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Volatile organic compounds (VOCs) are funda-mental elements of flavor, one of the most important fruit-quality traits. Despite its importance, this aspect is still poorly considered in assisted breeding programs, due to the lack of suitable and fast detection systems as well as validated func-tional markers. In this work, a full-sib parental mappin...
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... blend of aroma compounds in apple is subjected to important modifications of several fold change during the final phase of fruit ripening, as well as in postharvest, as reported by Fellman et al. (2003), Newcomb et al. (2006), and Schaffer et al. (2007). As aroma is an important trait for consumer appreciation, and thus marketability, we also performed a VOC survey over an apple collection represented by 124 accessions analyzing cut fruit instead of whole intact apples. To have a better idea of volatile change among these conditions, fruits from two apple cultivars (such as ‘ Golden Delicious ’ and ‘ Fuji ’ ) were initially assessed at harvest as well as after two months on both whole and cut fruit ( Fig. 4). This analysis showed that cut apples stored for two months produced the highest amount of VOCs, suggesting this as the most valuable type of sample to analyze. It is in fact interesting to note that fruit cutting highly stimulates changes in both the concentration and composition of volatile flavor compounds in the headspace. After cutting, or even more evidently after chewing, the food surface area exposed to air increases (de Roos 2003; Arvisenet et al. 2008), and several chemical lipoxygenase, lipase, and fatty acid hydroperoxide lyase; Contreras and Beaudry 2013). Taking this into consideration, apples from 124 cultivars were assessed after 2 months of cold storage and after cutting, to simulate the VOCs release occurring during human consumption. Apple cultivars (Table 2) were organized into hierarchical clusters based on VOCs assessed by PTR-ToF- MS (Fig. 5), which reveals a broad division of the cultivar dataset in two groups discriminated by the overall abun- dance of VOCs, mostly ester compounds. The results are in accordance with earlier findings reporting the impact of esters, alcohols, aldehydes and other carbonyl compounds in the determination of the apple aroma (Table 3; Newcomb et al. 2006; Schaffer et al. 2007; Ulrich and Dunemann 2012; Soukoulis et al. ...
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Citations
... In the context of using molecular markers in VOCs research, a full-sib parental mapping population was used to conduct a QTL survey to evaluate the VOC segregation found in apple fruit collected after a two-month postharvest storage period using a novel proton transfer reaction time of flight mass spectrometry (PTR-ToF-MS) [203]. A group of QTLs distributed over ten chromosomes in the population, such as linkage groups LG2, 3,4,5,11,13,14,15, and 16, were identified through the combined analysis of markers and phenotypic data. ...
... The QTLs found in these areas had LOD values ranging from 3.52 to 14.2, and their respective expressed variance percentages ranged from 26% to 68.9%. The majority of the detected QTLs were linked to sesquiterpenes, alcohols, and esters, according to the annotation of the masses associated with the VOCs profiled by PTR-ToF-MS [203]. Indeed, among all the compounds, esters are the largest class of VOCs in apple aroma and are produced either through the isoleucine pathway or the fatty acid pathway [204]. ...
... The allele 201 and major VOCs were found to be significantly correlated by the candidate gene association analysis. In contrast to acetate esters, which showed a trend in the opposite direction, the presence of allele 201 generally determined a significant increase in the production of volatiles [203]. However, Larsen et al. [205] discovered convincing associations for acetate esters, particularly butyl acetate and hexyl acetate, on chromosome 2 in the region, including the AAT1 gene, using GWA studies. ...
