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Abstract The effectiveness of initial low oxygen stress (ILOS) to control apple scald without postharvest application of scald inhibitors and fungicides was investigated. Scald susceptible (Granny Smith, Law Rome, Red Delicious) and not susceptible (Idared) fruits were harvested preclimacterically and treated at 1C with (1) ILOS pretreatment or (2) no pretreatment as control. Fruit were then stored at 0.5 1C in 3% O2 with 0% CO2, 1.5% O2 with 3% CO2, or 0.70.8% O2 with 3% CO2 in ow-through controlled atmosphere (CA) or in air. ILOS of 0.5 and 0.25% O2 for 2 weeks resulted in excellent control of scald when fruits were subsequently held in CA (3% O2, 0% CO2). Initial treatment with 0.25% O2 for 2 weeks, and when this treatment was supplemented with an additional 2 weeks of low O2 stress after 2 months of storage, were more effective treatment regimens for scald control than all other treatments. ILOS followed by CA storage at 1.5% O2 gave complete control of scald. A commercial test of initial low O2 stress conrmed its efcacy for controlling scald in several cultivars of apples. The production of a-farnesene and its volatile oxidation product 6-methyl-5-hepten-2-one (MHO) was inhibited by initial low O2 stress treatments and 1.5% O2 CA. The 0.25% O2 initial stress treatment caused stronger inhibition of a-farnesene and MHO production than 0.5% O2 initial stress treatment. The accumulation of MHO was related to scald development of apples. 2000 Elsevier Science B.V. All rights reserved.
Keywords: Apple; Supercial scald; Initial low O2 stress; Volatiles; a-Farnesene; 6-Methyl-5-hepten-2-one
1. Introduction Supercial scald of apples is a physiological disorder which, following long-term storage, has the potential to destroy the market value and utility of millions of tons of fresh apples annually (Wang and Dilley, 1999). The disorder is pre* Corresponding author. Tel.: + 1-517-3553286 or + 1-5173557483; fax: + 1-517-3530890. E -mail addresses: wangzhe1@pilot.msu.edu (Z. Wang), dilley@pilot.msu.edu (D.R. Dilley)
vented commercially by postharvest drench treatment with diphenylamine (DPA) (Smock, 1957). This drench treatment must include a fungicide to prevent decay development. Numerous countries have banned the use of DPA or importation of DPA-treated fruits, which mandates development of alternative methods to control scald. It has been proposed that apple fruit may possess at harvest or may acquire after harvest resistance to scald (Gallerani et al., 1990; Barden and Bramlage, 1994a,b). It is speculated that the skin browning associated with the disorder results
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from autooxidative processes that disrupt cell membranes and are brought about by depletion of endogenous antioxidants such as ascorbic acid or a-tocopherol during low temperature storage (Meir and Bramlage, 1988). Free-radicals such as hydroxyl (OH), hydroperoxyl (HO2) and superoxide (O 2 ) can attack and destroy lipid and protein macro molecular structures essential for cellular function (Anet, 1969; Halliwell and Gutteridge, 1989; Stadtman, 1993) and subsequently lead to scald symptom development. Extensive investigations in Australia (Anet and Coggiola, 1974 and references cited therein) indicated that scald results from the autoxidation of a volatile sesquiterpene hydrocarbon a-farnesene (2,6,10trimethyl-2,6,9,11-dodecatetraene) in the fruit skin (Meigh and Filmer, 1969). Based on in vitro studies, oxidation of a-farnesene was hypothesized to yield conjugated triene hydroperoxide free-radicals which injure the hypodermal cells and give rise to the symptoms of scald (Anet, 1969, 1972). However, Rowan et al. (1995) found that in vivo only trace amounts of these were detectable; the major oxidation products that accumulated in apple peel were two 2,7,8,11-tetraen6-ol (conjugated trienol) isomers. Scald development in Granny Smith, a cultivar very susceptible to scald, was correlated to the oxidation of a-farnesene to conjugated triene hydroperoxides (Huelin and Coggiola, 1970; Filmer and Meigh, 1971). Song and Beaudry (1996) found that 6-methyl-5-hepten-2-one (MHO), a volatile product of a-farnesene oxidation, caused a scald-like disorder when applied to peel tissue of susceptible apple fruit. Scald can be induced by the conjugated triene (CT) products of a-farnesene oxidation, and the concentration and time of appearance of these products is related to the severity of the disorder (Anet, 1972). The scald inhibitor chemicals DPA and ethoxyquin are effective inhibitors of lipid peroxidation, and ethanol vapor which controls scald (Scott et al., 1995a; Wang and Dilley, 1996, 1997; Wang et al., 1997; Ghahramani and Scott, 1998a) is a free-radical scavenger (Halliwell and Gutteridge, 1989). Little et al. (1982) reported that scald of Granny Smith apples was largely controlled by an initial low O2 stress followed by CA storage. Low O2
could exert its action partly by increasing endogenous ethanol production (Wang et al., 1997; Ghahramani and Scott, 1998b). We have tested the concept that initial low O2 stress prior to CA storage controls scald by inducing production of ethanol and inhibiting a-farnesene synthesis and its oxidation to yield MHO. Initial low O2 stress treatments were investigated to determine effective treatment regimens and to gain insight into the mechanism of scald control.
