Alcoholic Off-Flavor Disorders in Fresh Fruits

Ali, Maratab; Batool, Sara; Khalid, Nauman; Manzoor, Muhammad Faisal; Zhao, Xiuming; Li, Xiaoan; Li, Fujun; Xinhua, Zhang; Aadil, Rana Muhammad

doi:https://doi.org/10.1155/2023/3959653

Journal of Food Biochemistry

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Abstract Introduction Conclusion Data Availability Conflicts of Interest Authors’ Contributions Acknowledgments References Copyright Related Articles

Review Article | Open Access

Volume 2023 | Article ID 3959653 | https://doi.org/10.1155/2023/3959653

Alcoholic Off-Flavor Disorders in Fresh Fruits

Maratab Ali,^1,2Sara Batool,³Nauman Khalid,²Muhammad Faisal Manzoor,^4,5Xiuming Zhao,¹Xiaoan Li,¹Fujun Li,¹Zhang Xinhua,¹and Rana Muhammad Aadil⁶

Academic Editor: Ravi Pandiselvam

Received05 Apr 2023

Revised22 May 2023

Accepted24 Jun 2023

Published07 Jul 2023

Abstract

The off-flavor disorder is one of the most quality deteriorating and undesirable postharvest physiological disorders in fresh fruits. Over-biosynthesis and accumulation of ethanol metabolism-related metabolites such as acetaldehyde and ethanol have been associated with an alcoholic off-flavor disorder in various fresh fruits at both pre- and postharvest stages. Correspondingly, many studies have reported the association of such off-flavor disorders with several inducing factors, including anaerobic respiration, low O₂ stress, high CO₂ stress, and storage temperature stress, that upregulate ethanol metabolism in fruits. Furthermore, numerous metabolic and physiological mechanisms that govern ethanol metabolism under such factors are subsequently addressed. These mechanisms include the γ-aminobutyric acid (GABA) shunt pathway, mitochondrial energy metabolism, glycolysis, Krebs or TCA cycle, cytosolic malate metabolism, and starch and sugar metabolism. To summarize the relevant findings, the current paper reviews the literature on alcoholic off-flavor disorder, focusing on the role of significant underlying causes and key metabolic and physiological mechanisms in boosting ethanol metabolism in fresh fruits. In addition, recent measures that have been already taken or are in progress to control the higher activity of ethanol metabolism that may eventually result in limiting the alcoholic off-flavor disorder in harvested fresh fruits have also been discussed. Moreover, functions of metabolic mechanisms, including respiratory mechanisms, and other factors such as fruit genetic makeup, degree of maturity, and postharvest handling and storage conditions, are needed to be investigated in future work at both physiological and transcriptomics levels to reveal the additional relation to alcoholic off-flavor disorders in fresh fruits during ripening and storage.

1. Introduction

Fruits are one of the most desired horticulture commodities among consumers due to their high nutritional content, appealing color, and flavor [1–5]. However, due to a fluctuating ripening trend, most fresh fruits ripen rapidly after harvest, even at ambient and low temperatures. In turn, substantial-quality losses might occur in fruits, such as early fruit softening, decay incidence, weight loss, color degradation, flesh browning, skin pitting, nutritional breakdown, chilling injury, and alcoholic off-flavor development [6–8]. Such quality losses arise due to the ongoing metabolic adaptations that alter the intended qualities of fruits until they are unmarketable.

The perception of off-flavor sensations characterizes alcoholic off-flavor disorder as a result of an excess of fermentation metabolites in ethanol metabolism, such as acetaldehyde and ethanol [6, 9–11]. Acetaldehyde and ethanol concentrations in fruits are often present in trace amounts throughout normal ripening or postharvest storage but begin accumulating as fruits tend to ripen sharply under high respiration rates, elevated temperatures, and anaerobic respiration [12]. Such metabolites are involved in maintaining the postharvest quality of the fruit under aerobic respiration, notably during the volatile aroma production stage [13]. Also, they can improve fruit flavor, suppressing ethylene production and reducing fruit firmness [14]. Conversely, increased levels of these metabolites develop physiological disorders in fruit during postharvest storage as their accumulation may impart the formation of off-flavors in various horticultural crops such as mandarins, pear, dragon fruit, grapes, and kiwifruit.

Several studies have shown that anaerobic respiration [15], low O₂ stress [16], high CO₂ stress [17], and high-temperature stress [18] have a detrimental impact on ethanol metabolism, which leads inevitably to over biosynthesis or accumulation of acetaldehyde and ethanol, resulting in alcoholic off-flavor disorder in fruits. Subsequently, numerous metabolic physiological mechanisms, including the γ-aminobutyric acid (GABA) shunt pathway [6], mitochondrial energy metabolism [19], glycolysis and Krebs (TCA) cycle [12], cytosolic malate metabolism, and starch and sugar metabolism [20, 21], tend to alter or regulate the ethanol metabolism under the consequences of such stresses. The detrimental impact of undesirable modifications in such metabolic physiological pathways in response to stress stimuli, specifically accelerated ethanol metabolism, has been demonstrated by various studies.

To date, as per our knowledge, no review article on the alcoholic off-flavor disorder in fresh fruits has been published. Hence, this review article aims to discuss the comprehensive literature regarding alcoholic off-flavor disorders in fresh fruits. This article particularly addressed the primary causing factors involving physiological and metabolic mechanisms and postharvest strategies to overcome alcoholic off-flavor disorders at an agroindustrial level. Furthermore, a research gap that needs to be investigated at a molecular level in future research has also been pointed out.

2. Ethanol Biosynthesis and Metabolism

Ethanol is an essential aroma-active volatile compound naturally synthesized in fruits and vegetables during the maturing and ripening processes via ethanol metabolism [22]. Ethanol metabolism is a two-step pathway involving the decarboxylation of pyruvate into acetaldehyde and ethanol via the activities of pyruvate decarboxylase (PDC) and alcohol dehydrogenase (ADH), as depicted schematically in Figure 1. Furthermore, acetaldehyde could be further reduced to acetic acid via aldehyde dehydrogenase (ALDH) activity, and ethanol could be converted to ethyl acetate via an esterification reaction catalyzed by alcohol acetyltransferase (AAT) activity using acetyl-CoA as a key substrate [6, 23, 24]. Ethanol metabolism is concomitant with the continuous recycling and reoxidation of nicotinamide adenine dinucleotide + hydrogen (NADH) to nicotinamide adenine dinucleotide (NAD⁺) to generate adenosine triphosphate (ATP) [25–27]. Studies reported that ethanol metabolism is the only alternate pathway for ATP biosynthesis to fulfill the plant’s metabolic energy demand under hypoxic or anoxic stresses [12, 19].

Additionally, accelerated ethanol metabolism leads to over-accumulation and the induction of alcoholic off-flavor, which deteriorate the fruit flavor quality [19]. It has been reported that the high stimulation of ethanol metabolism is anticipated by cellular pH-stat. Pyruvate is catalyzed by lactate dehydrogenase (LDH) to lactic acid, and lactic acid production acidified cytoplasmic pH and consequently activates the first ethanol metabolism enzyme PDC [25, 27, 28]. The intensity of ethanol metabolism can vary based on PDC and ADH enzyme activities, resulting in desired or immature flavor development in various horticultural crops. For example, Botondi et al. [29] and Huan et al. [19] reported the correlation between excess ethanol and acetaldehyde production or accumulation due to enhanced activities of PDC and ADH, resulting in alcoholic off-flavor disorder in “Hayward” and “Bruno” kiwifruit.

Similarly, Zhang and Watkins [30] reported the occurrence of off-flavors compounds such as acetaldehyde and ethanol and enhanced PDC and ADH activity in strawberry fruit. Furthermore, Shi et al. [31] noticed enhanced acetaldehyde and ethanol contents due to upregulated PDC and ADH transcriptional levels in mandarin fruit. Our previous work also demonstrated that higher PDC and ADH activities result in higher acetaldehyde and ethanol accumulation, which leads to alcoholic off-flavor disorder in “Bruno” kiwifruit at room temperature storage [6, 12, 32].

In contrast, some previous studies negatively correlated PDC and ADH activities with acetaldehyde and ethanol biosynthesis. For example, Ponce-Valadez and Watkins [33] reported that the accumulation of acetaldehyde and ethanol negatively correlated with changes in PDC and ADH activities in “Jewel” and “Cavendish” strawberry fruit. Imahori et al. [34] reported that increased activity of PDC and ADH in bell pepper fruit was not correlated with acetaldehyde and ethanol accumulation. Our previous study concluded a negative correlation between kiwifruit fermentation metabolites and enzymatic activities [32]. Such a correlation could be based on the sensitivity range of each crop’s PDC and ADH expression under various stresses [35, 36].

2.1. Factors Leading to Upregulated Ethanol Metabolism in Fresh Fruits

The development of alcoholic off-flavor is limited during normal fruit growth or ripening, even at pre- and postharvest stages. Meanwhile, due to the upregulation of ethanol metabolism, fruits may be susceptible to the occurrence of alcoholic off-flavor, which may result in the deterioration of fruit flavor quality initiated by the following inducing factors:

2.2. Anaerobic Respiration

Anaerobic respiration is a process that initiates under the effect of reduced aerobic mitochondrial activity and lower ATP biosynthesis as a result of undesired stresses, activating ethanol metabolism to regenerate ATP by using NADH [37, 38]. Plenty of ethanol contents are produced and accumulated, resulting in alcoholic off-flavor development in various fruits [12, 19, 29, 39].

Moreover, a relationship between anaerobic respiration and alcoholic off-flavor disorder is described in detail by Saltveit [15] as follows: “Anaerobic respiration is associated with the regeneration of NAD⁺ and ATP to maintain the plant cell energy demand by moving the pyruvate flux to the ethanol fermentation/metabolism instead of the TCA cycle” (Figure 2). This process further leads to the accumulation of lactic acid in the lactate fermentation pathway, which results in the acidification of cells and activates the PDC in ethanol metabolism. PDC converts the pyruvate to acetaldehyde by removing the CO₂ and then to ethanol through the activity of the ADH to generate more NAD⁺ content. The anaerobic pathway accounted for only 20% of the energy synthesis ability, which is insufficient for plant survival. This fact upregulated ethanol metabolism for more ATP synthesis, resulting in an over-accumulation of ethanol and alcoholic off-flavor.

