivated titanium dioxide and chitosan to form a film to cover peppers of the 'Tejaswani' variety stored at room temperature (25°C). The treated peppers-maintained CSS along with higher firmness, cell integrity, hydration, and colour. De Chiara and colleagues [212] applied a powder composed of titanium dioxide and silicon dioxide in different proportions for the preservation of 'Camone' tomatoes. In tomatoes treated with the 80-20 ratio (TiO2-SiO2), ethylene was completely degraded after 14 days of study. However, the fruits in this treatment did not reach the red colour stage.

Titanium dioxide is categorized as a food additive by the U.S. Food and Drug Administration under code 21CFR73.575 and is inexpensive. This makes it cost-effective for application in fruit storage and transportation. However, there are some drawbacks to consider for its implementation:

• It requires very controlled storage conditions as the effectiveness of titanium dioxide depends on numerous variables such as humidity, temperature, or UV light intensity [137,174,213].
• It is not a selective method, as seen in the summary of the reaction triggered by titanium dioxide and UV radiation. In addition to ethylene, it acts on other organic compounds such as aromatic compounds, which can affect the organoleptic quality of the fruits [214].
• It can increase the storage temperature, which may lead to fruit damage [215].
• If UV radiation, essential for this method to be effective, directly impacts the fruits, it can lead to negative consequences such as loss of aromas, rupture of cell membranes, and degradation of structures [216].

5.3.4. Ozone (O₃)

Ozone, typically produced with UV lamps, is a strongly oxidizing molecule that can also be used as an ethylene eliminator [200]. Ozone, being a highly reactive gas, acts on the double bond of ethylene, breaking it and dissociating this phytohormone into CO₂ and water according to the following reaction: O₃ + C₂H₄ → CO₂ + H₂O The application of this method is straightforward. It involves passing a flow of air from fruit storage facilities (rich in ethylene) through an ozone generator and returning it to the warehouse free of ethylene [217]. Liang and colleagues [218] analysed ethylene production, firmness, vitamin C content, malondialdehyde content, levels of ascorbate peroxidase, peroxidase, and aromatic compound contents of 'Hard Pink' tomatoes stored for 25 days at 10°C and treated with 17.14 mg (m3)-1 of ozone for 1 hour at the start of the study. The results showed that ozone treatment delayed the loss of firmness, bioactive compounds, and aromatic compounds, as well as reducing ethylene production rate by 50% during the first 15 days. Toti and colleagues [219] treated 'Cantaloupe' melons with 0.3 ppm of ozone at 6°C for 13 days. Their results indicated that ozone treatment reduced the activity of cell wall enzymes (α-arabinopyranosidase, β-galactopyranosidase, and polygalacturonase) responsible for their degradation and hence softening of melons. Ethylene concentration was also reduced. With all this, the quality of the treated melons was maintained for a longer period. More recently, Gao and colleagues [220] treated 'Muscat Hamburg' grapes, despite being non-climacteric fruits, with 14.98 mg (m3)-1 of ozone. The results revealed a reduction in ethylene production close to 30% in the fruits subjected to the treatment. Additionally, there was maintenance of antioxidant capacity with higher concentrations of enzymes such as catalase (CAT), peroxidase (POD), superoxide dismutase (SOD), and polyphenol oxidase (PPO), capable of removing reactive oxygen species (ROS) from plant tissues. Furthermore, a lower incidence of fungal diseases was observed in the treated fruits. Although considered safe by the FDA (U.S. Food and Drug Administration) and not complicated for industrial implementation, ozone presents several drawbacks:

• Handling is very challenging due to its easy decomposition into oxygen [139].
• If certain levels are exceeded, ozone shifts from offering significant improvements to being harmful as it can destroy tissues, leading to wounds that can promote ethylene production due to stress [200].
• Even though it is safe at low concentrations, ozone begins to have harmful effects on human health from 5 ppm onwards, including vision problems, sensation of asphyxia, headaches, pulmonary oedema, and coma when concentrations reach 50 ppm [221].

5.3.5. Palladium

Palladium is a scarce metal with numerous applications, including its use as a catalyst in chemical reactions and in medicine for surgical material production. It also has the ability to absorb hydrogen into its metallic structure and purify hydrocarbons in the automotive industry. This latter application prompted its use as a potential ethylene eliminator through catalysis. Similar to other ethylene oxidants, palladium irreversibly decomposes ethylene into CO2 and water.

Bailén and colleagues [213] employed a mixture of granulated (20-60 mesh) activated carbon with 1% palladium on three tomato varieties, 'Beef', 'Mendez', and 'Raf'. Within the first 50 hours, nearly 60% ethylene elimination was achieved in all three varieties, along with better preservation of firmness and colour, both internally and externally. Mok and colleagues [222] applied a mixture of ZSM-5 zeolite and 0.36% palladium in the conservation of 'Fuji' apples, introduced into quartz tubes through which air from the storage chamber passed. The results showed nearly 100% ethylene elimination after 40 days of storage. Tzeng and colleagues [223] observed a 99% ethylene removal in banana preservation using a similar mixture and application method.

Mansourbahmani and colleagues [44] employed palladium with nano zeolites in various concentrations (0%, 1%, 2.5%, and 5%) and compared this ethylene removal method with other ethylene eliminators or inhibitors such as potassium permanganate, 1-MCP, and UV-C, concluding that the palladium method was the most effective.

Although this method does not have many drawbacks, its industrial application proves to be very complicated due to palladium being a very scarce and difficult-to-extract metal, and consequently extremely expensive. Additionally, the country with the highest production of this element worldwide is the Russian Federation, which is currently subject to numerous economic and commercial sanctions (due to the invasion of Ukraine), further increasing its price.

6. Practical implications and future directions

7. Conclusions

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