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Association Between Herbivore Resistance and Fruit Quality in Apple

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Association Between Herbivore Resistance and Fruit Quality in Apple HORTSCIENCE 46(1):12–15. 2011. (Kaplan et al., 2009; Koricheva et al., 1998) but often have a limited capacity for effective defense against pests as a result of allocation Association between Herbivore of resources to persistent plant organs and because of their relatively low intimacy of Resistance and Fruit Quality in Apple association with the economically most dam- 1 2 aging pests (Coley et al., 1985; Mattson et al., Sibylle Stoeckli , Karsten Mody , and Silvia Dorn 1988). Expression of plant defenses would, ETH Zurich, Institute of Plant Sciences/Applied Entomology, Schmelzbergstrasse however, be particularly important for peren- 9/LFO, CH-8092 Zurich, Switzerland nial crops such as apple orchards. Plants in such perennial systems are confronted with Markus Kellerhals a notable diversity of pests but also with ef- Agroscope Changins-Waedenswil Research Station ACW, P.O. Box, Schloss, fective natural enemies, which can be sup- CH-8820 Waedenswil, Switzerland ported by adequate pest management schemes incorporating plant defense to reduce pesticide Additional index words. Malus ·domestica, marker assisted selection, fruit quality, herbivore use (Beers et al., 2003; Dorn et al., 1999; resistance, resource allocation tradeoff Zehnder et al., 2007). Disease-resistant cultivars often have un- Abstract. Enhanced fruit quality, plant health, and productivity are major objectives in desirable fruit traits (Kellerhals et al., 2004b), apple breeding. The undesirable fruit quality traits frequently associated with pest- and which underlines a possible resource alloca- disease-resistant cultivars may be related to resource allocation tradeoffs. The objective tion tradeoff. Evaluation of fruit quality traits of the present study was to evaluate the association between insect resistance and fruit is highly relevant because the fruit is the quality in apple. The studied ‘Fiesta’ · ‘Discovery’ apple progeny was characterized by marketable product of an apple tree, and breed- reasonable fruit firmness and optimal sugar content and acidity but small fruit size. ing for disease and pest-resistant apple cultivars There was a positive correlation between codling moth (Cydia pomonella) fruit infes- should always be performed under consider- tation and fruit firmness. Additionally, a positive correlation was detected between shoot ation of marketable fruit quality (Brown and infestation by green apple aphid (Aphis pomi), fruit number as well as sugar content. In- Maloney, 2003). The objective of the present festation by the apple leaf miner moth (Lyonetia clerkella), the rosy apple aphid (Dysaphis study was to evaluate the association between plantaginea), the leaf-curling aphid (Dysaphis cf. devecta), and the apple rust mite (Aculus insect resistance and different fruit quality traits schlechtendali) was not significantly related to fruit quality traits. The positive relation- in apple. Enhanced knowledge on resource ship of increased infestation by some pest insects and quality-determining fruit allocation tradeoffs in apple may be considered characteristics such as firmness or sugar content points to a possibly increased necessity in breeding programs aiming at high-quality for plant protection measures in apple cultivars producing high-quality fruits. One disease- and pest-resistant apples. possible explanation of higher pest infestation in cultivars producing fruits with high quality is a tradeoff between resource allocation to defensive secondary metabolites or to fruit quality. By identifying a relationship between pest infestation and fruit quality, the present study highlights the need to consider pest resistance when breeding for high- Materials and Methods quality apple cultivars. The use of genetic markers for fruit quality and pest resistance in Plant material and study sites. Herbivore marker-assisted breeding may facilitate the combined consideration of fruit quality and resistance and fruit quality were assessed on pest resistance in apple breeding programs. a ‘Fiesta’ · ‘Discovery’ (Malus ·domestica Borkh.) F1 progeny. Progeny genotypes were bud-grafted on M27 rootstocks, and each of Host-plant resistance is a principal com- firmness, acidity, size, and color (Eigenmann 250 genotypes was planted in winter 1998– ponent of the management of pests and and Kellerhals, 2007). Preference is given to 1999 at three study sites in Switzerland diseases in an integrated cropping system fruit that is crisp (firmness 8 kgÁcm–2), sweet (Liebhard et al., 2003): Cadenazzo (Ticino; (Kellerhals, 2009). In addition to enhanced (sugar content 12 to 13 °Brix), and medium- lat. 46°09#35$ N, long. 8°56#00$ E; 203-m plant health, crop yield, tree architecture, sized (diameter 70 to 80 mm) with excellent altitude), Conthey (Valais; lat. 46°12#30$ N, storability, and consumer preference are the color and shelf life (Egger et al., 2009b). In long. 7°18#15$ E; 478-m altitude), and Wae- major aspects in apple breeding (Beers et al., apple (Malus ·domestica Borkh.), several denswil (Zurich; lat. 47°13#20$ N, long. 2003; Egger et al., 2009a, 2009b). Desirable quantitative trait loci (QTLs) or major genes 8°40#05$ E; 455-m altitude). The distance fruit traits include flavor, juiciness, sweetness, for disease (Calenge and Durel, 2006; Khan between trees was 0.5 m in Conthey and et al., 2006) and pest resistance (Bus et al., Waedenswil and 1.5 m in Cadenazzo. Trees 2008; Roche et al., 1997; Stoeckli et al., were planted in rows 3.5 m apart from each 2008b, 2008c, 2009; Wearing et al., 2003) other. Climate conditions at the three sites Received for publication 11 Aug. 2010. Accepted as well as fruit quality traits (Cevik et al., were characterized by highest temperature for publication 29 Sept. 2010. 