2.3. Low O₂ Stress

Oxygen concentration in fruit tissues is primarily based on each cultivar’s respiration rate, as the ratio of O₂ and CO₂ fluctuates continuously due to organic compounds’ consistent biochemical reactions and degradation [40]. The correlation between various physiological disorders in many horticultural crops was investigated by applying different modified gas conditions during storage. However, exposure of each cultivar to different concentrations of O₂ atmosphere can be beneficial or harmful depending on various factors such as storage temperature and cultivar type. Oxygen acts as an electron acceptor in mitochondrial cytochrome oxidase during aerobic pathways. However, in a scenario of a limited O₂ atmosphere, plant cells are exposed to fermentation pathways, energy deficits, and storage disorders [17, 41]. Reduced O₂ concentration appeared to have a beneficial relationship with decreased respiration rate, senescence, ethylene synthesis, and enzymatic oxidation at a certain level. In contrast, lowered O₂ induced physiological disorders such as alcoholic off-flavor, gummosis, fruit rots, fruit discoloration, superficial scald, split and shattered pits, gel breakdown, greasiness, and pit burning in many fruits, including pear, apple, mango, and stone fruits (nectarines, plum, cherry, apricot, and peach) during storage [16, 42–44].

The oxygen limit at which ethanol metabolism initiated and accumulation occurred was named the pasture point, or lower oxygen limit, and later referred to as fermentation induction point [16]. Recently, Park et al. [45] reported that the upregulated expression of ethanol metabolism-related genes MdPDC2 and MdADH1 led to higher production of acetaldehyde and ethanol in “Empire” apples during storage in a 0.5 kPa O₂ atmosphere compared to a 1.5 kPa O₂ atmosphere. Burdon et al. [46] reported increased PDC and ADH activities, resulting in a sharp accumulation of fermentation metabolites in the “Hass” avocado under the storage atmosphere with less than 0.5% O₂ concentration. Each crop has a specific tolerance limit for low O₂ levels, and below that, plants adopt a fermentation pathway [47, 48]. Previously, many researchers exposed various fruits to low O₂ atmosphere for a short period and concluded beneficial effects regarding quality. Meanwhile, susceptibility to alcoholic off-flavor disorder has been observed with long-term exposure [16, 19, 49, 50].

Regarding alcoholic off-flavor disorder in fruits under low O₂ stress, Wood et al. [51] reported that exposing fruits to anoxic conditions (20.9 kPa O₂) results in a higher concentration of acetaldehyde and ethanol in apple fruit. Ntsoane et al. [43] concluded that acetaldehyde and ethanol accumulation were significantly higher in “Shelly” mango stored at 5% O₂ than at 10% O₂ in controlled atmosphere storage. Moreover, Pintó et al. [52] concluded that the change in pomegranate flavor quality is caused by the accumulation of ethanol and acetaldehyde due to a low O₂ storage atmosphere. Under low O₂ stress, acetaldehyde is produced by the decarboxylation of pyruvate by PDC and then converted to ethanol by ADH by consuming NADH. Imahori et al. [53] reported that elevated PDC and ADH transcription and translation resulted in the synthesis of new mRNA and PDC and ADH protein in a low O₂ atmosphere. Thus, under low O₂ stress, activation of PDC and ADH is a critical response. Another effect of low O₂ stress is the accumulation of lactic acid that lowers cytoplasmic pH and inhibits LDH activity. This fact further activates PDC and ADH, resulting in excessive ethanol biosynthesis, as discussed above. Boeckx et al. [23] and Botondi et al. [29] reported the accumulation of acetaldehyde and ethanol and the occurrence of alcoholic off-flavor in “Jonagold” apple and “Hayward” kiwifruit under the storage of ultra-low oxygen atmosphere. Ethanol accumulation has been reported as a low-O₂ stress biomarker during horticultural crop storage [40].

2.4. High CO₂ Stress

Optimal ranges of CO₂ levels in fruit tissue or the storage environment may regulate fruit quality [53]. However, each crop’s sensitivity to elevated CO₂ injury varied according to genetic variation, resulting in different responses [54, 55]. For example, the storage atmosphere containing 20–30% CO₂ concentration resulted in maintaining the fruit firmness, ascorbic acid regulation, and titratable acidity level without deteriorating the flavor quality in several sweet cherry cultivars such as “Lapis,” “Van,” “Kristin,” “Stella,” “Sam,” and “Huldra” [56, 57]. On the other hand, Cozzolino et al. [58] described the occurrence of off-flavor in sweet cherry cv. “Ferrovia” during storage based on the accumulation of fermentation metabolites under a 20% CO₂ storage atmosphere. The beneficial effects of increased CO₂ may inhibit respiration rate, ethylene biosynthesis, and N-assimilation in crop tissues, even while promoting carbohydrate synthesis [59–61]. In contrast, Lu et al. [62] reported the occurrence of an alcoholic off-flavor in satsuma mandarin fruit during storage in a modified atmosphere accompanied by a 15% CO₂ concentration. Harman and McDonald [63] also reported the effect of a storage atmosphere consisting of 14% CO₂ concentration on the “Hayward” kiwifruit for 16 weeks and noticed the development of the alcoholic off-flavor disorder. Surprisingly, the optimum flavor was retained when fruit was stored at a lower CO₂ concentration during storage. The off-flavor sensation was identified due to ethanol over-accumulation in “Bartlett” pears, tomatoes, and lettuce stored in an elevated CO₂ storage atmosphere [64–68]. The mango cultivar “R2E2” was similarly reported to have an alcoholic off-flavor disorder after being stored under 8% CO₂ rather than 6% CO₂ during the controlled storage atmosphere [69]. Similarly, the storage atmosphere of 10 kPa CO₂ produced less ethanol and acetaldehyde in mango fruit than the storage atmosphere of 25 kPa CO₂ [70]. Wang et al. [71] evaluated the effect of modified atmosphere packing on the ethanol content of two sweet cherry varieties, “Lapins” and “Skeena,” resulting in off-flavor disorder due to ethanol buildup in the 10% CO₂ storage atmosphere rather than the 8% CO₂ storage atmosphere. Forney et al. [72] reported the accumulation of ethanol in several blueberry cultivars (such as “Duke,” “Aurora,” “Brigitta,” “Jersey,” and “Liberty”) under the storage condition of 25 kPa CO₂. The same findings were reported in the “Ottomanit” fig subjected to an elevated CO₂ atmosphere [73]. Alcoholic off-flavor was observed in the “Italia” table grapes stored in a high CO₂ (>20%) atmosphere [9]. Furthermore, increased expression of the ethanol metabolism-related gene VvADH was associated with increased accumulation of off-flavor aroma volatiles, including ethanol, in table grapes during storage under higher CO₂ conditions [74]. Overall, elevated CO₂ levels have mostly been associated with the deterioration of crop quality attributes, particularly flavor balance and sensory quality [75–79]. The overall impacts of O₂ and CO₂ stress conditions on ethanol metabolism-related metabolites, enzymes, and genes transcriptions in fresh fruits are summarized in Table 1.

2.5. Storage Temperature Stress

An optimal storage temperature range is critical to maintaining fresh fruit quality and extending shelf life by reducing pathological and physiological deterioration [80]. The ideal temperature for each crop’s storage varied depending on its origin or ripening behavior. Kader and Yahia [37] recognized the optimal temperature range of 20–24°C for the anticipated ripening of most harvested fruits. However, some fruits experienced alcoholic off-flavor disorder even at such temperature range (Table 2) along with others disorders such as freezing injury, heat injury, chilling injury, skin/flesh discoloration, softening, and AsA degradation [84, 85].

Studies were conducted on the effect of varied temperature environments on the alteration of volatile aromatic compounds in several fresh fruits. For example, Saberi et al. [81] noticed the alcoholic off-flavor in “Valencia” oranges at 20°C than 5°C during storage. Ali et al. [6] reported the alcoholic off-flavor occurrence in “Bruno” kiwifruit during storage at 24°C. Obenland et al. [82] reported a rapid decline in the sensory quality of “W. Murcott Afourer’s” mandarins due to an excess of alcohols and ethyl esters, resulting in alcoholic off-flavor disorder at 20°C compared to 5°C and 10°C. Similarly, Yang et al. [18] also concluded that high temperatures have a deteriorative effect, particularly due to the enhanced biosynthesis of volatile aromatic compounds (primarily in ethanol) in banana fruit during storage at 30°C than 20°C. Furthermore, Petracek et al. [83] likewise reported an increase in ethanol content in sweet cherry (Prunus avium L., cv. “Sams”) during storage at 20°C compared to storage at 5°C, 10°C, and 15°C, resulting in alcoholic off-flavor disorder. On the other hand, Zou et al. [86] demonstrated that lower storage temperatures ranging from 4°C to 10°C significantly impacted the transcript expression of genes ADH2, PDC1-like1, and PDC1-like2 in tomato fruit. In addition, the effects of storage temperature either on the accumulation or reduction of off-flavor compounds in a variety of fruits have been reported, including “Hort16A” kiwifruit [87], cantaloupe [88], strawberry [89], tomatoes [90], oranges [91], and nectarine [92].

3. Role of Physiological Mechanisms in Regulating Alcoholic Off-Flavor in Fresh Fruits

3.1. GABA Shunt Pathway

The GABA shunt pathway regulates many physiological mechanisms such as C : N balance, pH-stat, plant development, plant defense against insects, signal transduction, and osmoregulation [6, 93]. A short pathway generally metabolizes GABA via the TCA cycle that includes glutamate decarboxylase (GAD), GABA-transaminase (GABA-T), and succinate semialdehyde dehydrogenase (SSADH), called the GABA shunt pathway [94, 95], as depicted in Figure 3(a). So apart from GABA beneficial roles, its accumulation under stress is linked to various postharvest physiological disorders, including the incidence of pear fruit core breakdown [96], pear flesh and spathe browning [97, 98], reduced chilling tolerance in Natura zucchini fruit [99], and controlled atmosphere-related injury in “Honeycrisp” apples [100]. Enhanced GABA levels have also been linked to tomato surface pitting and soggy breakdown in “Honeycrisp” apples, respectively [101, 102].

