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This Article Appeared in a Journal Published by Elsevier This article appeared in a journal published by Elsevier. The attached copy is furnished to the author for internal non-commercial research and education use, including for instruction at the authors institution and sharing with colleagues. Other uses, including reproduction and distribution, or selling or licensing copies, or posting to personal, institutional or third party websites are prohibited. In most cases authors are permitted to post their version of the article (e.g. in Word or Tex form) to their personal website or institutional repository. Authors requiring further information regarding Elsevier’s archiving and manuscript policies are encouraged to visit: http://www.elsevier.com/authorsrights Author's personal copy Postharvest Biology and Technology 82 (2013) 51–58 Contents lists available at SciVerse ScienceDirect Postharvest Biology and Technology jou rnal homepage: www.elsevier.com/locate/postharvbio Impact of harvesting time and fruit firmness on the tolerance to fungal storage diseases in an apple germplasm collection a,∗ b a Masoud Ahmadi-Afzadi , Ibrahim Tahir , Hilde Nybom a Balsgård, Department of Plant Breeding, Swedish University of Agricultural Sciences, Fjälkestadsvägen 459, SE-29194 Kristianstad, Sweden b Department of Plant Breeding, Swedish University of Agricultural Sciences, Box 101, SE-23053 Alnarp, Sweden a r t i c l e i n f o a b s t r a c t Article history: Blue mold and bitter rot, caused by Penicillium expansum and Colletotrichum gloeosporioides, respectively, Received 16 October 2012 are two of the most devastating diseases during and after storage of apple. The present project was Accepted 3 March 2013 conducted to evaluate the level of tolerance to these diseases in apple germplasm, and investigate possible associations with other fruit characteristics such as harvest date, firmness at harvest, softening (loss Keywords: of firmness during storage) and sun-exposure. Apples were harvested at a maturation stage suitable Malus × domestica for storage, inoculated with spore suspensions of P. expansum (127 cultivars) or C. gloeosporioides (70 Blue mold cultivars), and stored for 6 or 12 weeks for early- and late-maturing cultivars, respectively. Fruit firmness Bitter rot was measured after harvest and after storage, and the difference was used as a measure of fruit softening. Fruit texture PLS-DA Average lesion diameter varied significantly among both early- and late-maturing cultivars. The amount of damage caused by the two diseases was significantly correlated across cultivars. Regression analyses indicated that lesion diameter was positively affected by fruit softening and negatively affected by harvest date and firmness at harvest. Impact of the independent variables was quantified with partial least squares discriminant analysis; approximately 40% of the genetic variation could be explained by these variables with harvest date being the most important. The effect of sun-exposure was analyzed on six bi-colored cultivars but the results were not conclusive. Cultivars that showed relatively small symptoms in spite of being early-maturing and/or only medium firm, may have other traits that are beneficial for storage and could therefore be especially useful in breeding programs. © 2013 Elsevier B.V. All rights reserved. 1. Introduction after a few days at room temperature (Jones and Aldwinckle, 1990). Presently, increasing concerns for environmental and health issues × Apple (Malus domestica Borkh.) is one of the economically limit the access to chemical compounds for efficient plant pro- most important fruit crops, and can have a strong impact on human tection. Moreover, prohibition of postharvest treatments in some health due to its high accessibility, comparatively low price and countries has resulted in increasingly serious problems with stor- high levels of antioxidant and phenolic compounds. Apples, in addi- age diseases (Jones and Aldwinckle, 1990; Gullino and Kuijpers, tion to being processed into juice, sauce, slices, vinegar or cider, 1994; Janisiewicz and Korsten, 2002). These problems are even are mainly consumed fresh or after storage (Folta and Gardiner, more pronounced in organic production; organically produced 2009). Presently, much of the commercially grown fruit is stored ‘Aroma’ fruit in Sweden suffered a 20-fold increase in fungal decay for 4 months or longer before being sold, and very good storage compared to conventionally grown fruit (Jönssson et al., 2010). potential is therefore an important requirement for apple cultivars Consequently, storage time must be substantially shortened for (Ferguson and Boyd, 2002). organically grown fruit, resulting in lower availability for con- Fungal diseases cause a wide array of postharvest damage on sumers and serious economic losses for growers. fruit, thus restricting storage potential and constituting one of Blue mold caused by Penicillium expansum is one of the most the main problems for commercial apple production. Attacks can common postharvest diseases of apple. This fungus produces be initiated either during the growing season