Breeding fruit species is time-consuming and expensive. With few exceptions, trees are likely the worst species to work with in terms of genetics and breeding. Most are characterized by large trees, long juvenile periods, and intensive agricultural practice, and environmental variability plays an important role in the heritability evaluations of every single important trait. Although vegetative propagation allows for the production of a significant number of clonal replicates for the evaluation of environmental effects and genotype × environment interactions, the spaces required for plant cultivation and the intensity of work necessary for phenotypic surveys slow down the work of researchers. Fruit breeders are very often interested in fruit traits: size, weight, sugar and acid content, ripening time, fruit storability, and post-harvest practices, among other traits relevant to each individual species. The translation of trait loci and whole-genome sequences into diagnostic genetic markers that are effective and affordable for use by breeders, who must choose genetically superior parents and subsequently choose genetically superior individuals among their progeny, is one of the most difficult tasks still facing tree fruit geneticists. The availability of updated sequencing techniques and powerful software tools offered the opportunity to mine tens of fruit genomes to find out sequence variants potentially useful as molecular markers. This review is devoted to analysing what has been the role of molecular markers in assisting breeders in selection processes, with an emphasis on the fruit traits of the most important fruit crops for which examples of trustworthy molecular markers have been developed, such as the MDo.chr9.4 marker for red skin colour in apples, the CCD4-based marker CPRFC1, and LG3_13.146 marker for flesh colour in peaches, papayas, and cherries, respectively.
... Fruit quality is affected by appearance, texture, juiciness and nutritional attributes (Abbott, 1999;Cappellin et al., 2015;Corollaro et al., 2014b;Endrizzi et al., 2015). Juiciness and texture have dominant roles in the determination of fruit quality in apple, since these are the two most appreciated characteristics by consumers (Daillant-Spinnler et al., 1996;King et al., 2000;Bourne, 2002).The relative importance of each quality trait component varies greatly among species. ...
Apple fruit quality is strongly influenced by the interplay between juiciness and texture. To better decipher the complexity underneath the control of such quality traits, a multidisciplinary approach combining the mechanic and acoustic profiling of texture, juice analysis, cell morphology, sensory and genetic analysis was carried out. The analyses were conducted after 1.5 months of cold storage on fourteen accessions employed in novel breeding schemes for texture and juiciness. The food matrix structure was exploited focusing on both the cell morphology (employing an optical microscope) and the intercellular space (using an X-ray computed micro-tomography scanner). The mechanical and acoustic properties of texture were profiled with a texture analyser, while the juice was extracted using a mechanical press. In parallel to the analytical assessments, fruit texture, juiciness and flavour were also evaluated by sensory analysis. The results highlighted a positive correlation between cell shape and the intercellular volume. Apple accessions distinguished by round cells were characterized by a reduced intercellular space, while cell with an angular cell shape had a higher intercellular space. While the cell shape was associated with juiciness, the firmness response was more influenced by cell size. The interplay between cellular morphology and juiciness was also investigated together with the allelotype variability of a genetic marker designed for MdPG1, a polygalacturonase gene known to control the regulation of fruit texture in apple. The highest juiciness was found in apples with both a high fraction of round cells and the presence of the MdPG1 allele associated with low softening rates. The elucidation of the role of cellular morphology in the control of fruit texture and juiciness, and their association with the MdPG1 alleles, provided valuable information for a more detailed and informative analysis of fruit quality, enabling a more precise characterization and selection of superior apple accessions.
... For these traits, robust marker-trait associations are needed in order to select the trait by selecting the linked marker. QTL can be identified through genetic mapping based on hybrid populations (Gardner et al., 2014;Harrison et al., 2016;Hernández et al., 2017) or through genome-wide association studies (GWAS), based on unrelated genotype populations (Cappellin et al., 2015;Di Guardo et al., 2017;Urrestarazu et al., 2017). ...