(a)

(b)

(c)

(d)

Regarding alcoholic off-flavor disorder, Deewatthanawong et al. [103] found that lower GABA-T activity under high CO₂ storage conditions induced the synthesis and accumulation of alcoholic off-flavor compound ethanol along with GABA accumulation as a stress marker in response to a high CO₂ environment in the “jewel” strawberry cultivar. Moreover, Mae et al. [104] also reported an increase in GABA levels due to upregulated GAD and limited GABA-T gene expressions in tomatoes stored in a hypoxic atmosphere. According to the preceding discussion, the relationship between GABA accumulation is mostly similar to the results of low O₂/high CO₂ or anaerobic respiration, which resulted in an ATP deficit and reduced aerobic metabolic pathways. Interestingly, Deewatthanawong and Watkins [101] justified the interlinking of GABA enhancement with the accumulation of fermentation metabolites. Our previous research concluded that the GABA shunt pathway might initiate ethanol metabolism, thus possibly causing the alcoholic off-flavor disorder in kiwifruit cv. “Bruno” [6]. In Figures 3(b)–3(d), it has been shown that inhibited activity of the GABA-T enzyme from 12 days (d) to onward in control kiwifruit cv. “Bruno” appears to be directly linked to limited decarboxylation of GABA and thus shows a sign of suppressed GABA shunt pathway that impacts the functionality of the TCA cycle and thus leads to activation of PDC and ADH, resulting in oversynthesis or accumulation of acetaldehyde and ethanol contents. Our previous study noticed this relationship by using 1-methylcyclopropene (1-MCP) to reduce ethanol metabolism by regulating the GABA shunt pathway under reduced climacteric respiration, avoiding the occurrence of alcoholic off-flavor disorder in kiwifruit during storage [6]. However, limited research has been conducted to investigate the GABA shunt pathway concerning ethanol fermentation, suggesting the need for an integrated approach to analyzing GABA shunt pathway-related gene expression, protein level, and involved metabolites at transcriptomic, proteomics, and metabolomics levels further to reveal the GABA shunt influence on ethanol fermentation.

3.2. Mitochondrial Energy Metabolism

The cell mitochondria are a primary site for maintaining the ATP or energy status pool required for regulating the plant’s physiological or biochemical reactions [105, 106]. The mitochondrial energy status is closely related to the activities of energy metabolism-related enzymes, including succinic dehydrogenase (SDH), cytochrome C oxidase (CCO), H⁺-adenosine triphosphatase (H⁺-ATPase), and Ca²⁺-adenosine triphosphatase (Ca²⁺-ATPase) [12]. Previously, decreased ATP and energy charge (EC) in various fruits were adversely associated with several physiological disorders i.e., chilling injury [107], tissue, pericarp, or peel browning [108], skin pitting [109], and ethanol accumulation or alcoholic off-flavor disorder [12, 19, 95]. An energy deficit occurs due to dysfunctional mitochondrial energy metabolism at the cellular level [19, 77].

In agreement, our previous study concluded a possible correlation between dysfunctional mitochondrial energy metabolism with reduced SDH, CCO, H⁺-ATPase, and Ca²⁺-ATPase activities with upregulated ethanol metabolism [12]. Moreover, Cukrov et al. [39] also reported the occurrence of alcoholic off-flavor in kiwifruit due to downregulated mitochondrial energy metabolism during storage. Consequently, Zhang et al. [110]; Huan et al. [19]; and Blanch et al. [77] further coincided with the susceptibility of alcoholic off-flavor disorder incidence under the same circumstances in strawberry and kiwifruit during storage.

3.3. Glycolysis and TCA Cycle

Glycolysis and the TCA cycle are the central respiratory pathways in horticultural crops. These pathways are involved in maintaining the ripening or senescence processes by regulating the energy status, carbon flux, NAD(P), and NAD(P)H levels through sequential cell reactions [111]. However, research has revealed that such respiratory pathways exhibit considerable variation under different conditions. The glycolysis ended up with pyruvate formation that acts as a primary substrate for the first enzyme of the TCA cycle, pyruvate dehydrogenase (PDH), and ethanol fermentation (PDC) [11, 12]. Meanwhile, the concentration and fate of pyruvate either in the TCA cycle or ethanol fermentation are profoundly influenced by a high CO₂/low O₂ atmosphere or anaerobic metabolism. For example, Ummarat et al. [11] concluded that the off-flavor sensation caused by the over-accumulation of ethanol correlated with enhanced pyruvate content in “Pixie” mandarin during an anaerobic atmosphere generated by waxing. Interestingly, it has been reported that enhanced pyruvate is preferred for alcoholic fermentation over the TCA cycle in anaerobic respiration [112]. Mannucci et al. [113] reported that anaerobic metabolism eventually converts pyruvate to acetaldehyde and ethanol, suggesting that pyruvate and ethanol contents are critically interlinked under stress conditions.

The increase or accumulation of glycolytic flux is interlinked with the enhanced activity of pyruvate kinases (PKs) or suppressed activities of TCA cycle enzymes that are eventually indicated by the accumulation of TCA cycle organic acid [12, 21, 114]. The functional TCA cycle is vital in generating a vast pool of ATP for regulating the cell metabolic mechanisms via oxidative phosphorylation or electron transport chains to maintain the desired ripening processes of horticultural crops [115–117]. Meanwhile, the TCA cycle has been reported to be replaced by glycolysis, which directs pyruvate to ethanol metabolism to generate sufficient ATP to keep cell functions alive under hypoxic conditions [12, 118, 119]. A number of previous research reported that ethanol metabolism occurs under stressed conditions as a result of a malfunctioned TCA cycle, which is associated with suppressed PDH, SDH, and GABA-T enzyme activities as well as increased levels of organic acids such as citric acid, succinic acid, and GABA content [12, 26, 120–122].

3.4. Cytosolic Malate Metabolism

In plants, malate is important in regulating the cytoplasmic pH, cell acidity, and carbon metabolism. Malate is synthesized by converting the phosphoenol-pyruvate (PEP) into oxaloacetate (OAA) by the activities of cytosolic enzymes, including phosphoenol-pyruvate carboxylase (PEPC) and NAD-linked malate dehydrogenase (NAD-MDH) [33]. It has been demonstrated that both OAA and malate can enter the mitochondrial TCA cycle to generate numerous organic acids. Malate can also be converted to pyruvate by the cytosolic NADP-linked malic enzyme (NADP-ME) via a dicarboxylate carrier [123].

Little research has been done into the possible link between cytosolicmalate metabolism and ethanol metabolism. In a recent study, Huan et al. [21] reported the promising involvement of higher expression of NADP-dependent malic enzymes (NADP-MEs) in increased ethanol production that eventually develops an alcoholic off-flavor in kiwifruit cv. “Bruno” during ambient storage conditions. Malate metabolism has previously been shown to induce ethanol metabolism by converting stored malate into pyruvate under various atmospheric conditions in harvested fruit such as grapes, berries, and strawberries [20, 123]. In contrast, Ponce-Valadez and Watkins [33] found a discrepancy between ethanol metabolism-related metabolites and enzymes involved in cytosolic malate metabolism. However, more omics-based research is needed to validate the role of cytosolic-malate metabolism in causing alcoholic off-flavor. A proposed schematic diagram of the relationship between cytosolic malate and ethanol metabolism is depicted in Figure 4.

3.5. Starch and Sugar Metabolism

Fruits primarily convert their starch stores into sugars as their primary energy source [124]. The flavor of ripe fruit is largely determined by the number of soluble sugars that accumulate during ripening [125]. Cultivars differ significantly in starch degradation and sugar composition at various stages of fruit development and ripening [126]. The starch in starch-based crops reportedly needs to be converted into simple sugars before it can be fermented into ethanol [127]. Previous research has indicated that sugars might be the most important substrates utilized by ethanol fermentation during fruit ripening [128]. β-amylase plays a crucial role in carbohydrate metabolism by depolymerizing α-glucan chains, thereby facilitating starch degradation during fruit ripening [129, 130], as evidenced in blueberries, as starch content drops at the same time as the rise in β-amylase gene expression [131]. It has been shown that the conversion of mango fruit starch to soluble sugars occurs simultaneously when the anaerobic respiration pathway is initiated [132]. Therefore, elevated starch degradation to sugars phenomenon may affect the biosynthesis of ethanol metabolism metabolites, which could initiate an off-flavor due to the over production and accumulation of ethanol content under anaerobic respiration.

Regarding this, Huan et al. [21] revealed that starch-to-sucrose metabolism (starch degradation) might induce alcoholic off-flavor disorder in “Bruno” kiwifruit during postharvest storage based on the findings associated with a dramatic decline in starch contents coincided with increased sucrose, fructose, and glucose levels accompanied with enhanced acetaldehyde and ethanol contents along with higher expressions of key genes such as starch phosphorylases (SPs), beta-amylases, UDP-glucose pyrophosphorylases (UGPases), sucrose synthases (SSs), and invertases (INVs) which are responsible for accelerating starch conversion to soluble sugars. This study provided transcriptome evidence that increased starch catabolism during fruit ripening may function as a substrate to promote ethanol fermentation, which induces an alcoholic off-flavor.

4. Measures to Control Alcoholic Off-Flavor Disorder Based on Changes in Ethanol Metabolism

4.1. Chemical Treatments

Numerous chemical treatments have been studied and applied to various fresh fruits to eradicate the susceptibility of alcoholic off-flavor disorder via suppressing the ethanol metabolism in both pre- and postharvest stages. Our recent study indicates that 1-MCP (1 μL·L⁻¹) treatment significantly reduced the occurrence of alcoholic off-flavor disorder in kiwifruit during storage at ambient conditions via inhibiting the ethanol metabolism-related enzymatic activities of PDC and ADH [12]. Subsequently, Huan et al. [133] also reported that 1 μL·L⁻¹ 1-MCP treatment could effectively reduce the alcoholic off-flavor disorder by suppressing ethanol metabolism in kiwifruit during ambient storage. Thewes et al. [134] concluded that applying 1-MCP together with dynamic controlled atmosphere (DCA) was particularly effective in lowering acetaldehyde and ethanol biosynthesis in apple fruit. Such findings were correlated with reduced PDC and ADH activity in apples, except during the mature stage, regardless of the cultivar. The lower levels of acetaldehyde and ethanol contents were also noticed in “Galaxy” apples under the effect of 1-MCP treatment and a dynamically controlled atmosphere based on respiratory quotient [135]. It has also been observed that 1-MCP could substantially lower the expression of PDC2 and ADH, reducing acetaldehyde and ethanol synthesis in apple fruit [136].