2.1. Initial low oxygen stress followed by CA storage 2.1.1. Plant materials and ILOS treatments Preclimacteric Law Rome apples were obtained from the Michigan State University, Clarksville Horticultural Experiment Station (MSU CHES) and Granny Smith apples were obtained from a grower in southwest Michigan, about 2 weeks prior to the onset of autocatalytic ethylene production. Forty to 60 fruit were placed in 20-l CA chambers in each of three replicates for each cultivar and storage period evaluation. Two levels of initial low O2 stress were employed: 0.5 and 0.25% O2 for 2 weeks at 1C. The control fruit did not receive low O2 stress. The low O2 stress atmospheres were generated with an air separator (Permea, Model 5, Permea, St Louis, MO) and distributed through a capillary ow board system for maintaining controlled atmospheres in a ow-through system. After the low O2 stress, the fruit were stored in dynamic (slowly ventilated) CA under 3 or 1.5% O2 at 1C, using a ow rate of 0.18 void volume exchanges per h. Some fruit so treated were again subjected to the same low O2 stress treatments after 2 months in CA. Other groups of fruit were stored in air or in 3 or 1.5% O2 CA without prior low O2 stress. 2.1.2. Scald index measurement Examination of fruit for scald index was made after 5 and 8 months of storage plus 1 week post-storage in air at 20C. Scald data are expressed as a scald index based on the percentage
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of fruit surface area affected, where no scald = 0, B 25% = 1, 2550% = 2, and \ 50% = 3. The scald index was normalized to 100 by multiplying the values by 100/3.
2.1.3. Analysis of 6olatiles To determine the effects of ILOS on ripening, scald development, and production of a-farnesene and its oxidation product MHO, fruit were removed from storage at appropriate intervals and transferred to 20C in air. Solid phase microextraction/gas chromatography/mass spectrometry (SPME/GC/MS) was employed to identify and quantify the volatiles including a-farnesene and MHO as described by Song et al. (1997). Five to eight fruit from each treatment were enclosed in 4-l glass jars at 20C for 1.5 h. The headspace was sampled by SPME equilibrating for 4 min to absorb volatiles and analyzed as described previously. At the same time intervals, samples (14 mg) of epicuticular wax were removed from the Granny Smith fruit and placed in glass vials (2 ml) for 3 h at 20C to determine the amount of a-farnesene and MHO partitioned in the epicuticular wax by SPME/GC/MS as described previously. 2.2. Continuous low O2 stress
Granny Smith apples as used for the ILOS studies were employed. Fruit were placed in 10-l glass desiccators at 1C and ventilated with 0.13, 0.25, 0.4, 0.5, 0.75, 1.0, 1.25 or 1.5% O2 or with air (20.7% O2). The low O2 atmospheres were made and distributed as described above. The ventilation rate through the chambers was 54 ml min 1 (approximately 0.6 void volume exchanges per h). Volatiles analyses commenced 1 week after the low O2 levels were established and continued at weekly intervals throughout the 8-month storage period. The volatiles prole of more than 60 compounds produced by intact fruit in a ow through system at 1C and while in various storage atmospheres was obtained. Fruit were examined for scald and other physiological disorders after removal from storage and again after 7 days at 20C.
3. Results
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gen level of ILOS progressively diminished the incidence and severity of scald in Granny Smith and Law Rome fruit subsequently stored in CA at 3% O2 without CO2; whereas fruit provided ILOS and stored at 1.5% O2 with CO2 did not develop a signicant level of scald (Fig. 1A,B). These results conrmed our studies in 1996/97 in which Granny Smith, Law Rome and Red Delicious apples stored in air at 1C without
ILOS developed severe scald and those held in CA storage at 3% O2 exhibited only partial control. ILOS at 0.5% O2 for 1 or 2 weeks with subsequent storage in CA at 3% O2 largely attenuated scald and 2 weeks at 0.5% O2 was more effective than 1 week (Wang et al., 1997). In the 1996/97 tests, CA storage of ILOS-treated fruit at 1.5% O2 completely controlled internal browning in both cultivars, whereas CA storage at 3% O2
Fig. 1. Initial low oxygen stress attenuates scald of Granny Smith and Law Rome apples stored in CA (3% O2 + 0% CO2 or 1.5% O2 + 3% CO2) for 5 months (A) and 8 months (B) at 0.5C plus 7 days at 20C in air. Scald index (0, none; 1, slight; 2, moderate; 3, severe) was normalized to 100 by multiplying the values by 100/3.