This book covers the biotechnology of all the major perennial fruit and nut species, including Actinidia spp., Anacardium occidentale , Mangifera indica , Pistacia vera , Annona spp., Cocos nucifera , Elaeis guineensis , Phoenix dactylifera , Ananas comosus , Carica papaya , Garcinia mangostana , Diospyros kaki , Vaccinium spp., Castanea spp., Carya illinoinensis , Juglans regia , Persea americana , Theobroma cacao , Musa spp., Psidium guajava , Olea europaea , Averrhoa carambola , Passiflora spp., Eriobotrya japonica , Fragaria × ananassa , Malus × domestica , Prunus persica , Prunus armeniaca , Prunus domestica , Prunus spp., Prunus dulcis , Pyrus spp., Cydonia spp., Rubus spp., Citrus spp., Dimocarpus longan , Litchi chinensis , and Vitis spp. This book also covers biotechnologies and also traditional ones, such as regeneration pathways, protoplast culture, in vitro mutagenesis, and ploidy manipulation that have been applied to many of these species. The species are organized by plant family to facilitate comparisons among related ones. Each species is discussed in relation to its family and its related wild forms, and most are accompanied by full colour illustrations. This book is a vital resource for those working on the improvement of perennial fruit, nut and plantation crops.
... To develop useful selection markers for apple, identification of causal genes for qualitative traits (e.g., ethylene production, skin color, apple scab resistance, columnar tree, and pale green lethal disorder in seedling) have been carried out mainly using linkage analyses and mapbased cloning (Ban et al. 2007, Harada et al. 2000, Orcheski et al. 2015, Otto et al. 2014, Takos et al. 2006, Vinatzer et al. 2001, Wolters et al. 2013. For the many quantitative traits (e.g., soluble sugar content, acidity, phenolic compounds, volatile organic compounds, firmness, bitter pit, and water core), which are commercially important but more difficult to analyze, QTLs have been detected using QTL mapping, genome-wide association study, and pedigreebased analyses (Bink et al. 2014, Cappellin et al. 2015, Chagné et al. 2012, Costa et al. 2010, Kenis et al. 2008, King et al. 2000, Kumar et al. 2013, Kunihisa et al. 2014, Liebhard et al. 2003. However, these studies were carried out using heterozygous materials, which might have lowered the QTL detection power. ...
Apple is an economically important crop, and various approaches to genetic analysis in breeding programs have been attempted, including the production of doubled haploid (DH) lines, which are genetically homozygous. In this study, we used a DH line for QTL analyses, for the first time in a fruit tree, expecting it to simplify the analysis of the inheritance of quantitative traits and thus to enhance QTL detection power. Using an F1 population from ‘Prima’ × ‘Apple Chukanbohon 95P6’ (DH), we constructed a genetic map of ‘Prima’, and identified 19 QTLs for 13 traits. These QTLs had comparatively high LOD scores and explained a large part of the variation of the phenotypes. In particular, acidity, juice browning, and skin splitting clearly segregated at a 1:1 ratio, consistent with the segregation of the alleles at the detected QTLs in linkage group 16; these traits appeared to be regulated by single genes, despite general consideration that they are quantitative traits. Using this simple genetic composition of the F1 population, we concluded that the skin splitting of apple fruit has recessive inheritance, and that the allele for splitting is tightly linked with those for high acidity and low juice browning in ‘Prima’.
... Although this represents a valuable and accurate technique, it is laborious and time consuming. Therefore, Proton Transfer Reaction-Time Of Flight-Mass Spectrometry (PTR-ToF-MS) might represent a valid alternative to profile VOCs in a more time-efficient way (Lindinger et al., 1998; Jordan et al., 2009) also in apple (Zini et al., 2005; Costa et al., 2013; Cappellin et al., 2015). Beside the headspace concentration of VOCs, the interaction between aromatic compounds and human receptors should also be considered (Farneti et al., 2015b). ...
Fruit quality represents a fundamental factor guiding consumers’ preferences. Among apple quality traits, volatile organic compounds and texture features play a major role. Proton Transfer Reaction-Time of Flight-Mass Spectrometry (PTR-ToF-MS), coupled with an artificial chewing device, was used to profile the entire apple volatilome of 162 apple accessions, while the fruit texture was dissected with a TAXT-AED texture analyzer. The array of volatile compounds was classed into seven major groups and used in a genome-wide association analysis carried out with 9142 single nucleotide polymorphisms (SNPs). Marker–trait associations were identified on seven chromosomes co-locating with important candidate genes for aroma, such as MdAAT1 and MdIGS. The integration of volatilome and fruit texture data conducted with a multiple factor analysis unraveled contrasting behavior, underlying opposite regulation of the two fruit quality aspects. The association analysis using the first two principal components identified two QTLs located on chromosomes 10 and 2, respectively. The distinction of the apple accessions on the basis of the allelic configuration of two functional markers, MdPG1 and MdACO1, shed light on the type of interplay existing between fruit texture and the production of volatile organic compounds.