Recently, Lv et al. [22] demonstrated that exogenous application of 100 μmol·L⁻¹ nordihydroguaiaretic acid (NDGA) on “Nanguo” pear fruit resulted in lower acetaldehyde and ethanol contents concomitant with downregulated expressions of PDC1, ADH1, and ADH2 genes. Our previous work also suppressed the ethanol metabolism via preharvest spraying of 5 mmol·L⁻¹ oxalic acid (OA), which resulted in the absence of alcoholic off-flavor disorder in kiwifruit “Bruno” during storage under ambient conditions [32]. In another study, Zhang et al. [137] demonstrated that fumigation with 10 μmol·L⁻¹ carbon monoxide (CO) on the winter jujube might restrain the ethanol metabolism-related metabolites (acetaldehyde and ethanol) under reduced PDC and ADH activities. Li et al. [17] reported that treating strawberries with 1 mM ATP in a 20% CO₂ storage atmosphere reduced acetaldehyde and ethanol over-accumulation (72% and 75% lower in ATP + CO₂-treated fruit, respectively) that was linked to lower PDC and ADH activities during fruit storage. In a recent study, the effects of melatonin on ethanol metabolism in kiwifruit were also investigated, resulting in reduced acetaldehyde and ethanol contents under suppressed PDC and ADH activities and downregulated expressions of potential genes such as AdADH1, AdPDC1, and AdPDC2 during storage [138]. These changes may eventually result in decreased synthesis of aromatic compounds such as acetaldehyde and ethanol, thus controlling the alcoholic off-flavor disorder in fresh fruits during storage.

4.2. Hypobaric and Other Treatments

Several studies have found that hypobaric treatments can help suppress or delay various physiological disorders in fresh fruits. To suppress the alcoholic off-flavor disorder, Huan et al. [19] reported the significant effects of hypobaric treatment (25 ± 5 kPa for 30 min/twice treatment) in alleviating the alcoholic off-flavor disorder by inhibiting the activities of PDC and ADH in kiwifruit during storage. Previously, it had been demonstrated that fresh fruits could reduce the occurrence of alcoholic off-flavor by reducing the ethanol metabolism metabolites during storage under modified super-atmospheric O₂ exposure [44, 139]. Similarly, Wood et al. [51] observed a decrease in acetaldehyde and ethanol levels under controlled storage conditions (O₂ kPa⁻¹ CO₂ kPa^−2.5) for “Golden Delicious” and “Jonagold” apple fruit. Furthermore, Chen et al. [140] reported the decrease in ethanol and acetaldehyde production in “Benihoppe” strawberries under low oxygen application (2 kPa O₂) during storage. Zuo et al. [141] investigated the effects of high relative humidity (RH: 98 ± 2%) on ethanol metabolism in zucchini fruit, identifying that it inhibited the activities of PDC and ADH, as well as CpPDC1 and CpADH1 gene transcripts, resulting in decreased acetaldehyde and ethanol levels during cold storage. Previously, certain coating formulations were reported to initiate ethanol metabolism, resulting in off-flavor disorder in various fruits due to a change in the internal gas atmosphere of the fruit.

Meanwhile, a recent study showed that Carnauba wax nano-emulsion (attributed to being an effective moisture barrier and relatively permeable to gases) as a coating material had the least impact on the occurrence of alcoholic off-flavor by avoiding the production of ethanol above the threshold level in citrus fruit during storage [142]. Moreover, Velazco et al. [143] also concluded that Brillaqua_F6 (18% solids (9.35% oxidized polyethylene wax and 5.7% shellac)), Citrosol_AK (18% solids (Carnauba E903 and shellac)), and Teycer_GLK (18% solids (Carnauba and shellac)) coatings on citrus fruit had promising effects about reduced acetaldehyde and ethanol contents, thus decreasing the susceptibility to alcoholic off-flavor disorder. The overall impacts of chemical, hypobaric, and other treatments on ethanol metabolism to suppress the alcoholic off-flavor disorder in fresh fruits are summarized in Table 3.

5. Conclusion and Future Work

Anaerobic respiration, low O₂ stress, high CO₂ stress, and storage temperature stress are the most important factors for inducing alcoholic off-flavor disorder by upregulating ethanol metabolism in fresh fruits. These factors further cause undesirable metabolic changes at cellular levels by altering physiological mechanisms such as the GABA shunt pathway, mitochondrial energy metabolism, glycolysis and TCA cycles, cytosolic-malate metabolism, and starch and sugar metabolism. These physiological metabolic mechanisms might further activate ethanol metabolism, resulting in over-biosynthesis and the accumulation of alcoholic off-flavor-related metabolites such as acetaldehyde and ethanol in fresh fruits. Several chemicals, hypobaric, and coatings treatments have been practiced to overcome alcoholic off-flavor disorders in fresh fruits during storage. Despite this, a wide research gap still prevails that needs further investigation at the omics level to evaluate or validate the impact of numerous physiological mechanisms on ethanol metabolism and reveal the detailed mechanistic relation. In addition, other mechanisms and factors, including respiratory mechanisms, fruit genetic makeup, degree of maturity, and postharvest handling and storage conditions, are also recommended for future work concerning the alcoholic off-flavor disorder in fresh fruits during storage. The overall thematic scheme representing the current review is shown in Figure 5.

Data Availability

The dataset supporting the conclusions of this article is included within the article.

Conflicts of Interest

The authors declare that there are no conflicts of interest.

Authors’ Contributions

Maratab Ali conceptualized the study and wrote the original draft. Sara Batool, Muhammad Faisal Manzoor, Rana Muhammad Aadil, and Xiuming Zhao wrote, reviewed, and edited the article was responsible for writing and review and editing. Nauman Khalid and Fujun Li performed visualization and wrote, reviewed, and edited the paper. Xiaoan Li was responsible for validation, writing, reviewing, and editing. Xinhua Zhang was responsible for conceptualization, writing and review, editing, funding, and supervision.

Acknowledgments

This work was supported by the National Natural Science Foundation of China (Grant no. 32172278) and Projects in the Natural Science Foundation of Shandong Province (Grant Nos. ZR2020KC011 and ZR2020MC149).