or at harvest and patulin, a carcinogenic mycotoxin which has attracted public con- postharvest handling, but symptoms are seldom visible until after cern due to its potential impact on human health (Beretta et al., a certain storage period and, for some diseases, they appear only 2000; Pianzzola et al., 2004; Barreira et al., 2010). Bitter rot, another important storage disease on fruit caused by Colletotrichum gloeosporioides and Colletotrichum acutatum, is both a pre- and ∗ a postharvest disease, especially under hot and humid condi- Corresponding author. Tel.: +46 445806; fax: +46 445830. tions, and can result in 50–80% losses in apples harvested from E-mail addresses: [email protected] (M. Ahmadi-Afzadi), [email protected] (I. Tahir), [email protected] (H. Nybom). orchards without effective plant protectant spraying (Jones and 0925-5214/$ – see front matter © 2013 Elsevier B.V. All rights reserved. http://dx.doi.org/10.1016/j.postharvbio.2013.03.001 Author's personal copy 52 M. Ahmadi-Afzadi et al. / Postharvest Biology and Technology 82 (2013) 51–58 Aldwinckle, 1990; Jurick et al., 2011). Concerns about posthar- 2.2. Analysis of level of resistance in apple germplasm vest fungal diseases, in particular blue mold and bitter rot, have increased significantly during the last decades, possibly due to a On the day of harvest or the following day, fruit were washed combination of global warming, increased organic production, pro- with distilled water in order to remove most of the naturally hibition of several previously used fungicides and the increasing occurring fungi (as previously demonstrated by Tahir et al., 2009), levels of resistance, e.g., P. expansum toward fungicides (Vinas˜ et al., wiped dry and then inoculated on opposite sides at a depth 5 −1 1991; Sholberg et al., 2005; Tahir and Jönsson, 2005; Weber, 2009). of 3 mm with 20 ␮L (1 × 10 conidia mL ) of P. expansum and Unfortunately, most of the currently grown apple cultivars are C. gloeosporioides, respectively. The pathogens used were iso- very sensitive to storage diseases. So far, no major genes provid- lated from naturally infected apples showing typical symptoms ing resistance against the storage diseases have been identified but of the diseases, maintained on petri dishes with potato dex- quantitatively inherited traits associated with chemical contents, trose agar augmented with penicillin G and streptomycin sulphate −1 fruit texture, structure of the fruit epidermis and ripening behav- (each at 200 mg L agar), and stored separately as pure cul- ◦ ior may affect the ability of different cultivars to withstand fungal tures at 4 C. Pathogen virulence over time was confirmed by attacks (Prusky et al., 2004; Blazek et al., 2007; Nybom et al., 2008; periodic transfers through apples. Spores of the pathogens were Johnston et al., 2009). Positive effects of increased firmness and removed from the surface of 10-day-old cultures and suspended less softening due to calcium spraying, have been documented in a in 5 mL sterile distilled water containing 0.05% (v/v) Tween 80. series of pre- and postharvest treatments of apple fruit, indicating The suspensions were filtered, and spore concentrations were 5 −1 × that fungal decay caused by P. expansum, Botrytis cinerea and Glom- adjusted to 1 10 conidia mL (Tahir et al., 2009; Weber and erella cingulata can be significantly reduced (Conway et al., 1991; Palm, 2010). Three replicates with 15 fruit in each were prepared Sams et al., 1993; Conway et al., 2002). with each cultivar-fungus combination, as well as two controls Variable levels of resistance to fungal decay have already been (fruit with no treatment, and wounded but not inoculated fruit, reported in some studies. Thus, evaluation of the resistance to blue respectively). mold in an apple germplasm collection from Kazakhstan, main- After inoculation, the fruit were stored in open-faced plastic ◦ tained in Geneva, NY, indicated the existence of greater genetic boxes at 2 C for either 6 weeks (2010: summer cultivars har- diversity among the Kazak apples than among cultivated apples vested from August 10 to September 9; 2012: from August 9 to (Janisiewicz et al., 2008). In another study, resistance to both blue September 7) or 12 weeks (2010: fall and winter cultivars har- mold and bitter rot was identified in wild apple germplasm from vested from September 13 to October 26; 2012: from September Kazakhstan (Jurick et al., 2011). 9 to October 9). Two alternatives for duration of storage were used Genetic differences in tolerance to storage diseases have also since early-maturing cultivars could have deteriorated and become been reported in some screenings of commercial cultivars (Spotts difficult to evaluate if stored longer than 6 weeks whereas late et al., 1999; Biggs and Miller, 2001) but data from large-scale cultivars might not have reached the climacteric stage if stored screenings, especially of cultivars suitable for a cooler climate, are less than 12 weeks, thereby making it difficult to observe dif- lacking. The aims of this study were (1) to evaluate the level of tol- ferences in firmness and disease incidence.
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