2010; Costa et al., 2008; Liebhard et al., in Cadenazzo (mean annual temperature: We thank Cesare Gessler (ETH Zurich), Mauro 2003) have been identified. This highlights the 10.5 °C, total annual rainfall: 1772 mm; 30- Jermini, Danilo Christen, and Reto Leumann potential usefulness of marker-assisted selec- (Agroscope Changins-Waedenswil Research Sta- year average; MeteoSwiss) and lowest tion ACW) for access to and maintenance of the tion in targeted apple breeding (Baumgartner amount of rainfall in Conthey (9.2 °C, 598 study orchards; Michelle Schmocker, Christoph et al., 2010; Kellerhals, 2009). mm) compared with Waedenswil (8.7 °C, Rohrer, Claudia Good. and Sandra Noser for help The most basic energy resources of green 1353 mm). Orchards were treated with fer- with fieldwork; Caroline Baumgartner for help plants are carbohydrates, proteins, and lipids. tilizers and herbicides, but no insecticides with mite filtration; Lorenz Kreis (Genossenschaft The allocation of resources to plant growth and fungicides were applied. Fruit thinning Migros Ostschweiz) for fruit trait determination; (e.g., tree growth, fruit yield, and quality) is was carried out by hand. In 2009, insecticide Robert Liebhard (Tecan AG) for access to the fruit costly and competes with the production of and fungicide treatments were carried out to quality data set; and Adriana Najar-Rodriguez for defensive compounds (e.g., phenolics, terpe- achieve acceptable fruit quality. useful comments to an earlier version of the article. noids, or glucosinolates) (Gayler et al., 2004; 1Present address: Agroscope Changins-Waedens- Herbivore resistance. Herbivore infesta- wil Research Station ACW, P.O. Box, Schloss, Koricheva, 2002). This tradeoff between de- tion per tree was used as a measure of CH-8820 Waedenswil, Switzerland. mands for growth and defense is especially resistance to two lepidopteran, three aphid, 2To whom reprint requests should be addressed; relevant in perennial crop plants. These crops and one mite species. Herbivore assessment e-mail [email protected]. are bred and grown for maximized yield was carried out on 160 apple genotypes in 12 HORTSCIENCE VOL. 46(1) JANUARY 2011 | BREEDING,CULTIVARS,ROOTSTOCKS, AND GERMPLASM RESOURCES study Years 1 and 2 (2005 and 2006). The Statistical analysis. Spearman’s rank cor- determined for 50% of the randomly chosen number of codling moth (Cydia pomonella relation test was used to analyze the associ- fruits. The corresponding percentiles for L.) larval penetrations in fruits was inspected ation between herbivore resistance and fruit sugar content were 10 and 11°Brix and for only on fruits attached to the tree, not on quality, and the Benjamini-Hochberg correc- acidity 7 gÁL–1 and12gÁL–1 malic acid, fallen fruits (Stoeckli et al., 2009). All leaf tion for Type I errors in multiple tests was respectively. Mean fruit number, fruit weight, mines of the apple leaf miner (Lyonetia applied (Verhoeven et al., 2005). All statis- fruit firmness, sugar content, and acidity were clerkella L.) were counted for each individ- tics were carried out with R 2.7.0 for Mac not significantly correlated with each other ual tree by examining every leaf at the end of OS X (http://www.stat.ethz.ch/CRAN/). (P > 0.05; Spearman’s rank test). July in 2005. At this time of the year, mines Fruit trait comparison between sites and from all generations (two to three per year) Results across years. Fruit firmness was significantly should be visible and the quantified infesta- correlated between the site Waedenswil tion level should therefore be representative Herbivore abundance. Mean RI of the (study Year 3) and the two other study sites for the whole year (Baggiolini et al., 1992). progeny plants (average from several surveys (Year 2000; Cadenazzo: n = 33, rs = 0.588, The rosy apple aphid (Dysaphis plantaginea covering two to three study sites and 2 P < 0.001; Conthey: n = 22, rs = 0.512, P = Pass.) was assessed by evaluation of the consecutive years) was 0.7 ± 0.1 (all study 0.015; Spearman’s rank test). Apple fruit in number of colonies (Stoeckli et al., 2008b). species) and ranged between completely Cadenazzo and Conthey were comparable The number of red-curled leaves was taken uninfested trees (RI = 0%; all herbivores considering sugar content (n = 93, rs = as a measure of leaf-curling aphid (Dysaphis species) and 10.6% (D. plantaginea), 8.7% 0.522, P < 0.001; Spearman’s rank test), but cf.
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