References

H. A. Khalil, M. F. Abdelkader, A. Lo’ay et al., “The combined effect of hot water treatment and chitosan coating on mango (Mangifera indica L. cv. Kent) fruits to control postharvest deterioration and increase fruit quality,” Coatings, vol. 12, no. 1, p. 83, 2022.
View at: Publisher Site | Google Scholar
M. F. Manzoor, A. Hussain, N. Naumovski et al., “A narrative review of recent advances in rapid assessment of anthocyanins in agricultural and food products,” Frontiers in Nutrition, vol. 9, Article ID 901342, 2022.
View at: Publisher Site | Google Scholar
A. Rahaman, X. A. Zeng, M. A. Farooq et al., “Effect of pulsed electric fields processing on physiochemical properties and bioactive compounds of apricot juice,” Journal of Food Process Engineering, vol. 43, no. 8, 2020.
View at: Publisher Site | Google Scholar
M. Shehzad, H. Rasheed, S. A. Naqvi et al., “Therapeutic potential of date palm against human infertility: a review,” Metabolites, vol. 11, no. 6, p. 408, 2021.
View at: Publisher Site | Google Scholar
Y. Sultanbawa and D. Sivakumar, “Enhanced nutritional and phytochemical profiles of selected underutilized fruits, vegetables and legumes,” Current Opinion in Food Science, vol. 46, Article ID 100853, 2022.
View at: Publisher Site | Google Scholar
M. Ali, M. A. Raza, S. Li et al., “1-MCP regulates ethanol fermentation and GABA shunt pathway involved in kiwifruit quality during postharvest storage,” Horticultural Plant Journal, vol. 7, no. 1, pp. 23–30, 2021.
View at: Publisher Site | Google Scholar
M. D. Antunes, C. Gago, and A. J. H. Guerreiro, “Postharvest handling of horticultural products,” Horticulturae, vol. 8, p. 726, 2022.
View at: Publisher Site | Google Scholar
J. Tian, S. Xie, P. Zhang, Q. Wang, J. Li, and X. Xu, “Attenuation of postharvest peel browning and chilling injury of banana fruit by Astragalus polysaccharides,” Postharvest Biology and Technology, vol. 184, Article ID 111783, 2022.
View at: Publisher Site | Google Scholar
M. Cefola, A. Damascelli, V. Lippolis et al., “Relationships among volatile metabolites, quality and sensory parameters of ‘Italia’table grapes assessed during cold storage in low or high CO2 modified atmospheres,” Postharvest Biology and Technology, vol. 142, pp. 124–134, 2018.
View at: Publisher Site | Google Scholar
D. Obenland and M. L. Arpaia, “Effect of harvest date on off-flavor development in mandarins following postharvest wax application,” Postharvest Biology and Technology, vol. 149, pp. 1–8, 2019.
View at: Publisher Site | Google Scholar
N. Ummarat, M. L. Arpaia, and D. Obenland, “Physiological, biochemical and sensory characterization of the response to waxing and storage of two Mandarin varieties differing in postharvest ethanol accumulation,” Postharvest Biology and Technology, vol. 109, pp. 82–96, 2015.
View at: Publisher Site | Google Scholar
M. Ali, M. A. Raza, S. Li, C. Huan, and X. Zheng, “1-Methylcyclopropene treatment controls ethanol accumulation associated with regulation of mitochondrial energy metabolism in kiwifruit (Actinidia deliciosa) cv. “Bruno” during storage at room temperature,” Journal of Food Biochemistry, vol. 44, no. 7, 2020.
View at: Publisher Site | Google Scholar
J. X. Shi, R. Porat, R. Goren, and E. E. Goldschmidt, “Physiological responses of ‘Murcott’mandarins and ‘Star Ruby’grapefruit to anaerobic stress conditions and their relation to fruit taste, quality and emission of off-flavor volatiles,” Postharvest Biology and Technology, vol. 38, no. 2, pp. 99–105, 2005.
View at: Publisher Site | Google Scholar
E. Pesis, “The role of the anaerobic metabolites, acetaldehyde and ethanol, in fruit ripening, enhancement of fruit quality and fruit deterioration,” Postharvest Biology and Technology, vol. 37, no. 1, pp. 1–19, 2005.
View at: Publisher Site | Google Scholar
M. E. Saltveit, “Respiratory metabolism,” in Postharvest Physiology and Biochemistry of Fruits and Vegetables, pp. 73–91, Elsevier, Amsterdam, Netherlands, 2019.
View at: Publisher Site | Google Scholar
C. Techavuthiporn, M. Thammawong, and K. Nakano, “Effect of short-term anoxia treatment on endogenous ethanol and postharvest responses of broccoli florets during storage at ambient temperature,” Scientia Horticulturae, vol. 277, Article ID 109813, 2021.
View at: Publisher Site | Google Scholar
D. Li, L. Li, Y. Xu et al., “Exogenous ATP attenuated fermentative metabolism in postharvest strawberry fruit under elevated CO2 atmosphere by maintaining energy status,” Postharvest Biology and Technology, vol. 182, Article ID 111701, 2021.
View at: Publisher Site | Google Scholar
X. Yang, J. Song, S. Fillmore, X. Pang, and Z. Zhang, “Effect of high temperature on color, chlorophyll fluorescence and volatile biosynthesis in green-ripe banana fruit,” Postharvest Biology and Technology, vol. 62, no. 3, pp. 246–257, 2011.
View at: Publisher Site | Google Scholar
C. Huan, H. Li, Z. Jiang, S. Li, S. Shen, and X. Zheng, “Effect of hypobaric treatment on off-flavour development and energy metabolism in ‘Bruno’kiwifruit,” LWT--Food Science and Technology, vol. 136, Article ID 110349, 2021.
View at: Publisher Site | Google Scholar
F. Famiani, D. Farinelli, A. Palliotti, S. Moscatello, A. Battistelli, and R. P. Walker, “Is stored malate the quantitatively most important substrate utilised by respiration and ethanolic fermentation in grape berry pericarp during ripening?” Plant Physiology and Biochemistry, vol. 76, pp. 52–57, 2014.
View at: Publisher Site | Google Scholar
C. Huan, X. Du, L. Wang et al., “Transcriptome analysis reveals the metabolisms of starch degradation and ethanol fermentation involved in alcoholic off-flavour development in kiwifruit during ambient storage,” Postharvest Biology and Technology, vol. 180, Article ID 111621, 2021.
View at: Publisher Site | Google Scholar
J. Lv, D. Xu, Y. Zhang et al., “Effects of abscisic acid (ABA) on ethanol fermentation during postharvest ripening of Nanguo pear fruit (Pyrus ussuriensis Maxim.),” Scientia Horticulturae, vol. 288, Article ID 110388, 2021.
View at: Publisher Site | Google Scholar
J. Boeckx, M. Hertog, A. Geeraerd, and B. Nicolaï, “Regulation of the fermentative metabolism in apple fruit exposed to low-oxygen stress reveals a high flexibility,” Postharvest Biology and Technology, vol. 149, pp. 118–128, 2019.
View at: Publisher Site | Google Scholar
Q. T. Ho, P. Verboven, A. Ambaw, B. E. Verlinden, and B. M. Nicolaï, “Transport properties of fermentation metabolites inside ‘Conference’pear fruit,” Postharvest Biology and Technology, vol. 117, pp. 38–48, 2016.
View at: Publisher Site | Google Scholar
P. Ahmad and M. N. V. Prasad, “Drought stress induced reactive oxygen species and anti-oxidants in plants,” Abiotic Stress Responses in Plants: Metabolism, Productivity and Sustainability, Springer Science & Business Media, Berlin, Germany, 2011.
View at: Publisher Site | Google Scholar
L. T. Bui, G. Novi, L. Lombardi et al., “Conservation of ethanol fermentation and its regulation in land plants,” Journal of Experimental Botany, vol. 70, no. 6, pp. 1815–1827, 2019.
View at: Publisher Site | Google Scholar
M. Mithran, E. Paparelli, G. Novi, P. Perata, and E. Loreti, “Analysis of the role of the pyruvate decarboxylase gene family in A rabidopsis thaliana under low‐oxygen conditions,” Plant Biology, vol. 16, no. 1, pp. 28–34, 2014.
View at: Publisher Site | Google Scholar
I. R. White, R. S. Blake, A. J. Taylor, and P. S. Monks, “Metabolite profiling of the ripening of Mangoes Mangifera indica L. cv.‘Tommy Atkins’ by real-time measurement of volatile organic compounds,” Metabolomics, vol. 12, no. 3, p. 57, 2016.
View at: Publisher Site | Google Scholar
R. Botondi, V. Russo, and F. Mencarelli, “Anaerobic metabolism during short and long term storage of kiwifruit,” Postharvest Biology and Technology, vol. 64, no. 1, pp. 83–90, 2012.
View at: Publisher Site | Google Scholar
J. Zhang and C. B. Watkins, “Fruit quality, fermentation products, and activities of associated enzymes during elevated CO2 treatment of strawberry fruit at high and low temperatures,” Journal of the American Society for Horticultural Science, vol. 130, no. 1, pp. 124–130, 2005.
View at: Publisher Site | Google Scholar
J. X. Shi, E. E. Goldschmidt, R. Goren, and R. Porat, “Molecular, biochemical and anatomical factors governing ethanol fermentation metabolism and accumulation of off-flavors in mandarins and grapefruit,” Postharvest Biology and Technology, vol. 46, no. 3, pp. 242–251, 2007.
View at: Publisher Site | Google Scholar
M. Ali, M. Liu, Z. Wang, S. Li, T. Jiang, and X. Zheng, “Pre-harvest spraying of oxalic acid improves postharvest quality associated with increase in ascorbic acid and regulation of ethanol fermentation in kiwifruit cv. Bruno during storage,” Journal of Integrative Agriculture, vol. 18, no. 11, pp. 2514–2520, 2019.
View at: Publisher Site | Google Scholar
M. Ponce‐Valadez and C. B. Watkins, “Fermentation and malate metabolism in response to elevated CO2 concentrations in two strawberry cultivars,” Physiologia Plantarum, vol. 134, no. 1, pp. 121–133, 2008.
View at: Publisher Site | Google Scholar
Y. Imahori, M. Kota, Y. Ueda, M. Ishimaru, and K. Cachin, “Regulation of ethanolic fermentation in bell pepper fruit under low oxygen stress,” Postharvest Biology and Technology, vol. 25, no. 2, pp. 159–167, 2002.
View at: Publisher Site | Google Scholar
J. Strommer, “The plant ADH gene family,” The Plant Journal, vol. 66, no. 1, pp. 128–142, 2011.
View at: Publisher Site | Google Scholar
J. Zhang, S. Huang, G. Wang, J. Xuan, and Z. Guo, “Overexpression of Actinidia deliciosa pyruvate decarboxylase 1 gene enhances waterlogging stress in transgenic Arabidopsis thaliana,” Plant Physiology and Biochemistry, vol. 106, pp. 244–252, 2016.