... The protonated m/z = 47.049, TI as ethanol Cappellin et al. 2014;Taiti et al. 2015a), and the protonated m/ z = 61.028, TI as acetic acid/acetates Taiti et al. 2015a), are both generally considered in olive oils as compounds deriving from alteration due to a long time of olive storage before processing (Morales et al. 2000;Angerosa et al. 1996a) the protonated m/z = 75.044, ...
Olive oil samples were obtained from six culti-
vars grown in different environments, and graded by
chemical analyses as extra virgin (EVOOs). These were
evaluated for flavors and off-flavors, and relative VOCs
spectrum as dete rmined by PTR–ToF–MS. A hierarchical
clustering of Panel test data separated olive oil in three
groups, one including the samples with perceived off-flavor
(VOOs), regardless of cultivar and environment. The
Pearson’s correlation coefficients between the mass data
from PTR–ToF–MS and the sensory characteristics per-
ceived by the Panel test were determined. A mass-to-sen-
sory attributes correlation index was calculated. A color-
coded card was built up based on the intensities (ncps) of
five selected prot onated mass data that was able to distin-
guish EVOOs from VOOs olive oil samples.
... Each graph is divided by a line, at time 0, in two phases, respectively, before and after the "chewing moment": (1) headspace analysis of the intact apple flesh cylinder and (2) headspace analysis of "chewed" apple flesh (between 0 and 120 s). Adapted from Farneti, B., Khomenko, I., Cappellin, L., Ting, V.J.L., Costa, G., Biasioli, F., Costa, F., 2015. Dynamic volatile organic compound fingerprinting of apple fruit during processing. ...
The instrumental characterization of volatile organic compounds (VOCs) is essential to have a precise, reliable, and reproducible estimation of food aroma and, therefore, of the overall product quality. In this article we will list the main characteristics of PTR-MS (proton transfer reaction mass spectrometry) and its application in the agri-food chain research. The high time resolution brought by this technology allows real-time monitoring of fast food processes and rapid and noninvasive VOC fingerprinting.
... Our objective here was to detect QTLs responsible for the superior characteristics of 'Fuji', by using sequence information of 'Fuji' and many of its relatives. QTLs for several traits have been previously reported on the genetic map of 'Fuji', or the integrated maps of the populations generated by crossing 'Fuji' with other cultivars; i.e., a QTL for firmness (Costa et al. 2010), QTLs for some other physical texture components (Longhi et al. 2012), and QTLs for some volatile organic compounds (Cappellin et al. 2015). Here, we focused on the degree of watercore and flesh mealiness as an index of fruit shelf-life. ...
‘Fuji’ is one of the most popular and highly-produced apple cultivars worldwide, and has been frequently used in breeding programs. The development of genotypic markers for the preferable phenotypes of ‘Fuji’ is required. Here, we aimed to define the haplotypes of ‘Fuji’ and find associations between haplotypes and phenotypes of five traits (harvest day, fruit weight, acidity, degree of watercore, and flesh mealiness) by using 115 accessions related to ‘Fuji’. Through the re-sequencing of ‘Fuji’ genome, total of 2,820,759 variants, including single nucleotide polymorphisms (SNPs) and insertions or deletions (indels) were detected between ‘Fuji’ and ‘Golden Delicious’ reference genome. We selected mapping-validated 1,014 SNPs, most of which were heterozygous in ‘Fuji’ and capable of distinguishing alleles inherited from the parents of ‘Fuji’ (i.e., ‘Ralls Janet’ and ‘Delicious’). We used these SNPs to define the haplotypes of ‘Fuji’ and trace their inheritance in relatives, which were shown to have an average of 27% of ‘Fuji’ genome. Analysis of variance (ANOVA) based on ‘Fuji’ haplotypes identified one quantitative trait loci (QTL) each for harvest time, acidity, degree of watercore, and mealiness. A haplotype from ‘Delicious’ chr14 was considered to dominantly cause watercore, and one from ‘Ralls Janet’ chr1 was related to low-mealiness.