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did not. Internal browning incidence (probably senescence-related) in Granny Smith fruit was 74 and 49% for fruit stored in air and 3% O2, respectively, and this was reduced by about one-half by ILOS followed by CA at 3% O2 and was not seen in fruit from 1.5% O2 with or without ILOS. Also core browning (a slight discoloration of tissue immediately adjacent to the seed cavity that is related to immaturity at harvest) was seen in Law Rome subjected to the greatest degree of O2 stress and only when stored at 1.5% O2 CA (57%). Slight core browning in Granny Smith was not related to CA treatments. The incidence of decay was most prevalent in the air control fruit for both cultivars (11 and 15%, respectively) and was reduced to about 2% by ILOS treatments. ILOS treatments resulted in high retention of esh rmness of both cultivars when followed by CA at 1.5% O2. Initial esh rmness was 82 and 89 N for Granny Smith and Law Rome respectively, and for ILOS followed by CA at 1.5% O2 was 76 and 80 N and for 3% O2 was 70 and 65 N, respectively. Granny Smith apples without ILOS produced more a-farnesene after 8 months of storage in 1.5% O2 CA than those with ILOS treatments (Fig. 2) and also accumulated more a-farnesene in the epicuticular wax (Wang, 1998). Control fruit stored in 3% O2 produced lower amounts of a-farnesene than ILOS treated fruit (Fig. 2). Apples with ILOS at 0.25% O2 produced about one-half as much a-farnesene than those with 0.5% ILOS treatment after 5 and 8 months in 1.5% O2 storage. Fruit receiving 0.25% O2 ILOS treatments produced about one-half as much a-farnesene and MHO as those with 0.5% O2 stress for all CA storage conditions for both Granny Smith and Law Rome cultivars in air at 20C after 5 months of storage (Wang, 1998). In contrast, after 8 months of storage, the potential of fruit to produce MHO with 0.25% O2 initial stress treatments had increased for 3% O2, but not 1.5% O2 storage (Fig. 3). This indicates that the a-farnesene production of apples was inhibited by ILOS treatments and also that ILOS at 0.25% O2 and/or 1.5% O2 CA more strongly inhibited a-farnesene production than storage without ILOS and/or CA at the 3% O2 level. Fruit given the
combination of 0.25% O2 ILOS treatment and CA at 1.5% O2 had an approximately threefold lower content of MHO than did 0.5% O2 ILOS and CA at 3% the O2 level.
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Fig. 2. Effect of initial low oxygen stress on a-farnesene levels in the headspace of Granny Smith (A, B) and Law Rome (C, D) apples stored in 1.5% (A, C) and 3% (B, D) O2 CA for 8 months at 0.5C.
age period (Fig. 5C). MHO was not detected at O2 levels below 1.25% and was generally detectable only at 1.5% O2 or above. The relationship observed between the levels of O2 favoring ethanol production and reduced levels of MHO suggest that ethanol metabolism may be directly related to keeping MHO at low levels. Alterna-
tively, MHO production may involve an oxidative enzyme with a fairly low afnity for molecular O2. The mechanism for scald control by ILOS remains to be resolved. Scald developed on the air-stored fruit after 4 months of storage and on the 1.5% O2 stored fruit after 10 months of storage (data not shown.)
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sied after a week at 20C. Fruit from ultra low CA (0.7% O2 throughout) were scald-free (Fig. 6). Some Red Delicious and Law Rome fruit had pigment purpling on removal from 0.7% O2 storage but this largely cleared after a few days in air at 20C. All cultivars from CA room c 2 (1.5% O2 throughout) exhibited scald of varying degrees with an overall scald index average of 0.52. Fruit
Fig. 3. Effect of initial low oxygen stress on 6-methyl-5-hepten-2-one (MHO) levels in the headspace (A, B) and epicuticular wax (C, D) of Granny Smith apples stored in 1.5% (A, C) and 3% (B, D) O2 CA for 8 months at 0.5C.
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Fig. 4. Change in production rates of ethanol (A), ethyl acetate (B) and acetaldehyde (C) in Granny Smith apples at low levels of oxygen and in air storage at 0.5C.
from CA room 3 and 4 (ILOS for 18 days at 0.55 and 0.61% O2, respectively) had very low scald index values of 0.09 and 0.03. Galia Beauty Rome was the only cultivar with appreciable scald (data not shown). These fruit were very immature at harvest.