View at: Publisher Site | Google Scholar
A. Kader and E. Yahia, “Postharvest biology of tropical and subtropical fruits,” in Postharvest Biology and Technology of Tropical and Subtropical Fruits, pp. 79–111, Elsevier, Amsterdam, Netherlands, 2011.
View at: Publisher Site | Google Scholar
Y. Kubo, “Ethylene, oxygen, carbon dioxide, and temperature in postharvest physiology,” in Abiotic Stress Biology in Horticultural Plants, pp. 17–33, Springer, Berlin, Germany, 2015.
View at: Publisher Site | Google Scholar
D. Cukrov, S. Brizzolara, and P. Tonutti, “Physiological and biochemical effects of controlled and modified atmospheres,” in Postharvest Physiology and Biochemistry of Fruits and Vegetables, pp. 425–441, Elsevier, Amsterdam, Netherlands, 2019.
View at: Publisher Site | Google Scholar
H. Gao, W. Hu, Y. Guan, Y. Ji, X. Yang, and M. Hou, “Respiratory metabolism and quality in postharvest sweet cherries (Prunus avium L.) in response to high CO₂ treatment,” Journal of Food Processing and Preservation, vol. 46, no. 10, 2022.
View at: Publisher Site | Google Scholar
M. Buccheri, V. Picchi, M. Grassi, D. Gandin, G. Bianchi, and R. L. Scalzo, “Dynamic changes of antioxidants and fermentative metabolites in apple peel in relation to storage, controlled atmosphere, and initial low oxygen stress,” Scientia Horticulturae, vol. 288, Article ID 110312, 2021.
View at: Publisher Site | Google Scholar
A. R. Malik, R. Raja, and R. Javaid, “Physiological disorders in stone fruits,” in Production Technology of Stone Fruits, pp. 189–209, Springer, Berlin, Germany, 2021.
View at: Publisher Site | Google Scholar
M. L. Ntsoane, A. Luca, M. Zude-Sasse, D. Sivakumar, and P. V. Mahajan, “Impact of low oxygen storage on quality attributes including pigments and volatile compounds in ‘Shelly’mango,” Scientia Horticulturae, vol. 250, pp. 174–183, 2019.
View at: Publisher Site | Google Scholar
A. Wright, J. Delong, J. Arul, and R. Prange, “The trend toward lower oxygen levels during apple (Malus× domestica Borkh) storage,” The Journal of Horticultural Science and Biotechnology, vol. 90, no. 1, pp. 1–13, 2015.
View at: Publisher Site | Google Scholar
D. Park, Y. A. Shoffe, B. E. Algul, and C. B. Watkins, “Fermentative metabolism of three apple cultivars during storage under low partial pressures of oxygen,” Postharvest Biology and Technology, vol. 193, Article ID 112037, 2022.
View at: Publisher Site | Google Scholar
J. Burdon, N. Lallu, C. Yearsley, D. Burmeister, and D. Billing, “The kinetics of acetaldehyde and ethanol accumulation in ‘Hass’ avocado fruit during induction and recovery from low oxygen and high carbon dioxide conditions,” Postharvest Biology and Technology, vol. 43, no. 2, pp. 207–214, 2007.
View at: Publisher Site | Google Scholar
S. Chockchaisawasdee, J. B. Golding, Q. V. Vuong, K. Papoutsis, and C. E. Stathopoulos, “Sweet cherry: composition, postharvest preservation, processing and trends for its future use,” Trends in Food Science & Technology, vol. 55, pp. 72–83, 2016.
View at: Publisher Site | Google Scholar
N. Nakamura, D. V. S. Rao, T. Shiina, and Y. Nawa, “Respiration properties of tree-ripe mango under CA condition,” Japan Agricultural Research Quarterly: Japan Agricultural Research Quarterly, vol. 38, no. 4, pp. 221–226, 2004.
View at: Publisher Site | Google Scholar
J. Goliáš, A. Němcová, A. Čaněk, and D. Kolenčíková, “Storage of sweet cherries in low oxygen and high carbon dioxide atmospheres,” Horticultural Science, vol. 34, no. 1, pp. 26–34, 2007.
View at: Publisher Site | Google Scholar
G. Polenta, C. Budde, and R. Murray, “Effects of different pre-storage anoxic treatments on ethanol and acetaldehyde content in peaches,” Postharvest Biology and Technology, vol. 38, no. 3, pp. 247–253, 2005.
View at: Publisher Site | Google Scholar
R. M. Wood, F. R. Thewes, M. Reynaud et al., “Apple fruit recovery from anoxia under controlled atmosphere storage,” Food Chemistry, vol. 371, Article ID 131152, 2022.
View at: Publisher Site | Google Scholar
E. Pintó, I. Lentheric, M. Vendrell, and C. Larrigaudière, “Role of fermentative and antioxidant metabolisms in the induction of core browning in controlled-atmosphere stored pears,” Journal of the Science of Food and Agriculture, vol. 81, no. 3, pp. 364–370, 2001.
View at: Publisher Site | Google Scholar
Y. Imahori, K. Matushita, M. Kota, Y. Ueda, M. Ishimaru, and K. Chachin, “Regulation of fermentative metabolism in tomato fruit under low oxygen stress,” The Journal of Horticultural Science and Biotechnology, vol. 78, no. 3, pp. 386–393, 2003.
View at: Publisher Site | Google Scholar
B. Madani, A. Mirshekari, and Y. Imahori, “Physiological responses to stress,” in Postharvest Physiology and Biochemistry of Fruits and Vegetables, pp. 405–423, Elsevier, Amsterdam, Netherlands, 2019.
View at: Publisher Site | Google Scholar
G. H. Teixeira, L. C. Cunha Júnior, A. S. Ferraudo, and J. F. Durigan, “Quality of guava (Psidium guajava L. cv. Pedro Sato) fruit stored in low-O2 controlled atmospheres is negatively affected by increasing levels of CO2,” Postharvest Biology and Technology, vol. 111, pp. 62–68, 2016.
View at: Publisher Site | Google Scholar
S. P. Tian, A. L. Jiang, Y. Xu, and Y. S. Wang, “Responses of physiology and quality of sweet cherry fruit to different atmospheres in storage,” Food Chemistry, vol. 87, no. 1, pp. 43–49, 2004.
View at: Publisher Site | Google Scholar
L. Wang and S. Vestrheim, “Controlled atmosphere storage of sweet cherries (Prunus avium L.),” Acta Agriculturae Scandinavica Section B Soil and Plant Science, vol. 52, no. 4, pp. 136–142, 2002.
View at: Publisher Site | Google Scholar
R. Cozzolino, A. Martignetti, M. Cefola et al., “Volatile metabolites, quality and sensory parameters of “Ferrovia” sweet cherry cold stored in air or packed in high CO₂ modified atmospheres,” Food Chemistry, vol. 286, pp. 659–668, 2019.
View at: Publisher Site | Google Scholar
J. Dong, X. Li, G. Nazim, and Z. Duan, “Interactive effects of elevated carbon dioxide and nitrogen availability on fruit quality of cucumber (Cucumis sativus L.),” Journal of Integrative Agriculture, vol. 17, no. 11, pp. 2438–2446, 2018.
View at: Publisher Site | Google Scholar
S. Singh and R. Pal, “Controlled atmosphere storage of guava (Psidium guajava L.) fruit,” Postharvest Biology and Technology, vol. 47, no. 3, pp. 296–306, 2008.
View at: Publisher Site | Google Scholar
G. H. Teixeira and J. F. Durigan, “Effect of controlled atmospheres with low oxygen levels on extended storage of guava fruit (Psidium guajava L. ‘Pedro Sato’),” HortScience, vol. 45, no. 6, pp. 918–924, 2010.
View at: Publisher Site | Google Scholar
Y. Lu, D. Li, L. Li et al., “Effects of elevated CO₂ on pigment metabolism of postharvest Mandarin fruit for degreening,” Food Chemistry, vol. 318, Article ID 126462, 2020.
View at: Publisher Site | Google Scholar
J. Harman and B. McDonald, “Controlled atmosphere storage of kiwifruit. Effect on fruit quality and composition,” Scientia Horticulturae, vol. 37, no. 4, pp. 303–315, 1989.
View at: Publisher Site | Google Scholar
D. Ke and A. A. Kader, “Tolerance of `Valencia' oranges to controlled atmospheres, as determined by physiological responses and quality attributes,” Journal of the American Society for Horticultural Science, vol. 115, no. 5, pp. 779–783, 1990.
View at: Publisher Site | Google Scholar
D. Ke, E. Yahia, M. Mateos, and A. A. Kader, “Ethanolic fermentation of `Bartlett' pears as influenced by ripening stage and atmospheric composition,” Journal of the American Society for Horticultural Science, vol. 119, no. 5, pp. 976–982, 1994.
View at: Publisher Site | Google Scholar
M. Mateos, D. Ke, M. Cantwell, and A. A. Kader, “Phenolic metabolism and ethanolic fermentation of intact and cut lettuce exposed to CO₂-enriched atmospheres,” Postharvest Biology and Technology, vol. 3, no. 3, pp. 225–233, 1993.
View at: Publisher Site | Google Scholar
E. Pesis and R. Marinansky, “Inhibition of tomato ripening by acetaldehyde vapour or anaerobic conditions prior to storage,” Journal of Plant Physiology, vol. 142, no. 6, pp. 717–721, 1993.
View at: Publisher Site | Google Scholar
R. Porat and E. Fallik, “Production of off-flavours in fruit and vegetables under fermentative conditions,” in Fruit and Vegetable Flavour, pp. 150–164, Elsevier, Amsterdam, Netherlands, 2008.
View at: Publisher Site | Google Scholar
H. Lalel and Z. Singh, “Controlled atmosphere storage of ‘Delta R2E2’ mango fruit affects production of aroma volatile compounds,” The Journal of Horticultural Science and Biotechnology, vol. 81, no. 3, pp. 449–457, 2006.
View at: Publisher Site | Google Scholar
E. M. Yahia, “Modified and controlled atmospheres for tropical fruits,” Stew Post Review, vol. 5, no. 6, pp. 1–10, 2006.
View at: Publisher Site | Google Scholar
Y. Wang, J. Bai, and L. E. Long, “Quality and physiological responses of two late-season sweet cherry cultivars ‘Lapins’ and ‘Skeena’to modified atmosphere packaging (MAP) during simulated long distance ocean shipping,” Postharvest Biology and Technology, vol. 110, pp. 1–8, 2015.
View at: Publisher Site | Google Scholar
C. F. Forney, M. A. Jordan, K. M. Pennell, and S. Fillmore, “Controlled atmosphere storage impacts fruit quality and flavor chemistry of five cultivars of highbush blueberry (Vaccinium corymbosum),” Postharvest Biology and Technology, vol. 194, Article ID 112073, 2022.
View at: Publisher Site | Google Scholar
A. Bahar and A. Lichter, “Effect of controlled atmosphere on the storage potential of Ottomanit fig fruit,” Scientia Horticulturae, vol. 227, pp. 196–201, 2018.
View at: Publisher Site | Google Scholar