... In apple (Malus x domestica Borkh.) the aroma profile results from the interaction of more than 270 VOCs (Farneti et al., 2014; Nijssen, van Ingen-Visscher, & Donders, 2011; ) synthesized by intact fruit prior to consumption, as well as in response to cellular disruption caused by biting and mastication (Contreras & Beaudry, 2013). Among them, esters, alcohols and aldehydes are recognized as the most relevant volatiles responsible for aroma in apple (Cappellin et al. 2014; Dunemann et al. 2012; Holland et al., 2005;). The volatile emission behavior greatly depends by the fruit integrity. ...
... Among the various possibilities proposed and investigated for a rapid quantification and identification of VOCs by direct injection, PTR-ToF (Time of Flight)-MS is one of the most used, since it allows on-line measurement of a mixture of VOCs in a straightforward, fast and high sensitivity fashion (Biasioli, Gasperi, Yeretzian, & M€ ark, 2011 ). Furthermore PTR-ToF-MS has already been exploited in apple , validating the capacity of this technique to characterize the aroma profile in apple fruit (Cappellin et al. 2014; Farneti et al. 2014). The aim of this work was the development of a fast and reliable system to study the volatile aroma profile of apple fruit, imitating the release of volatiles in the human mouth during fruit consumption without taking into account the several parameters varying between different consumers, such as number of bites, chewing speed, and liquid lubricant. ...
Genetic diversity is the primary source of variability in any crop improvement program, and the diverse germplasm of any crop species represents an important genetic resource for gene or allele mining to meet future needs. Huge genetic and phenotypic diversity is present in the apple gene pool, even though, breeding programs have been mainly focused on a few traits of interests, which have resulted in the reduction of the diversity in the cultivated lines of apple. Therefore, the present study was carried out on 70 diverse apple genotypes with the objective of analyzing the genetic diversity and to identify the genetic loci associated with important fruit quality traits. A total of 140 SSR primers were used to characterize the 70 genotypes of apples, out of which only 88 SSRs were found to be polymorphic. The PIC values varied from 0.03 to 0.75. The value of MI, EMR, and RP varied from 0.03 to 3.5, 0.5 to 5.0, and 1.89 to 6.74, respectively. The dendrogram and structure analysis divided all the genotypes into two main groups. In addition to this, large phenotypic variability was observed for the fruit quality traits under study indicated the suitability of the genotypes for association studies. Altogether 71 novel MTAs were identified for 10 fruit quality traits, of which 15 for fruit length, 15 for fruit diameter, 12 for fruit weight, 2 for total sugar, 2 for TSS, 4 for reducing sugar, 5 for non-reducing sugar, 5 for fruit firmness, 5 for fruit acidity and 6 for anthocyanin, respectively. Consistent with the physicochemical evaluation of traits, there was a significant correlation coefficient among different fruit quality characters, and many common markers were found to be associated with these traits (fruit diameter, length, TSS, total sugar, acidity and anthocyanin, respectively) by using the different modeling techniques (GLM, MLM). The inferred genetic structure, diversity pattern and the identified MTAs will be serving as resourceful grounds for better predictions and understanding of apple genome towards efficient conservation and utilization of apple germplasm for facilitating genetic improvement of fruit quality traits. Furthermore, these findings also suggested that association mapping could be a viable alternative to the conventional QTL mapping approach in apple.