4. Discussion
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al., 1982; Van der Merwe et al., 1997; Wang et al., 1997; Ghahramani and Scott, 1998b) although other investigators nd that ILOS is sometimes ineffective (Truter et al., 1994). The level and duration of ILOS as well as the temperature and the subsequent O2 regimens during CA may be important factors determining the efcacy of ILOS. The treatment regimen of ILOS prior to 1.5% O2 CA storage resulted in excellent retention of fruit rmness and overall quality while CA
storage at 3% O2 following low O2 stress did not. The decline in the incidence of scald as the degree of initial low O2 stress increased is consistent with scald reduction obtained following ethanol vapor treatment and suggests that the effects may be related to some aspect of ethanol metabolism. That apples produce ethanol when the O2 level is below the extinction point of alcohol fermentation (ca 1%) suggests that a storage regimen of initial low O2 stress may induce apples to produce
Fig. 5. Change in production rates of a-farnesene (A,B) and its oxidation product MHO (C) in Granny Smith apples at low levels of oxygen and in air storage at 0.5C (fruit did not produce MHO when O2 was lower than 1.25%).
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ILOS treatment may be a consequence of the ethanol produced and/or metabolism induced to reduce the concentration of some metabolites or ameliorate their actions. CA storage following ILOS, at O2 levels known to reduce scald, may be expected to further attenuate scald incidence and severity.
Fig. 6. Effects of initial low oxygen stress and oxygen levels in CA storage (1.5% O2 + 0.5% CO2) at the MSU CHES facility on controlling scald of apples stored for 8 months at 0.1C plus 7 days at 20C in air. Treatment designation: A, 1.5% O2 CA as control; B, 0.55% O2 initial stress for 18 days then 1.5% O2 CA; C, 0.61% O2 initial stress for 18 days then 1.5% O2 CA; D, 0.7% ultra-low O2 CA continuously. Scald index (0, none; 1, slight; 2, moderate; 3, severe) was normalized to 100 by multiplying the values by 100/3.
ethanol and be an effective means to control scald even after the O2 level is raised above the extinction point during subsequent storage. Apples receiving continuous 0.25% O2 stress produced ethanol (as measured in the atmosphere exiting slowly ventilated chambers of fruit) soon after the O2 level was reduced below the extinction point as would be expected (Fig. 4). The internal ethanol level, although not measured in this experiment, would rise in advance of ethanol being detected in the gas atmosphere surrounding the fruits. Moreover, initial ethanol vapor treatment reduced scald (Scott et al., 1995a; Wang et al., 1997) in apples subsequently stored at reduced O2 levels and this was also observed even when fruit were subsequently stored in air (Wang and Dilley, 1996). Thus, a transient exposure of apples to ethanol vapor applied exogenously imparts a residual scald-reducing effect. Reduction of scald incidence by the
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gas mixtures at a ow rate that is just sufcient to remove CO2 produced in respiration and provide a constant CO2 and O2 level. This amount of fresh atmosphere ushing is apparently sufcient to remove the trace amounts of scald-related volatile substances that may partition into the epicuticular wax (Matich et al., 1998). Such substances may include a-farnesene and 6-methyl-5-hepten-2-one derived from the biological oxidation of a-farnesene (Filmer and Meigh, 1971; Spicer et al., 1993). Ventilation would not be expected to favor loss of conjugated trienols derived from a-farnesene (Whitaker et al., 1997). In theory, removal of volatiles by use of a scrubber is feasible. However, the ability of mineral oil impregnated wraps (Brooks et al., 1919) and vegetable oil coatings (Scott et al., 1995b) to reduce scald is thought to be a consequence of volatiles partitioning into them. Our data are consistent with our observation that fruit do not scald during 8-month storage when ow-through CA is employed at 1.5% O2 with 3% CO2. Moreover, scald is largely controlled in commercial storage at 1.5% O2 with 3% CO2 employing air separators to remove CO2 by ushing (Dilley, 1990). ILOS of ca 0.25 0.5% O2 for 14 days at 0C followed by warming to 2C for 3 days can effectively control scald of apples not treated with DPA when fruit are subsequently stored at 1.5% O2 at 0C employing a static CA regimen. CA storage at 0.7% O2 completely controlled scald (Fig. 6) conrming the results of Lau (1990). However, fruit of some cultivars exhibited symptoms of anaerobic fermentation damage. Since initial low O2 stress followed by CA at 1.5% O2 controlled scald effectively and maintained it at commercially acceptable levels, this CA storage regimen may provide a safe and effective means to store apples not treated with DPA. Avoiding DPA drenching would also avoid the use of postharvest fungicides, which is an issue of growing concern. Our present results support the concept that the benecial effects of ILOS may be related to ethanol metabolism in some manner. A similar conclusion was recently reported by Ghahramani and Scott (1998b) who suggested that control of scald by ILOS may be due to increased production of ethanol and consequent reduction in
a-farnesene and conjugated triene accumulation. The residual effect of ethanol vapor treatment in reducing scald of apples subsequently stored in air (Wang and Dilley, 1996) suggests that ethanol may act apart from the effect of low O2 alone.
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