I. Maoz, M. De Rosso, T. Kaplunov et al., “Metabolomic and transcriptomic changes underlying cold and anaerobic stresses after storage of table grapes,” Scientific Reports, vol. 9, no. 1, p. 2917, 2019.
View at: Publisher Site | Google Scholar
M. Blanch, I. Álvarez, M. T. Sanchez-Ballesta, M. I. Escribano, and C. Merodio, “Involvement of fatty acids in the response to high CO₂ and low temperature in harvested strawberries,” Postharvest Biology and Technology, vol. 147, pp. 196–205, 2019.
View at: Publisher Site | Google Scholar
M. Blanch, R. Rosales, R. Mateos et al., “Effects of high CO2 levels on fermentation, peroxidation, and cellular water stress in Fragaria vesca stored at low temperature in conditions of unlimited O2,” Journal of Agricultural and Food Chemistry, vol. 63, no. 3, pp. 761–768, 2015.
View at: Publisher Site | Google Scholar
M. Blanch, R. Rosales, F. Palma, M. T. Sanchez-Ballesta, M. I. Escribano, and C. Merodio, “CO2-driven changes in energy and fermentative metabolism in harvested strawberries,” Postharvest Biology and Technology, vol. 110, pp. 33–39, 2015.
View at: Publisher Site | Google Scholar
M. Cefola and B. Pace, “High CO2-modified atmosphere to preserve sensory and nutritional quality of organic table grape (cv.‘Italia’) during storage and shelf-life,” European Journal of Horticultural Science, vol. 81, no. 4, pp. 197–203, 2016.
View at: Publisher Site | Google Scholar
C. Lumpkin, J. K. Fellman, D. R. Rudell, and J. P. Mattheis, “‘Fuji’apple (Malus domestica Borkh.) volatile production during high pCO₂ controlled atmosphere storage,” Postharvest Biology and Technology, vol. 100, pp. 234–243, 2015.
View at: Publisher Site | Google Scholar
E. Fallik and Z. Ilic, “Pre-and postharvest treatments affecting flavor quality of fruits and vegetables,” in Preharvest Modulation of Postharvest Fruit and Vegetable Quality, pp. 139–168, Elsevier, Amsterdam, Netherlands, 2018.
View at: Publisher Site | Google Scholar
B. Saberi, J. B. Golding, J. R. Marques et al., “Application of biocomposite edible coatings based on pea starch and guar gum on quality, storability and shelf life of ‘Valencia’ oranges,” Postharvest Biology and Technology, vol. 137, pp. 9–20, 2018.
View at: Publisher Site | Google Scholar
D. Obenland, S. Collin, J. Sievert, and M. L. Arpaia, “Mandarin flavor and aroma volatile composition are strongly influenced by holding temperature,” Postharvest Biology and Technology, vol. 82, pp. 6–14, 2013.
View at: Publisher Site | Google Scholar
P. D. Petracek, D. W. Joles, A. Shirazi, and A. C. Cameron, “Modified atmosphere packaging of sweet cherry (Prunus avium L., ev.‘Sams’) fruit: metabolic responses to oxygen, carbon dioxide, and temperature,” Postharvest Biology and Technology, vol. 24, no. 3, pp. 259–270, 2002.
View at: Publisher Site | Google Scholar
O. J. Caleb, O. A. Fawole, R. R. Mphahlele, and U. L. Opara, “Impact of preharvest and postharvest factors on changes in volatile compounds of pomegranate fruit and minimally processed arils–Review,” Scientia Horticulturae, vol. 188, pp. 106–114, 2015.
View at: Publisher Site | Google Scholar
E. M. Yahia, Postharvest Technology of Perishable Horticultural Commodities, Woodhead Publishing, Sawston, UK, 2019.
J. Zou, J. Chen, N. Tang et al., “Transcriptome analysis of aroma volatile metabolism change in tomato (Solanum lycopersicum) fruit under different storage temperatures and 1-MCP treatment,” Postharvest Biology and Technology, vol. 135, pp. 57–67, 2018.
View at: Publisher Site | Google Scholar
C. S. Günther, K. B. Marsh, R. A. Winz et al., “The impact of cold storage and ethylene on volatile ester production and aroma perception in ‘Hort16A’ kiwifruit,” Food Chemistry, vol. 169, pp. 5–12, 2015.
View at: Publisher Site | Google Scholar
M. C. Do Nascimento Nunes, “Impact of environmental conditions on fruit and vegetable quality,” Stew Post Rev, vol. 4, no. 4, pp. 1–14, 2008.
View at: Publisher Site | Google Scholar
J. F. Ayala-Zavala, S. Y. Wang, C. Y. Wang, and G. A. González-Aguilar, “Effect of storage temperatures on antioxidant capacity and aroma compounds in strawberry fruit,” LWT-Food Science and Technology, vol. 37, no. 7, pp. 687–695, 2004.
View at: Publisher Site | Google Scholar
J. Bai, E. A. Baldwin, Y. Imahori, I. Kostenyuk, J. Burns, and J. K. Brecht, “Chilling and heating may regulate C6 volatile aroma production by different mechanisms in tomato (Solanum lycopersicum) fruit,” Postharvest Biology and Technology, vol. 60, no. 2, pp. 111–120, 2011.
View at: Publisher Site | Google Scholar
V. E. Perotti, H. A. Del Vecchio, A. Sansevich et al., “Proteomic, metabalomic, and biochemical analysis of heat treated Valencia oranges during storage,” Postharvest Biology and Technology, vol. 62, no. 2, pp. 97–114, 2011.
View at: Publisher Site | Google Scholar
J. Cano-Salazar, M. López, C. H. Crisosto, and G. Echeverría, “Volatile compound emissions and sensory attributes of ‘Big Top’nectarine and ‘Early Rich’peach fruit in response to a pre-storage treatment before cold storage and subsequent shelf-life,” Postharvest Biology and Technology, vol. 76, pp. 152–162, 2013.
View at: Publisher Site | Google Scholar
Y. Wang, Z. Luo, L. Mao, and T. Ying, “Contribution of polyamines metabolism and GABA shunt to chilling tolerance induced by nitric oxide in cold-stored banana fruit,” Food Chemistry, vol. 197, pp. 333–339, 2016.
View at: Publisher Site | Google Scholar
N. Bouche and H. Fromm, “GABA in plants: just a metabolite?” Trends in Plant Science, vol. 9, no. 3, pp. 110–115, 2004.
View at: Publisher Site | Google Scholar
D. Li, L. Li, G. Xiao et al., “Effects of elevated CO₂ on energy metabolism and γ-aminobutyric acid shunt pathway in postharvest strawberry fruit,” Food Chemistry, vol. 265, pp. 281–289, 2018.
View at: Publisher Site | Google Scholar
R. Pedreschi, C. Franck, J. Lammertyn et al., “Metabolic profiling of ‘Conference’pears under low oxygen stress,” Postharvest Biology and Technology, vol. 51, no. 2, pp. 123–130, 2009.
View at: Publisher Site | Google Scholar
M. S. Aghdam, R. Naderi, A. Jannatizadeh, M. Babalar, M. A. A. Sarcheshmeh, and M. Z. Faradonbe, “Impact of exogenous GABA treatments on endogenous GABA metabolism in anthurium cut flowers in response to postharvest chilling temperature,” Plant Physiology and Biochemistry, vol. 106, pp. 11–15, 2016.
View at: Publisher Site | Google Scholar
C. Franck, J. Lammertyn, Q. T. Ho, P. Verboven, B. Verlinden, and B. M. Nicolaï, “Browning disorders in pear fruit,” Postharvest Biology and Technology, vol. 43, no. 1, pp. 1–13, 2007.
View at: Publisher Site | Google Scholar
F. Palma, F. Carvajal, M. Jamilena, and D. Garrido, “Contribution of polyamines and other related metabolites to the maintenance of zucchini fruit quality during cold storage,” Plant Physiology and Biochemistry, vol. 82, pp. 161–171, 2014.
View at: Publisher Site | Google Scholar
G. Z. Chiu, B. J. Shelp, S. R. Bowley, J. R. DeEll, and G. G. Bozzo, “Controlled atmosphere-related injury in ‘Honeycrisp’apples is associated with γ-aminobutyrate accumulation,” Canadian Journal of Plant Science, vol. 95, no. 5, pp. 879–886, 2015.
View at: Publisher Site | Google Scholar
R. Deewatthanawong and C. Watkins, “Accumulation of γ-aminobutyric acid in apple, strawberry and tomato fruit in response to postharvest treatments,” Acta Horticulturae, vol. 877, pp. 947–952, 2010.
View at: Publisher Site | Google Scholar
R. S. Leisso, D. A. Buchanan, J. Lee et al., “Chilling‐related cell damage of apple (Malus× domestica Borkh.) fruit cortical tissue impacts antioxidant, lipid and phenolic metabolism,” Physiologia Plantarum, vol. 153, no. 2, pp. 204–220, 2015.
View at: Publisher Site | Google Scholar
R. Deewatthanawong, P. Rowell, and C. B. Watkins, “γ-Aminobutyric acid (GABA) metabolism in CO₂ treated tomatoes,” Postharvest Biology and Technology, vol. 57, no. 2, pp. 97–105, 2010.
View at: Publisher Site | Google Scholar
N. Mae, Y. Makino, S. Oshita et al., “Accumulation mechanism of γ-aminobutyric acid in tomatoes (Solanum lycopersicum L.) under low O₂ with and without CO₂,” Journal of Agricultural and Food Chemistry, vol. 60, no. 4, pp. 1013–1019, 2012.
View at: Publisher Site | Google Scholar
J. Li, X. Zhou, B. Wei, S. Cheng, Q. Zhou, and S. Ji, “GABA application improves the mitochondrial antioxidant system and reduces peel browning in ‘Nanguo’pears after removal from cold storage,” Food Chemistry, vol. 297, Article ID 124903, 2019.
View at: Publisher Site | Google Scholar
L. Song, J. Wang, M. Shafi et al., “Hypobaric treatment effects on chilling injury, mitochondrial dysfunction, and the ascorbate–glutathione (AsA-GSH) cycle in postharvest peach fruit,” Journal of Agricultural and Food Chemistry, vol. 64, no. 22, pp. 4665–4674, 2016.
View at: Publisher Site | Google Scholar
S. Cheng, B. Wei, Q. Zhou, D. Tan, and S. Ji, “1-Methylcyclopropene alleviates chilling injury by regulating energy metabolism and fatty acid content in ‘Nanguo’pears,” Postharvest Biology and Technology, vol. 109, pp. 130–136, 2015.
View at: Publisher Site | Google Scholar
D. Tao, J. Wang, L. Zhang, Y. Jiang, and M. Lv, “1-Methylcyclopropene alleviates peel browning of ‘Nanguo’pears by regulating energy, antioxidant and lipid metabolisms after long term refrigeration,” Scientia Horticulturae, vol. 247, pp. 254–263, 2019.
View at: Publisher Site | Google Scholar
Q. Zhou, C. Zhang, S. Cheng, B. Wei, X. Liu, and S. Ji, “Changes in energy metabolism accompanying pitting in blueberries stored at low temperature,” Food Chemistry, vol. 164, pp. 493–501, 2014.
View at: Publisher Site | Google Scholar
X. Zhang, D. Li, Y. Wang et al., “Fumigation of SO2 in combination with elevated CO2 regulate sugar and energy metabolism in postharvest strawberry fruit,” Postharvest Biology and Technology, vol. 192, Article ID 112021, 2022.
View at: Publisher Site | Google Scholar
N. Wang, H. Fang, Q. Yang, Z. Liu, H. Feng, and S. Ji, “Exogenous melatonin alleviated leaf yellowing via inhibiting respiration and ethylene biosynthesis during shelf life in pakchoi,” Plants, vol. 11, no. 16, p. 2102, 2022.
View at: Publisher Site | Google Scholar
L.-N. Sun, M.-C. Liu, S.-H. Zhu, J. Zhou, and M.-L. Wang, “Effect of nitric oxide on alcoholic fermentation and qualities of Chinese winter jujube during storage,” Agricultural Sciences in China, vol. 6, no. 7, pp. 849–856, 2007.
View at: Publisher Site | Google Scholar
A. Mannucci, A. Serra, D. Remorini et al., “Aroma profile of Fuji apples treated with gelatin edible coating during their storage,” LWT-Food Science and Technology, vol. 85, pp. 28–36, 2017.
View at: Publisher Site | Google Scholar
B. Biais, B. Beauvoit, J. William Allwood et al., “Metabolic acclimation to hypoxia revealed by metabolite gradients in melon fruit,” Journal of Plant Physiology, vol. 167, no. 3, pp. 242–245, 2010.
View at: Publisher Site | Google Scholar
A. R. Fernie, F. Carrari, and L. J. Sweetlove, “Respiratory metabolism: glycolysis, the TCA cycle and mitochondrial electron transport,” Current Opinion in Plant Biology, vol. 7, no. 3, pp. 254–261, 2004.
View at: Publisher Site | Google Scholar
R. Steuer, A. N. Nesi, A. R. Fernie, T. Gross, B. Blasius, and J. Selbig, “From structure to dynamics of metabolic pathways: application to the plant mitochondrial TCA cycle,” Bioinformatics, vol. 23, no. 11, pp. 1378–1385, 2007.
View at: Publisher Site | Google Scholar
S. Zhang, H. Lin, Y. Lin et al., “Energy status regulates disease development and respiratory metabolism of Lasiodiplodia theobromae (Pat.) Griff. & Maubl.-infected longan fruit,” Food Chemistry, vol. 231, pp. 238–246, 2017.
View at: Publisher Site | Google Scholar
J. Bailey-Serres, T. Fukao, D. J. Gibbs et al., “Making sense of low oxygen sensing,” Trends in Plant Science, vol. 17, no. 3, pp. 129–138, 2012.
View at: Publisher Site | Google Scholar
J. Chen, Y. Hu, R. Yan, and H. Hu, “Effect of high carbon dioxide injury on the physiological characteristics of fresh-cut garlic scapes,” Scientia Horticulturae, vol. 250, pp. 359–365, 2019.
View at: Publisher Site | Google Scholar
A. Chrysargyris, A. Nikou, and N. Tzortzakis, “Effectiveness of Aloe vera gel coating for maintaining tomato fruit quality,” New Zealand Journal of Crop and Horticultural Science, vol. 44, no. 3, pp. 203–217, 2016.
View at: Publisher Site | Google Scholar
R. Deewatthanawong, J. F. Nock, and C. B. Watkins, “γ-Aminobutyric acid (GABA) accumulation in four strawberry cultivars in response to elevated CO2 storage,” Postharvest Biology and Technology, vol. 57, no. 2, pp. 92–96, 2010.
View at: Publisher Site | Google Scholar
Y. Lin, M. Chen, H. Lin et al., “DNP and ATP induced alteration in disease development of Phomopsis longanae Chi-inoculated longan fruit by acting on energy status and reactive oxygen species production-scavenging system,” Food Chemistry, vol. 228, pp. 497–505, 2017.
View at: Publisher Site | Google Scholar
M. V. Jensen, J. W. Joseph, S. M. Ronnebaum, S. C. Burgess, A. D. Sherry, and C. B. Newgard, “Metabolic cycling in control of glucose-stimulated insulin secretion,” American Journal of Physiology-Endocrinology And Metabolism, vol. 295, no. 6, pp. E1287–E1297, 2008.
View at: Publisher Site | Google Scholar
R. B. Jordan, E. F. Walton, K. U. Klages, and R. J. Seelye, “Postharvest fruit density as an indicator of dry matter and ripened soluble solids of kiwifruit,” Postharvest Biology and Technology, vol. 20, no. 2, pp. 163–173, 2000.
View at: Publisher Site | Google Scholar
M. Li, F. Feng, and L. Cheng, “Expression patterns of genes involved in sugar metabolism and accumulation during apple fruit development,” PLoS One, vol. 7, no. 3, 2012.
View at: Publisher Site | Google Scholar
G. Vizzotto and R. Falchi, “Genetics of sugar and starch metabolism,” in The Kiwifruit Genome, pp. 189–204, Springer, Berlin, Germany, 2016.
View at: Publisher Site | Google Scholar
H. Velásquez-Arredondo, A. Ruiz-Colorado, and S. De Oliveira junior, “Ethanol production process from banana fruit and its lignocellulosic residues: energy analysis,” Energy, vol. 35, no. 7, pp. 3081–3087, 2010.
View at: Publisher Site | Google Scholar
M. V. Albertini, E. Carcouet, O. Pailly, C. Gambotti, F. Luro, and L. Berti, “Changes in organic acids and sugars during early stages of development of acidic and acidless citrus fruit,” Journal of Agricultural and Food Chemistry, vol. 54, no. 21, pp. 8335–8339, 2006.
View at: Publisher Site | Google Scholar
J. R. O. Do Nascimento, A. V. Júnior, P. Z. Bassinello et al., “Beta-amylase expression and starch degradation during banana ripening,” Postharvest Biology and Technology, vol. 40, no. 1, pp. 41–47, 2006.
View at: Publisher Site | Google Scholar
T. Maria, G. Tsaniklidis, C. Delis et al., “Gene transcript accumulation and enzyme activity of β-amylases suggest involvement in the starch depletion during the ripening of cherry tomatoes,” Plant Gene, vol. 5, pp. 8–12, 2016.
View at: Publisher Site | Google Scholar
J. H. Lee, D. J. Yu, S. J. Kim, D. Choi, and H. J. Lee, “Intraspecies differences in cold hardiness, carbohydrate content and β-amylase gene expression of Vaccinium corymbosum during cold acclimation and deacclimation,” Tree Physiology, vol. 32, no. 12, pp. 1533–1540, 2012.
View at: Publisher Site | Google Scholar
Y. Nolasco-González and J. Osuna-García, “Calidad postcosecha en mango ‘kent’ con fertilización normal y alta,” Revista Mexicana de Ciencias Agrícolas, vol. 8, no. 19, pp. 4009–4022, 2017.
View at: Publisher Site | Google Scholar
C. Huan, J. Zhang, Y. Jia et al., “Effect of 1-methylcyclopropene treatment on quality, volatile production and ethanol metabolism in kiwifruit during storage at room temperature,” Scientia Horticulturae, vol. 265, Article ID 109266, 2020.
View at: Publisher Site | Google Scholar
F. R. Thewes, A. Brackmann, R. de Oliveira Anese, V. Ludwig, E. E. Schultz, and M. R. P. Berghetti, “1-methylcyclopropene suppresses anaerobic metabolism in apples stored under dynamic controlled atmosphere monitored by respiratory quotient,” Scientia Horticulturae, vol. 227, pp. 288–295, 2018.
View at: Publisher Site | Google Scholar
R. D O. Anese, A. Brackmann, F. R. Thewes et al., “Impact of dynamic controlled atmosphere storage and 1-methylcyclopropene treatment on quality and volatile organic compounds profile of ‘Galaxy’ apple,” Food Packaging and Shelf Life, vol. 23, Article ID 100443, 2020.
View at: Publisher Site | Google Scholar
X. Yang, J. Song, L. Du et al., “Ethylene and 1-MCP regulate major volatile biosynthetic pathways in apple fruit,” Food Chemistry, vol. 194, pp. 325–336, 2016.
View at: Publisher Site | Google Scholar
S. Zhang, Y. Shi, and B. Deng, “Exogenous carbon monoxide treatment delayed the ethanol metabolism and fruit softening of postharvest jujube,” As J. Agric. Food Sci., vol. 3, no. 2, 2015.
View at: Google Scholar
J. Cheng, A. Zheng, H. Li et al., “Effects of melatonin treatment on ethanol fermenation and ERF expression in kiwifruit cv. Bruno during postharvest,” Scientia Horticulturae, vol. 293, Article ID 110696, 2022.
View at: Publisher Site | Google Scholar
X. Lu, G. Meng, W. Jin, and H. Gao, “Effects of 1-MCP in combination with Ca application on aroma volatiles production and softening of ‘Fuji’apple fruit,” Scientia Horticulturae, vol. 229, pp. 91–98, 2018.
View at: Publisher Site | Google Scholar
J. Chen, D. Zhang, X. Zhang et al., “Comprehensive responses of aroma production in ‘Benihoppe’strawberry to low oxygen associated with the changes of key gene expressions and energy levels,” Journal of the Science of Food and Agriculture, vol. 103, no. 4, pp. 1856–1863, 2023.
View at: Publisher Site | Google Scholar
X. Zuo, S. Cao, N. Ji et al., “High relative humidity enhances chilling tolerance of zucchini fruit by regulating sugar and ethanol metabolisms during cold storage,” Postharvest Biology and Technology, vol. 189, Article ID 111932, 2022.
View at: Publisher Site | Google Scholar
M. Miranda, X. Sun, C. Ference et al., “Nano-and micro-carnauba wax emulsions versus shellac protective coatings on postharvest citrus quality,” Journal of the American Society for Horticultural Science, vol. 146, no. 1, pp. 40–49, 2021.
View at: Publisher Site | Google Scholar
G. F. Velazco, F. Scaparoni, M. Sisquella et al., “Effects of different comercial coatings on postharvest citrus fruit quality for export,” Agro Urug, vol. 25, no. 1, pp. 1–22, 2021.
View at: Publisher Site | Google Scholar

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Copyright © 2023 Maratab Ali et al. This is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

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Journal of Food Biochemistry

Alcoholic Off-Flavor Disorders in Fresh Fruits

Abstract

1. Introduction

2. Ethanol Biosynthesis and Metabolism

2.1. Factors Leading to Upregulated Ethanol Metabolism in Fresh Fruits

2.2. Anaerobic Respiration

2.3. Low O2 Stress

2.4. High CO2 Stress

2.5. Storage Temperature Stress

3. Role of Physiological Mechanisms in Regulating Alcoholic Off-Flavor in Fresh Fruits

3.1. GABA Shunt Pathway

3.2. Mitochondrial Energy Metabolism

3.3. Glycolysis and TCA Cycle

3.4. Cytosolic Malate Metabolism

3.5. Starch and Sugar Metabolism

4. Measures to Control Alcoholic Off-Flavor Disorder Based on Changes in Ethanol Metabolism

4.1. Chemical Treatments

4.2. Hypobaric and Other Treatments

5. Conclusion and Future Work

Data Availability

Conflicts of Interest

Authors’ Contributions

Acknowledgments

References

Copyright

2.3. Low O₂ Stress

2.4. High CO₂ Stress